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
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FIELDIANA: BOTANY
2a. Calyx tube 7-12 mm long with lobes to 3 mm long; inflorescences with 3-15 distantly spaced
flowers, ovary and fruit pilose . c. enneantha
2b. Calyx tube 0.5-2 mm long with lobes to 1 mm long; inflorescences with 15-50 proximate
flowers, ovary and fruit glabrous .3
3a. Leaf blades 11-28 x 5-15 cm; corolla tube 10-13 mm long, distinctly pubescent; in the
Caribbean lowlands, 0-300 mm elevation c. hondensis
3b. Leaf blades 6-19 x 3-8 cm; corolla tube ca. 9 mm long, glabrous or minutely puberulent;
cloud forests, (600-) 1 200-2000 m elevation C. austin-smithii
4a. Leaves subsessile, domatia of tufted hairs or pits often present along the midvein 5
4b. Leaves with petioles usually more than 4 mm long, domatia absent along veins beneath (rarely
present in C. chiriquiensis 7
5a. Corolla usually minutely sericeous, calyx tube 2-3 mm long; domatia usually narrow de- pressions along the midvein above the vein axils [leaf blades 7-18 x 3-9 Cm; Caribbean
lowlands of northern Costa Rica] C impetiolaris
5b. Corolla glabrous or minutely papillate puberulent; calyx tube 1-2 mm long; domatia of shallow
puberulent depressions in the vein axils or absent 6
6a. Inflorescences 3-10 cm long, paniculate with distinct lateral branches; leaf blades 7-16 cm long, domatia of puberulent pits in the vein axils; Caribbean lowlands of southernmost Costa
Rica C. sp. A aff. curvigemmia
6b. Inflorescences ca. 3 cm long, subcapitate (paniculate but with short, closely spaced, lateral branches); leaf blades more than 1 5 cm long, domatia present or absent; lowland rain forest
C. sp. B aff. curvigemmia
7a. Flowering portion of the inflorescences elongate-racemiform, distinctly longer than broad ..... 8 7b. Flowering portion of the inflorescences open paniculate, corymbiform to umbelliform or pyramidal,
usually with length equaling breadth 10
8a. Stipules narrowly oblong to linear, to 2 cm long and 4 mm thick, acute at the apex and Ficus- like, enclosing apices of stems or lateral branches in early stages; leaves usually less than 7
cm broad [corolla 1 5 mm long] C. caroliana
8b. Stipules not linear or narrowly oblong, not resembling the stipules of Ficus, flattened and
enclosing the shoot apex but quickly caducous; leaves usually more than 8 cm broad ... 9
9a. Calyx tube (limb) ca. 2 mm long; fruit 9-17 mm diam., ellipsoid to obovoid; leaf blades
chartaceous, short- to long-acuminate, petioles to 15 mm long; stipules usually bluntly obtuse
at apex C. talamancana
9b. Calyx tube 5-7 mm long; fruits 10-28 mm diam., ellipsoid-oblong; leaf blades subcoriaceous, short-acuminate or rounded at apex, petioles to 40 mm long; stipules usually broadly rounded
distally C. latifolia
lOa. Plants of montane cloud forest formations (600-) 1 200-2000 m elevation; stipules often persisting
11
1 Ob. Plants of lowland evergreen formations, 0-600 m elevation; stipules usually caducous 13
1 la. Corolla tube ca. 6 mm long [leaf blades 3-5 m broad, 5-7 major 2° veins]
C. chiriquiensis
1 Ib. Corolla tube 9-18 mm long 12
12a. Leaf blades 2-4 cm broad and 4-6 major 2° veins; Chiriqui Highlands, Panama
C. nebulosa
12b. Leaves usually 3-8 cm broad and with 6-9 major 2° veins; Costa Rica . C. austin-smithii 13a. Leaves usually drying very dark or black, membranaceous to thin-chartaceous; flowers usually
4-parted; stipules often with 2 minute (0.3 mm) lobes on each side C. nigrescens
1 3b. Leaves usually drying greenish or brownish, usually stiffly chartaceous; flowers 5-parted; stipules
various 14
14a. First node of the inflorescence usually with 4 lateral branches; fruits with 2 seeds, dorsal surface
of seed with longitudinal ridges; corolla lobes 3-7 mm long; calyx lobes 0.3-1 mm long
Psychotria eurycarpa
14b. First node of the inflorescence usually with 2 lateral branches; fruits usually single seeded (rarely 2-seeded), seed smooth on the dorsal surface; corolla lobes 6-20 mm long; stipules without lobes; calyx lobes present or absent C. psychotrioides
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 1 1 1
Coussareaaustin-smithiiStandl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1286. 1938. Psychotria tutensis Dwyer, Ann. Missouri Bot. Gard. 67: 434. 1980. Figure 47.
Coussarea caroliana Standl., Field Mus. Nat. Hist., Bot. Ser. 22: 178. 1940. C. veraguensis Dwyer, Ann. Missouri Bot. Gard. 67: 134. 1980. Figure 47.
Shrubs or small trees 3-6(-10) m tall, leafy stems 1.5- 6 mm thick, glabrous or very minutely (0.05 mm) pa- pillate in early stages, with thickened nodes and longi- tudinally striate (dried); stipules 4-8 mm long, 4-7 mm broad, triangular with an acuminate or narrowed apex, rarely with a small (0.5 mm) U-shaped sinus at the tip, glabrous, persisting or deciduous. Leaves opposite, pet- ioles 8-30 mm long, 0.6-2 mm thick, glabrous or mi- nutely puberulent; leaf blades (5-)7-19 cm long, (2-)3- 8.5 cm broad, elliptic to narrowly elliptic-oblong, ellip- tic-obovate or narrowly ovate, apex acute to short-acu- minate, base acute to obtuse and often slightly decurrent on petiole, drying chartaceous, dark green or dark brown above, glabrous above, glabrous or minutely (0.1-0.2 mm) puberulent on the veins beneath, 2° veins 6-9/side, without domatia. Inflorescences 5-10 cm long, equally broad, open paniculate or umbelliform (3-branched, rarely with 4 branches from the first node), peduncles 12-20 mm long, 1-2 mm thick, glabrous or very mi- nutely papillate-puberulent, primary branches 1-3 cm long and opposite or alternate, flowers cymose or in distal groups of (1) 2 or 3, bracts absent or minute (sometimes borne 3-7 mm up along the lateral branches), pedicels 0-8 mm long, purplish. Flowers fragrant, hypanthium 1-1.5 mm long, ca. 1.5 mm diam., obconic-tubular, gla- brous, calyx tube 0.5-1 mm long and ca. 3 mm broad, broadly cupulate, calyx lobes to 0.5 mm high (and broad- ly triangular) or not developed and the calyx entire; co- rolla salverform, white, glabrous or minutely papillate- puberulent externally, tube 9-10 mm long (to 1 5 mm in life?), 1.2-2.2 mm diam., lobes 4(-5), ca. 7 mm long, 2 mm broad and fleshy, narrowly oblong; style branches 1.8 mm long. Fruits 10-12 mm diam., globose, greenish white with pale longitudinal lines and becoming red, purple, or black, persisting calyx less than 0.5 mm high.
Plants of evergreen montane cloud forest for- mations of the Caribbean slopes and continental divide, from (600-)1200 to 2000 m elevation. Flowering in June-November (peaking in August); fruiting in July and November-February. This species is known only from the Cordilleras de Guanacaste and Tilaran, the northern slopes of the Meseta Central (San Ramon-Zarcero), and above the Rio Reventazon, in northern and central Costa Rica.
Coussarea austin-smithii is recognized by its usually open-corymbiform or umbelliform inflo- rescences without bracts, flowers with short broad- ly cupulate calyx with poorly developed lobes, long narrow corolla lobes, and its cloud forest habitat. This is our only Coussarea species growing above 1 800 m elevation.
Shrubs or small treelets, 1 .5-4(-6) m tall, leafy branch- lets 1 .2-5 mm thick, glabrous; stipules 1 2-20 mm long, 2-4 mm diam., united and forming a slender cap over the shoot apex, glabrous, acute at apex, usually splitting down one side and caducous. Leaves opposite, petioles 4-14(-18) mm long, 1-3 mm thick, glabrous, slightly sulcate above; leaf blades 8-14(-17) cm long, 3-7(-9.5) cm broad, elliptic-oblong, narrowly elliptic-oblong, el- liptic to ovate-elliptic or elliptic-obovate, apex short- acuminate, base acute, drying chartaceous and usually grayish green, glabrous above and small linear cystoliths often visible, glabrous beneath, 2° veins 7-10/side and weakly loop-connected near the margin. Inflorescences 3-10 cm long, 3-5 cm broad, racemiform or spiciform, peduncles 1-3 cm long, glabrous, the proximal 2 lateral branches opposite or subopposite, to 6 mm long, distal flowers or flower clusters sessile or subsessile, bracts ab- sent or minute (0.3 mm), pedicels 0-3 mm long. Flowers glabrous, hypanthium 1-2 mm long (not clearly differ- entiated from the pedicel or calyx tube), calyx tube 1- 1.5 mm long, 1.8-2.5 mm broad at apex, entire; corolla white, tube 6-10 mm long, 0.7-2 mm diam., lobes 4, 4- 8 mm long. Fruits 12-15 mm long, 7-10 mm diam., ellipsoid-oblong to oblong-obovoid, drying yellowish and minutely white-lenticellate, the persisting calyx 0.5-1 mm high.
Plants of evergreen rain forest formations on the Caribbean slope and central Cordilleras, from 350 to 1500(-1800) m elevation. Flowering in Janu- ary-April; probably fruiting throughout the year. The species ranges from the Cordillera de Guana- caste eastward to the western part of the Cordillera de Talamanca and western and central Panama.
Coussarea caroliana is recognized by its lower montane habitat, lack of puberulence, racemiform inflorescence without bracts, entire calyx tube, and distinctive fruit. The elongate Ficus-like stipules forming a cap over the shoot apices are distinctive; they are often seen at the base of an inflorescence enclosing the apices of new lateral shoots. None of our other species of Coussarea have such stip- ules. The dried leaves are often a characteristic grayish green beneath. Specimens with smaller leaves and immature fruits may resemble Rudgea cornifolia.
Several specimens with larger (18-26 x 6.5-12 cm) leaf blades and larger (30 x 13 mm) fruits on long (14 cm) infructescences are tentatively placed here. All are from the Caribbean lowlands: Gray- urn et al. 8754 CR, Gomez- Laurito 8785 (sterile) CR, and Opler 340 CR, F. They may represent an
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FIELDIANA: BOTANY
unrecognized species or merely an extreme form of C. caroliana.
Coussarea chiriquiensis (Dwyer) C. M. Taylor, comb. nov. Rudgea chiriquiensis Dwyer, Ann. Missouri Bot. Gard. 67: 476. 1980.
Shrubs ca. 4 m tall, leafy stems 0.7-4 mm thick, gla- brous; stipules 2-4 mm long, broadly triangular or rounded, entire distally or with 2-4 short stiff lobes, usually with thickened teeth within, the base persisting. Leaves with petioles 5-16 mm long, 0.5-1.3 mm thick, glabrous; leaf blades (5-)6-12 cm long, (2-)3-5 cm broad, ovate-elliptic to elliptic or lanceolate, apex acute to acu- minate, base obtuse to acute, drying chartaceous, dark above, glabrous, 2° veins 5-7/side. Inflorescences soli- tary and terminal, (3-)5-7 cm long, (2-)4-9 cm broad, peduncles 9-33 mm long, lateral branches of the first node opposite, often longer than the peduncle and equal- ing the rachis, glabrous, flowers sessile. Flowers glabrous, hypanthium ca. 1 mm long and 0.9 mm diam., tubular, calyx 0.5-1 mm long, broadly cupulate, calyx lobes mi- nute or broadly triangular; corolla 6 mm long and 1.5 mm diam. in bud. Fruits not seen.
The original description of Coussarea chiri- quiensis was based on a single collection (Croat 37071 MO) from about 1500 m elevation, above San Felix, Chiriqui, Panama. The colleters or teeth within the stipule may have been mistaken for the distal stipular teeth that distinguish Rudgea, We tentatively place Burger et al. 10702 (CR, F, MO) here, which is similar but has pit-domatia and an entire stipule sheath. It was collected on the Pacific slope beneath Monteverde, Puntarenas, at ca. 1 400 m elevation. All this material is very similar in overall appearance to Coussarea nebulosa and Faramea ovalis.
Coussarea enneantha Standl., J. Wash. Acad. Sci. 18: 282. 1928. Figure 46.
Shrubs or small trees, 1.5-7 m tall, leafy stems 2-4 mm thick, glabrous or with thin pale brownish hairs 0.3- 0.7 mm long; stipules united and forming a short (2-3 mm) sheath with rounded to truncated distal margin. Leaves opposite, petioles 14-36 mm long, 0.6-1.4 mm thick, sparsely to densely pilose, leaf blades 9-17 cm long, 3-8 cm wide, elliptic to elliptic-oblong, apex acute to long-acuminate with tip 7-14 mm long, base obtuse or acute, drying thin- to stiffly chartaceous and concol- orous, glabrescent above, pubescent on the veins be- neath, 2° veins 8-10/side and weakly loop-connected near the margin. Inflorescences 2.5-8 cm long, to 8 cm broad, peduncles to 6 cm long, ca. 0.7 mm thick and with thin erect hairs 0.3-0.7 mm long, with 3-5 distant
flowers, peduncle often bearing 3 sessile or long-pedi- cellate flowers (or with 2 lateral flowers and a slender rachis bearing 3 remote flowers), bracts absent, pedicels to 2 cm long, pilose. Flowers with hypanthium 1-3 mm long and densely hirsute with erect or ascending yellow- ish brown hairs 0.3-0.7 mm long, calyx tube 4-8 mm long and 1.5-2.5 mm diam., sparsely pubescent, calyx lobes 4, (3-)4-8 mm long, ca. 2 mm broad, narrowly oblong; corolla salverform, white, tube 20-30 mm long and 3-4 mm diam., with ascending hairs ca. 0.7 mm long, corolla lobes 4, 12-15 mm long and 2 mm wide, glabrous. Fruit to 3 cm long and 1 5 mm diam., ellipsoid, densely hirsute, the persisting calyx 10-17 mm long.
Plants of evergreen rain forest formations, from near sea level to 1000 m elevation. This species is only known from Panama, but a collection from the Fish Creek Mountains in Bocas del Toro Prov- ince suggests that this species may be found in the Talamanca Valley of Costa Rica. The pubescent hypanthium, elongate calyx tube, and few-flow- ered inflorescences with long slender pedicels are very distinctive, but the flowers appear to vary greatly in size. The open few-flowered inflores- cences resemble those of Faramea occidentalis and F. paucijlora.
Coussarea hondensis (Standl.) C. M. Taylor & W. Burger, Selbyana 12: 138. 1 99 1 . Psychotria hon- densisStandLJ. Wash. Acad. Sci. 18: 183. 1928. P. ostaurea Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 143. 1967. Figure 48.
Shrubs or small trees, 2-10(-15) m tall, trunks to 20 cm dbh, leafy stems 3-9 mm thick, with fine soft hairs 0. 1-0.4 mm long or minutely papillate-puberulent; stip- ules 7-15 mm long and 4-8 mm broad at the base, triangular to narrowly oblong, puberulent, acute or bi- dentate with teeth to 2 mm long, often persisting. Leaves with petioles 10-35(-70) mm long, 1-3 mm thick, pu- bescent; leaf blades 11-29 cm long, 5-16 cm broad, broadly elliptic to broadly elliptic-obovate or ovate-ob- long, apex acute or short acuminuate apex, tip to 10(-14) mm long, base acute to broadly obtuse, drying stiffly chartaceous and brownish or greenish, glabrous above, with short (ca. 0.2 mm) soft hairs on the veins and sur- faces beneath, 2° veins 7-10/side. Inflorescences 6-15 cm long, to 10(-15) cm broad, paniculate or corymbi- form with opposite, alternate or clustered lateral branch- es, peduncles 4-10 cm long, 1 .5-2.5 mm thick, minutely puberulent, bracts 0.5-1 mm long or caducous, flowers in distal groups of 1-3, pedicels 0-3 mm long. Flowers distylousand nocturnal, hypanthium 1-2 mm long, 1.3- 1.7 mm diam., turbinate, minutely puberulent, calyx tube 0.7-2 mm long, cupulate or spreading and 3 mm broad, lobes minute or absent; corolla salverform, white, minutely puberulent externally, tube 10-18 mm long, 0.8-3 mm diam., glabrous within, lobes 4-5, (4-)7-12
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
113
mm long, 1.5-2 mm broad, narrowly oblong, acute; an- thers ca. 5 mm long. Fruits 12-24(-20?) mm long, ca. 10(-15?) mm diam., ellipsoid-oblong, becoming red- purple, persisting calyx less than 0.5 mm high.
Plants of evergreen rain forest formations in the Caribbean lowlands, from near sea level to 500 m elevation (to 900 m in Panama). Flowering in May- September; fruiting in June-August and October- January. The species ranges from Tortuguero in northern Costa Rica and the Osa Peninsula, south- ward to Code Province in Panama.
Coussarea hondensis is recognized by the mi- nute soft puberulence on vegetative and flowering parts, the large long-petiolate and broadly elliptic leaves, the few- or many-branched and umbelli- form inflorescences, broad and usually entire ca- lyx, and the oblong fruit. An atypical collection (Burger & Malta 4729 CR, F) with almost glabrous leaves from above Golfito is tentatively placed here.
Coussarea impetiolaris J. D. Smith, Bot. Gaz. 37: 418. 1904. Figure 46.
Coussarea impetiolaris is recognized by the sub- sessile leaves with slightly auriculate leaf bases, the few-branched glabrous inflorescences, puber- ulent corolla tubes, and the unusual fruit drying pale in color and with round or oblong wart-like lenticels. The longitudinally elongate domatia with hairs along the sides or hairs along the midvein are a distinctive feature when present. This species was misinterpreted in the past to include material from Guatemala (now segregated as C. imitans L. O. Williams with more puberulent hypanthium- calyx and dark green fruit). Coussarea curvigem- mia Dwyer of central Panama with smaller flowers is also closely related; see the discussions under Coussarea spp. A & B aff. C. curvigemmia. Ma- terial from the Osa Peninsula shows considerable variation and may indicate that the material placed under Coussaria sp. B is only an extreme form of C. impetiolaris; see the discussion under Coussa- rea sp. B. This species may be mistaken for a Rudgea.
Coussarea jiminezii J. D. Smith is a species of Viburnum (Caprifoliaceae).
Small trees to 1 7 m tall, leafy branchlets 1-3 mm thick, glabrous and drying greenish; stipules 2-4 mm long, tri- angular, glabrous and coriaceous, deciduous. Leaves sub- sessile, petioles 1-4 mm long, 0.7-1.8 mm thick, gla- brous; leaf blades 7-1 8 cm long, 3-9 cm broad, narrowly to broadly elliptic, elliptic-oblong or oblanceolate, apex acuminate with tip 5-15 mm long, gradually narrowed to an acute base and slightly auriculate on the petiole, leaves drying chartaceous and greenish or grayish, gla- brous above and below, but with hairs along the edges of pit dormatia in or near the vein axils beneath, 2° veins 4-7 /side. Inflorescences 3-5 cm long, to 7 cm broad, paniculate and often with 1 or 2(-3) pairs of opposite branches and 9-1 5 or more flowers, peduncles 10-26(-40) mm long, ca. 1.2 mm thick, glabrous, lateral branches to 1 2(-l 5) mm long, flowers sessile, bracteoles 0.3-1 mm long. Flowers with hypanthium ca. 2 mm long, calyx tube little differentiated from the hypanthium, 2-3 mm long, 2-3 mm diam. distally, glabrous or minutely pa- pillate-puberulent, calyx lobes 0.2-0.3 mm high; corolla white, minutely sericeous externally, tube (8-)l 1-16(-20) mm long, lobes 4, 6-9 mm long, 2 mm broad at the base; anthers ca. 8 mm long. Fruits 1 5-20 mm long and 14-15 mm diam., broadly ellipsoid and slightly flattened laterally, surface smooth and pale yellowish white with distinctive white (becoming brown) tuberculate lenticels 0.5-1.4 mm long, persisting calyx ca. 2 mm high.
Plants of the lowland Caribbean rain forest for- mations, from 15 to 500 m elevation. Flowering in March-July; fruiting in January, March, and June-November. The species is found in the Ca- ribbean lowlands, the Osa Peninsula, and Panama.
Coussarea latifolia Standl., J. Wash. Acad. Sci. 1 8: 281. 1928. Figure 48.
Small trees, 6-1 5 m tall, perhaps dioecious, leafy stems 4-12 mm thick, glabrous, quadrangular; stipules ca. 5 mm long, rounded distally, glabrous, deciduous. Leaves opposite, petioles 9-30 mm long, 2-4 mm thick, terete, glabrous; leaf blades 17-30 cm long, 9-19 cm broad, broadly elliptic to broadly elliptic-obovate or ovate-el- liptic, apex abruptly narrowed to the short-acuminate, tip 4-10 mm long, base obtuse, drying stiffly chartaceous or subcoriaceous, grayish green, glabrous above and be- low, 2° veins 7-9/side, domatia absent. Inflorescences 2-6 cm long, 3-6 cm broad, racemose in form with cy- mose flower clusters on short (6-1 5 mm) lateral branches or with pedicellate flowers from the central rachis, pe- duncles 2-30 mm long, 2-3.5 mm thick and glabrous, bracts minute, pedicels 1-5 mm long. Flowers glabrous, hypanthium 3-4 mm long, obovoid, poorly differenti- ated from the calyx tube, calyx tube 5-7 mm long, 4-5 mm diam., calyx lobes not developed; corolla salver- form, yellowish white, tube ca. 10 mm long (perhaps not fully expanded), lobes 4, 13-15 mm long; anthers 4-5 mm long. Fruits 25-45 mm long (including persisting calyx 2-6 mm long), 10-28 mm diam., ellipsoid-oblong, green with white spots and becoming yellow.
Plants of evergreen rain forest formations of the Caribbean lowlands, from 5 to 300 m elevation (to 1 000 m in the central highlands of Panama). Flowering in April-May; fruiting in July, Septem- ber-October, and December. The species ranges
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FIELDIANA: BOTANY
from central Costa Rica (Reventazon valley) to Colombia.
Coussarea latifolia is recognized by its large broadly elliptic leaves on prominent thick petioles and with relatively few secondary veins, the lack of pubescence, the racemose inflorescences, and long calyx tube. We have seen only the following collections from Costa Rica: Grayum et al. 8754 MO, Shank & Molina 4422 F, and Tonduz 9574 us holotype. This species is difficult to separate from the much more often collected C. talaman- canum in the absence of flowers or mature fruit.
Coussarea nebulosa Dwyer, Ann. Missouri Bot. Card. 67: 131. 1980.
Shrubs, ca. 3 m tall, leafy stems 1-5 mm thick, gla- brous, becoming grayish, terete and smooth; stipules 1- 3 mm long, 2-3 mm broad, with a broad U-shaped sinus and 2 small (0.7 mm) lobes, deciduous. Leaves with petioles 4-16 mm long, 0.5-1 mm thick, glabrous; leaf blades 6-13 cm long, 2-4 cm broad, elliptic-lanceolate to narrowly elliptic-oblong, apex tapering gradually to the acuminate, base acute, drying stiffly chartaceous and dark olive green above, glabrous above and below, 2° veins 4-6/side. Inflorescences solitary and terminal, 2.5- 7 cm long, 1.5-6 cm broad, paniculate with opposite lateral branches, peduncle 6-46 mm long, 0.7-1.3 mm thick, glabrous, bracts ca. 2 mm long, subulate, flowers mostly sessile in distal triads. Flowers glabrous exter- nally, hypanthium 0.8-1.3 mm long, ca. 1 mm diam., calyx tube 0.5-0.8 mm high, lobes 0.2-0.5 mm long; corolla white, tube 10-1 8 mm long, lobes ca. 6 mm long. Fruits unknown.
Coussarea nebulosa is a species of the Chiriqui Highlands known only from near Boquete at about 1200-1600 m elevation. The smaller leaves, un- usual stipules (for the genus), glabrous parts, and small inflorescences are distinctive. There are two other similar small-leaved species in the Chiriqui Highlands: Coussarea chiriquensis and Faramea ovalis.
Coussarea nigrescens C. M. Taylor & Hammel, Selbyana 12: 134. 1991.
Shrubs or small treelets, 2-7 m tall, leafy stems 1.2- 4 mm thick, glabrous or minutely (0.05 mm) papillate- puberulent, drying dark; stipules united to form a short sheath 0.8-2 mm long, at first broadly triangular but becoming truncated or with 2 minute lobes ca. 0.3 mm long, glabrous, the base persisting as a short collar above the node. Leaves with petioles 6-35 mm long, 0.7-1.7 mm thick, glabrous or minutely papillate-puberulent.
drying black; leaf blades 8-2 1 cm long, 3-9 cm broad, elliptic to elliptic-oblong or slightly ovate-elliptic, apex abruptly narrowed and short-acuminate with tip 2-8 mm long, base cuneate to obtuse, drying membranaceous to thin-chartaceous and blackish above, glabrous and lus- trous above, glabrous or minutely puberulent along the midvein beneath, 2° veins 6-10/side, weakly loop-con- nected near the margin in the distal half of the leaf. Inflorescences solitary and terminal, 4-1 1 cm long, 3- 1 1 cm broad, open umbelliform panicles with 2 or 4 branches at the first node, peduncle 15-50 mm long, 0.7-2.2 mm thick, glabrous or papillate-puberulent, dry- ing black, bracts subtending the 1° branches 2-6 mm long, flowers sessile in distal cymes or glomerules of 3- 7, distal bracts ca. 1 mm long. Flowers minutely (0.05 mm) papillate-puberulent externally, hypanthium 0.6-1 mm long, calyx cup only 0.5 mm long, becoming rotate, lobes minute; corolla tubular, white, tube 14-24 mm long and 1.2-2 mm diam., lobes 4 or 5, 6-10 mm long, lanceolate; ovary with well-developed septum and 2 loc- ules. Fruits 9-26 mm long, 6-18 mm diam., purple- black, glabrous; pyrenes 1 or 2, globose to ellipsoid.
Plants of evergreen rain forest formations of the Caribbean lowlands and southern Pacific low- lands, from 20 to 800 m elevation. Flowering in July and October-November; fruiting in Febru- ary. This species has been collected in the de- partment of Zelaya, Nicaragua, near La Selva and Puerto Viejo de Sarapiqui, Cerro Nara east of Que- pos, and Corcovado National Park in Costa Rica. This species is known only from Costa Rica and Nicaragua.
Coussarea nigrescens is recognized by the many parts drying dark or blackish, the very thin leaves, short collar-forming stipules, and slender corolla tubes that are minutely papillate-puberulent. This species may be related to C. nebulosa of the Chi- riqui Highlands with smaller stiffer leaves that do not dry so dark, larger calyx lobes, lack of puber- ulence, and higher-elevation habitat.
Coussarea psychotrioides Taylor & Hammel, Sel- byana 12: 135. 1991. Figure 47.
Shrubs or small trees, 2.5-7(-12) m tall, trunks to 20 cm dbh, leafy stems 1.5-4 mm thick, glabrous, nodes often conspicuously thickened: stipules 0.3-2 mm long, truncate or slightly bilobed, quickly caducous and leav- ing a short cupulate ring around the stem just above the distal nodes, glabrous. Leaves with petioles 6-20 mm long, 0.6-1.7 mm thick, glabrous, sulcate above; leaf blades 8-18 cm long, (2.5-)3-8.5 cm broad, narrowly elliptic to elliptic-oblong or elliptic-ovate, apex short- acuminate (rarely acute or obtuse), tip 5-12 mm long, base obtuse to acute, drying chartaceous and often yel- lowish green or greenish brown above, glabrous above and below, 2° veins 6-9/side and loop-connected near
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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the margin in the distal half of the lamina. Inflorescences 3-10 cm long and 4-10 cm broad, broadly paniculate to umbelliform, peduncles 1-3 cm long, 1-2 mm thick, glabrous, first branching node usually with 2 branches and these often coequal with the continuing rachis (= umbelliform), bracts 0.4-3 mm long or absent, flowers usually subsessile in distal 3-7-flowered cymes or glom- erules, pedicels 0-3.5 mm long. Flowers glabrous, noc- turnal and distylous, hypanthium 0.7-2 mm long, 0.5- 1.5 mm diam., obconic or turbinate, calyx tube 0.2-1 mm long, calyx lobes not clearly developed; corolla white or tinged with pink, salverform or slightly funnelform, tube 12-18 mm long, 1-2 mm diam. near the base and 2-3 mm near the mouth, often curved, lobes 5(-6), 6- 12(-20) mm long, 0.8-2 mm broad. Fruits 14-20 mm long, 10-15 mm diam., ooblong or ovoid and abruptly rounded at apex and base, blue-black in life, persistent calyx not elevated or less than 0.5 mm high and 4 mm diam.; pyrenes 1 or 2, smooth or sulcate adaxially.
Plants of poorly drained areas in lowland rain forest formations, 50-600(-900) m elevation. Flowering in February-July; fruiting in Septem- ber-February. The species is known only from the Caribbean lowlands of northern Costa Rica and the Osa Peninsula.
Coussarea psychotrioides is recognized by its re- stricted lowland habitat, glabrous parts, somewhat umbelliform inflorescences, and slightly pink flow- ers with long corolla lobes. The leaves are quite variable in shape but tend to dry greenish. This species is common at La Selva, where crushed leaves are said to have a slight odor of wintergreen. This species resembles Psychotria eurycarpa, which has larger stipules and calyx lobes, shorter corolla lobes, and an earlier flowering period (at La Selva). This species was studied by Bawa and Beach (1983) and referred to as Coussarea sp. (voucher JHB 1467); that Beach collection is also the type (ho- lotype DUKE). Specimens from the Osa Peninsula often have longer ( 1 6-20 mm) corolla lobes.
Coussarea talamancana Standl.. Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1288. 1938. Figure 48.
Shrubs or small trees, 2-5 m tall, dioecious, leafy stems 2-8 mm thick, glabrous, often drying pale yellowish green; stipules 6-16 mm long (to 22 mm below the inflores- cences), to 1 2 mm broad, united to form a sheath around the shoot apex, broadly obtuse to rounded distally, gla- brous, coriaceous, caducous. Leaves opposite, petioles 4-13 mm long, 1-2.5 mm thick, glabrous; leaf blades 12-28(-34) cm long, 4-18(-22) cm broad, broadly ellip- tic to elliptic-obovate or elliptic-suborbicular, apex abruptly narrowed to the acuminate, tip 6-18(-25) mm long, base broadly obtuse to acute, drying chartaceous, grayish green, glabrous above and below, 2° veins 7-10/ side, domatia absent. Inflorescences 3-7(-10) cm long, 3-5 cm broad, racemose panicles with short opposite
lateral branches, peduncles 5-13 mm long, 1.2-3.5 mm thick, glabrous, bracts minute or absent, pedicels 0-3 mm long. Flowers functionally unisexual, glabrous ex- ternally, hypanthium ca. 1-2 mm long (not clearly dis- tinguished from the calyx tube), calyx tube ca. 2 mm long, becoming 3-5 mm broad, cupulate, calyx lobes minute or not developed; corolla salverform, white, tube 4-6 mm long, 1.5-2 mm diam., lobes 4, 4-6 mm long and 1.2 mm broad, narrowly oblong. Fruits 14-22 mm long, 9-1 7(-24) mm diam., ellipsoid to obovoid, becom- ing white with spongy exocarp in final stages; pyrene solitary.
Plants of evergreen formations of the Caribbean lowlands and the southern Pacific slope, from near sea level to 700 m elevation. Flowering in Janu- ary-July; fruiting in every month but May. The species ranges along the Caribbean lowlands from northern Costa Rica to Bocas del Toro Province in Panama and in southern Puntarenas province.
Coussarea talamancana is recognized by the large broad leaves on short petioles, glabrous parts, united ovate-elliptic stipules, short racemiform in- florescences, smaller unisexual flowers, and larger spongy-white fruit. The type material (Cooper 10466 F) has long (to 25 mm) narrow drip tips, whereas some other Costa Rican material has shorter (to 1 0 mm) tips, but other characteristics are very similar and suggest that the material placed here is conspecific.
Coussarea sp. A aff. C. curvigemmia Dwyer, Phy- tologia 38: 215. 1978. Figure 47.
Shrubs or small trees, 2-8 m tall, leafy stems 1.5-3.5 mm thick, glabrous, grayish or yellowish green when dry; stipules 0.5-3 mm long, ovate and rounded distally or reduced to an entire ridge, glabrous. Leaves subsessile or with petioles 1-3 mm long, 0.5-1.5 mm thick, gla- brous; leaf blades 7-16 cm long, 2.5-5 cm broad, nar- rowly elliptic-oblong to elliptic-obovate, apex acumi- nate, tip ca. 1 6 mm long, gradually narrowed to the acute or cuneate base and usually slightly auriculate at the petiole, drying thin-chartaceous and grayish green, gla- brous above and below, with tufts of hairs in depressions (domatia) in the vein axils, 2° veins 4-7/side. Inflores- cences 4-7 cm long, 2-6 cm broad, paniculate with a single main rachis and short (3-12 mm) opposite or subopposite lateral branches, peduncles 1-2.2 cm long, glabrous and drying pale yellowish, flowers in distal pairs or triads, bracts subtending the flowers absent or less than 0.5 mm long. Flowers glabrous externally except for short (0.1 mm) erect whitish hairs on the ovary, hypanthium ca. 1 mm long, calyx tube 1-2 mm long, lobes 0.2-0.5 mm long, narrowly dentate; corolla nar- rowly tubular-salverform, white, tube 6-10 mm long, 0.7-1.5 mm diam., lobes 4, ca. 6-7 mm long; anthers 4-5 mm long, linear. Fruits not known (probably similar to those of C. curvigemmia: 10-13 x 5-8 mm, oblong and slightly flattened, whitish).
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Plants of evergreen rain forest of the Caribbean lowlands, from near sea level to 300 m elevation. Flowering in April and June (Barringer et al. 2642 &3615cR,F, Gdmez-Laurito 8388 CR). This spe- cies is known only from near Suretka in the Tal- amanca Valley in southern Limon Province.
Coussarea sp. A aff. C. curvigemmia is recog- nized by the thin subsessile leaves often slightly auriculate at the base and with weakly defined domatia, delicate whitish inflorescences, and small flowers with puberulent ovary and slender gla- brous corolla tubes. Coussarea curvigemmia, of central Panama, differs from the material placed here in having clearly outlined ellipsoid pit-doma- tia with few or no hairs, glabrous ovaries, and minutely papillate-puberulent corollas that dry dark. Both taxa have distinctive thin, slightly curved corolla tubes, and it may be that they are conspecific. This material also resembles Cous- sarea impetiolaris with more robust inflorescences and some species of Faramea.
Coussarea sp. B aff. C. curvigemmia. Figure 48.
Another species with very similar flowers may be represented by Burger & Gentry 8960 F from the Osa Peninsula and Bunting & Licht 793 F from the lower Rio San Juan, Nicaragua. Both of these collections have smaller (4 cm) compact capitate inflorescences and larger (20 cm) subsessile ob- ovate leaves with long (20 mm) narrow drip tips and with slightly auriculate leaf bases. However, the Nicaraguan collection has a pedunculate inflo- rescence and the Osa collection has a subsessile inflorescence, which may be immature. In con- trast, Liesner 3225 and Hammel et al. 18604 (all at CR) are intermediate with typical C. impetiolaris and indicate that the unusual specimens may be bridged by intermediates, in which case the de- scription given for C. impetiolaris needs to be ex- panded to include the collections placed here.
Coutarea Aublet
REFERENCE— A. Aiello, A reexamination of Portlandia (Rubiaceae) and associated taxa. J. Ar- nold Arbor. 60: 38-124. 1979.
Shrubs or small trees, branchlets terete, glabrous or puberulent, often with conspicuous elongate lenticels; stipules interpetiolar, short and acute, persisting. Leaves decussate or somewhat distichous, petiolate; leaf blades chartaceous, entire, some species with domatia. Inflo-
rescences terminal or apparently axillary (terminal on short axillary shoots with poorly developed leaves), flow- ers usually in open cymose groups of 3 or solitary, pe- duncles short, pedicels subtended by narrow bracts. Flowers bisexual, monomorphic, large and showy, bi- laterally symmetrical due to curvature of the corolla tube and asymmetric stamens, hypanthium turbinate, calyx lobes 5-6(-8), narrow, often unequal: deciduous; corolla funnelform to campanulate and often inflated on the lower side, white to rose or purple, corolla tube slightly curved, with a glabrous throat, corolla lobes 5-6(-8), imbricate in bud; stamens 5 or 6, inserted near the base of the corolla tube, filaments long and often twisted in bud, anthers basi fixed, linear, exserted or included: ovary 2-locular, placentas borne on the septum, ovules many in each locule. Fruits capsules, ovoid to obovoid or ob- long, flattened, coriaceous or woody, 2-locular, dehiscing loculicidally from the apex (down the center of the broad face of the capsule) to form 2 valves; seeds many, im- bricate and ascending, flattened and broadly winged with a thin margin around the circumference.
A genus of 6-10 species ranging from southern Mexico to Argentina; only 1 species is found in Mexico and Central America. The large curved corolla tubes, long free filaments, and the capsules splitting down the middle of their flattened sides make the genus distinctive. Only a few species of Rubiaceae in our flora have similarly large flowers (cf. figs. 15 and 31).
Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl. Bras. 6, pt. 6: 196. 1889. Portlandia hexandra Jacq., Enum. PI. Carib. 16. 1760; Sel. Stirp. 63, pi. 1 82, f. 20. 1 763. C. speciosa Aubl., PI. Guian. 1: 314, pi. 122. 1775. Figure 31.
Shrubs or small trees, 2-8(-18) m tall, leaf branchlets 1.7-4 mm thick, glabrous or sparsely puberulent, be- coming dark brown with elongate whitish lenticels; stip- ules 1.5-4(-5) mm long, ca. 2 mm broad at the base, intrapetiolarand forming a short (0.5- 1.5 mm) tube above the petioles, distally triangular and acute, glabrous in Central America. Leaves with petioles 2-10(-15) mm long, 0.7-1 .5 mm wide, glabrous or minutely puberulent; leaf blades 5-1 5 cm long, 2-9 cm broad, ovate to broadly elliptic or ovate-oblong, apex acute, short-acuminate or caudate-acuminate, base obtuse to rounded and sub- truncate (acute), drying thin -chartaceous or membra- naceous, glabrous or minutely puberulent on the veins above, glabrous or minutely (0.2-0.4 mm) puberulent beneath, 2° veins (4-)6-10/side, with tufts of minute hairs (domatia) in vein axils beneath. Inflorescences few- branched and with (l-)3-9 flowers, peduncles 3-30 mm long, bracts 3-5 mm long, pedicels 2-1 5 mm long, merg- ing gradually with the base of the flower, glabrous or sparsely and minutely puberulent. Flowers to 10 cm long and 4 cm broad, mostly glabrous (in Central America), hypanthium 4-7 mm long, 2-3 mm diam., calyx tube
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1.5-2.5 mm long, 4-5 mm broad, calyx lobes 4-6, 4- 9(-12) mm long, lanceolate to linear, deciduous; corolla funnelform-campanulate, white or tinged with pink, gla- brous externally, tube 45-80 mm long, 1 0-20 mm diam. distally, slightly curved, inflated before anthesis, lobes 4-6, 1-2 cm long, 7-10 mm broad, apex ovate-triangular and obtuse to acute; stamens 6, filaments to 8 cm long, anthers 14-19 mm long, 0.5-1 mm thick, yellow, ex- serted; style to 85 mm long. Fruits 25-45 mm long, 15- 28 mm broad, ca. 8 mm thick, oblong-obovate, woody when mature, surfaces dark brown and glabrous, with or without small white punctate lenticels, the valves slightly split in 2 at apex; seeds 7-14(-20) mm long, 6- 9 mm abroad, oblong to suborbicular, body of the seeds 3-4 mm diam., wing pale brown.
Trees and shrubs of both deciduous and ever- green forest formations, from near sea level to 900 m elevation. Flowering in late June-October; fruiting in November-April in southern Central America. This species appears to be much more common in seasonally deciduous forests than in evergreen forests in Costa Rica. The species ranges from southern Mexico and Belize to Argentina.
Coutarea hexandra is recognized by its thin short-petiolate leaves, few-flowered inflorescences with large flowers, the curved corolla tube, and the large flattened capsules with winged seeds. The fruits are unusual in that they split down the center of the broadly flattened halves (fig. 31). The flow- ers appear to be filled with gas before anthesis, and they are often pendulous at anthesis. The floral biology has been discussed by Haber and Frankie (1989). In northern Central America, bitter prin- ciples in the bark have been used medicinally, especially for malaria (Mabberley, 1987).
Crusea Schlectendal & Chamisso
REFERENCE— W. R. Anderson, A monograph of
the genus Crusea (Rubiaceae). Mem. New York Bot. Gard. 22: 1-128. 1972.
Annual or perennial herbs, sometimes woody at the base, decumbent or erect, stems terete or 4-angled with longitudinal ribs, pubescent; stipules interpetiolar and united to adjacent petioles to form a thin sheath, with a distal cross- vein and 2-12 setae, persisting. Leaves op- posite and decussate, sessile or petiolate, petioles adnate to the stipular sheath; leaf blades mostly ovate to lan- ceolate with strongly ascending pinnate venation, entire, domatia absent. Inflorescences terminal or axillary, cap- itate, verticillate or of congested dichasial cymes, sessile to long-pedunculate, subtended by 2, 4, or 8 (more) leaf- like bracts, pedicels short (to 2 mm) or absent. Flowers bisexual and radially symmetrical, homostylous, usually 4-parted, calyx lobes 4 or reduced to 2-3, with minute glands in the base of the sinuses between the lobes; co- rolla funnelform to campanulate, white to pink, red, or purple, corolla lobes 4, valvate in bud; stamens 4, fila- ments adnate to the upper half of the tube and free be- neath apex of the tube, anthers dorsifixed, exserted; ovary 2-locular, placenta elongate from the center of the sep- tum, with 1 ovule in each locule, stigma 2-lobed or sub- capitate. Fruits of 2 dry 1 -seeded mericarps (cocci) borne on the sides of and separating from a persisting bifid or fenestrated carpophore, mericarps indehiscent and ecos- tate, calyx dehiscing circumscissily or persisting; seeds with the persisting placenta on the adaxial face.
A genus of 13 species ranging from Arizona, U.S.A., through Mexico and Central America to western Panama. The herbaceous habit, congested subsessile flowers, often lanceolate leaves with strongly ascending secondary veins, narrow co- rolla tube (in our species) and unusual fruit help distinguish this genus. The mericarps are easily mistaken for seeds because of their smooth round- ed brownish surfaces and longitudinal adaxial sul- ci. The genus reaches its southern limit in Costa Rica and western Panama and is represented by only a few collections from Costa Rica. This treat- ment is based on Anderson's detailed monograph.
Key to the Species of Crusea
la. Flowers bright pink to magenta; leaf blades 1-5 cm broad, to 13 cm long 2
Ib. Flowers white or white-tipped with pink; leaf blades 0.5-3.5 cm broad, to 8 cm long 3
2a. Corolla tube 20-38 mm long, stigma lobes 1-4 mm long; secondary veins arising from the proximal half of the midvein; 1 800-3000 m elevation C. coccinea
2b. Corolla tube 5-18 mm long, stigma lobes 0-0.5 mm long; secondary veins arising from the
proximal '/: or 2/3 of the midvein; 0-200 m elevation C. hispida
3a. Corolla tube 5-1 1 mm long, stigma lobes 0.2-0.6 mm long; plants usually found around 2000 m
elevation in Costa Rica C. longiflora
3b. Corolla tube 1.8—4 mm long, stigma lobes 0.1-0.3 mm long; plants found in deciduous and partly
deciduous areas below 1 200 m elevation C. parviflora
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FIELDIANA: BOTANY
Crusea coccinea DC., Prodr. 4: 567. 1830. C. coc- cinea var. chiriquiensis W. R. Anderson, Mem. New York Hot. Card. 22: 45. 1972. Figure 31.
Decumbent or low perennial herbs to 1 m tall, often rooting at the nodes, leafy stems 1-3.5 mm thick, gla- brous or sparsely puberulent, quadrangular or terete; stipule sheath 4-14 mm long (including the setae), 1-6 mm wide, glabrous or puberulent, setae 3-12 and to 12 mm long and linear, evenly spaced or in a central group with fused bases. Leaves with petioles 4-20(-25) mm long, glabrous or puberulent; leaf blades 2.2-10(-l 3) cm long, l-4(-5) cm broad, narrowly elliptic to elliptic or ovate, apex acute to short- or long-acuminate, base acute and decurrent on petiole, drying stiffly chartaceous and dark, often slightly scabrous with the major veins im- pressed above, glabrous or sparsely puberulent above, sparsely hispidulous with short (0.3 mm) hairs along the veins beneath, 2° veins 3-6/side, strongly ascending, sub- parallel and arising from the proximal half of the mid- vein, minutely punctate on both surfaces. Inflorescences bracteate/involucrate heads or with verticillate flowers in the node below the terminal head, 10-15 mm diam., with 15-100 flowers in the head but few-10 flowering at the same time, involucral bracts 2 or 4, 2-4 cm long, leaf-like and often sessile within the expanded petioles and st i pular sheath of the subtending node, often with additional smaller bracts. Flowers with hypanthium 1 .4- 3 mm long, glabrous or puberulent distally, calyx lobes to 8 mm long, narrowly triangular, corolla deep red to pink, purple, or lavender, funnelform, tube (8-)20-38 mm long, gradually expanded from a narrow (3 mm) base, lobes (3.5-)5-l 1 mm long; stamens with free por- tion of the filaments ca. 4-10 mm long, filiform, anthers (2-)2.5-3.8 mm long; style as long as the corolla tube. Fruits with a broad flat carpophore, 4-7 mm long with lobes 0.7-2 mm long, cocci (1.7-)3-6 mm long, 2-2.7 mm broad, rounded-turbinate to turbinate, brown, calyx often coming off as a complete whorl (circumscissile).
Crusea hispida (Miller) Robinson, Proc. Amer. Acad. Sci. 45: 409. 1910. Crucianella hispida Miller, Gard. Diet. ed. 8, no. 4. 1768.
Erect annual herbs to 0.6 m tall, many-branched, leafy stems 0.5-4 mm thick, terete with stiff" unicellular trans- parent or whitish hairs 1.5-3 mm long, spreading or retrorse from a thickened base, smaller (0.2 mm) hairs sometimes also present; stipule sheath 2-5 mm long, 4- 10 mm broad, with 3-7 setae, conspicuously hispid. Leaves opposite, often with smaller axillary leaves from the same node, petioles 5-25 mm long, hispid; leaf blades 4-9(-l 1) cm long, l-3.6(-4.8) cm broad, narrowly ovate- elliptic to lanceolate or narrowly ovate, apex tapering gradually and acuminate, base acute to obtuse, drying thin-chartaceous, both surfaces with thin whitish hairs ca. 1.3 mm long, 2° vein 5-6/side, strongly ascending. Inflorescences solitary terminal bracteate capitulae with 40-100 closely crowded sessile flowers, subtended by 8 (4) leaf-like bracts and many linear hispid bracteoles. Flowers with glabrous hypanthium, calyx tube 0.3-1 mm long, lobes 2-6 mm long, subulate, margins with stiff hispid hairs; corolla salverform, light pink to dark red or purple, tube 5.5-12(-18) mm long, ca. 0.3 mm diam. (dried), lobes 2-5 mm long; anthers 1-1.5 mm long, exserted. Fruits with cocci 2-3.5 mm long, 1.1-2.3 mm broad, yellowish to dark brown.
A species of open grassy sites, ranging from Mexico to El Salvador and collected only recently in Costa Rica at a single locality: Costa de Pajaros, Bahia de Nicoya, Puntarenas (C. M. Taylor 249 and Wilbur 31715, both at DUKE). This population was in flower in July; it is variety hispida (with shorter corolla tubes). The broader petiolate leaves, colorful flowers, and unusual pubescence distin- guish this species.
Herbs of montane evergreen forest formations of the Pacific slope and central highlands, from 1 800 to 3000 m elevation. Probably flowering and fruiting mostly in the wet season and beginning of the dry season: May-January. The species ranges from western Mexico to western Panama.
Crusea coccinea is recognized by the herbaceous habit, short stipular sheath with long narrow setae, narrow leaves with subparallel secondary veins, and large pink to lavender flowers. Plants of Costa Rica and Chiriqui, Panama, belong to variety chi- riquensis W. R. Anderson. This variety is similar to variety coccinea in having larger corollas, an- thers, and mericarps, but variety chiriquensis dif- fers in having leaves with major veins deeply im- pressed above, a greater number of stipular setae, and pink to magenta flowers. While often collected in the Chiriqui Highlands, these plants have rarely been collected in Costa Rica.
Crusea longiflora (Willd. ex Roem. & Schult.) W. R. Anderson, Mem. New York Bot. Gard. 22: 89. 1 972. Spermacoce longiflora Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. C. brachy- phylla Schlechtend. & Cham., Linnaca 5: 165. 1830.
Erect annual herbs to 50 cm tall, stems terete or less often quadrangular, pubescent to pilose with hairs 1-2 mm long; stipule sheath 1 .4-6.5 mm long, 2-7 mm broad, with 3-7 distal setae, and 0-2 inconspicuous sessile lat- eral colleters, longest setae 0.5-3(-5) mm long, equaling or shorter than the sheath, with thin hairs 0.5-2 mm long. Leaves sessile or subsessile with petioles to 5 mm long; leaf blades 8-50 mm long, 3-2 1 mm wide, narrowly to broadly elliptic, or ovate, apex acute to obtuse, base abruptly narrowed, drying chartaceous and scabrous, sparsely hispidulous, 2° veins 2-3/side, strongly ascend- ing. Inflorescences 1-2 cm diam., small terminal brac- teate heads (or with lateral heads reduced to an axillary cluster of 1 or a few flowers), with up to 75(-l 00) flowers
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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in a head, involucral bracts 4 or 8. Flowers sometimes cleistogamous and resembling small unopened flower buds, hypanthium 0.7-1 . 1 mm long, glabrous, calyx tube 0.5-1 mm long, lobes 1-3.5 mm long and 0.2-0.8 mm wide, broadly to narrowly triangular, often ciliate along the edge; corolla white (rarely pink or lavender), tube 5- 1 1 mm long, narrow at the base and only 0.8 mm broad at apex, papillose externally, with long straight hairs within distally, lobes 1 .6-3.6 mm long, 0.8-1 .5 mm broad, nar- rowly elliptic, becoming strongly reflexed, with few long hairs at the base within; stamens with filaments 1.5-3.8 mm long, anthers 0.6- 1.3 mm long; style 5-16 mm long, glabrous, stigmatic lobes 0.2-0.6 mm long. Fruits with a bifid carpophore to 1 .5 mm long, the mericarps (cocci) 1.1-2.3 mm long, ellipsoid-cylindrical, whitish to yel- low-brown or dark brown, calyx often coming off as a whorl.
Herbaceous plants of montane evergreen for- mations, from ca. 1000 to 2700 m elevation. Flow- ering in July-December; fruiting in September- December (over the entire range). This species ranges from northern Mexico through Guatemala, with isolated occurrences in the highlands of Hon- duras and Costa Rica.
Crusea longiflora is distinguished by its erect annual habit, short stipular sheath with setae, small and narrow (often subsessile) leaves, the narrow corolla tube, and the small rounded mericarps. The long (1-2 mm) slender unicellular hairs also help distinguish this species. It has been collected infrequently in the Central Highlands at about 2200 m elevation. This species is common in Mexico.
Crusea parviflora Hook. & Arnott, Bot. Beechey Voy. 430, pi. 99. 1840. Figure 4.
Erect, trailing or decumbent herbs to about 50 cm high and 1 m long, annual or perennial, leafy stems 0.5-3.5 mm thick, with 4 longitudinal ridges or wings in early stages, with thin whitish hairs 0.3-1 .2 mm long but often glabrescent; stipule sheath 1-3 m long, 1.5-4 mm wide, often with minute spots, with 3-5 setae 1-7 mm long and with hairs to 1 mm long. Leaves with petioles 3- 9(-15) mm long, with lateral wings continuous with the lamina margin, with thin hairs on both surfaces; leaf blades 2.4-7(-9) cm long, 1-3.6 cm broad, narrowly to broadly elliptic or lanceolate, apex bluntly to sharply acute, base gradually (abruptly) narrowed and decurrent on petiole, drying thin-chartaceous and often grayish green, pilose on both surfaces with thin whitish hairs 0.3-1 mm long, 2° veins 3-5/side. Inflorescences brac- teate heads, with 10-many flowers per head and a ma- jority of the flowers in bloom at one time, the heads terminal or axillary to distal leaves, subglobose and 1- 2 cm broad, often on long (to 1 5 cm) peduncles, invo- lucral bracts to 3 cm long and leaf-like, pedicels short. Flowers with small (0.5-1 mm) glabrous hypanthium, calyx tube 0.5-1.3 mm long, lobes 0.6-2.5 mm long, 0.2-0.5 mm broad and linear-triangular, with thin hairs
on the outer surface, calyx persisting on the fruit; corolla white or the lobes tipped with pink, funnelform, tube \.8-4 mm long, 0.7-1.4 mm broad at the throat, usually glabrous externally and with longer hairs on the distal portion inside, lobes 1.3-2.8 mm long, 0.5-1 mm broad, narrowly triangular, erect to spreading (not reflexed), usually with small hairs externally at the tips; stamens with filaments 1.3-4.5 mm long, glabrous and becoming retracted into the corolla after anthesis; style 2.7-8 mm long, glabrous, stigmatic lobes 0.1-0.3 mm long. Fruits with mericarps 0.8-1.2 mm long, 0.6-0.9 mm broad, oblong-cylindrical or subglobose, surface slightly pitted.
Weedy plants of deciduous and evergreen for- mations, from 50 to 1600(-2000) m elevation in Central America. Flowering in October-April; mature fruits in November-April. The species ranges from western Mexico along the Pacific slope to isolated localities in Honduras, Nicaragua, and Costa Rica.
Crusea parviflora is recognized by its short weedy habit, quadrangular or slightly winged young stems, short stipular sheath with few setae, capitate in- florescences with usually only 4 broad bracts, and unusual carpophore and seed-like mericarps. The short-pedicellate flowers contrast with the sessile flowers of similar-looking species of Mitracarpus and Spermacoce. In our area it has only been found in Guanacaste Province below 500 m elevation.
Declieuxia Humboldt, Bonpland & Kunth
REFERENCE— J. H. Kirkbride, Jr., A revision of the genus Declieuxia (Rubiaceae). Mem. New York Bot. Gard. 28: 1-87. 1976.
Herbs or subshrubs, perennial and often woody at the base, branches terete or angular, glabrous or puberulent; stipules interpetiolar, subulate or reduced to a line bear- ing 1-3 linear lobes. Leaves opposite or whorled, sessile or short-petiolate, leaf blades entire and usually small, linear to elliptic, deltoid or orbicular, usually coriaceous, domatia absent. Inflorescences of terminal or axillary panicles (compound cymes), often spike-like or race- miform, solitary to 3 at the end of the stem, pedunculate, branching often dichotomous, flowers in distal cymose groups, bracts and bracteoles present or absent. Flowers bisexual and radially symmetrical, small, mostly gla- brous, hypanlhium turbinate to subglobose or obovoid, slightly compressed laterally, calyx lobes 4 (2), equal or unequal, small and persisting; corolla funnelform to tu- bular, white to blue or purple, tube pilose-villose in the throat, corolla lobes 4, short, spreading or reflexed, val- vate in bud; stamens 4, inserted in the corolla throat or between the lobes, filaments slender, anthers dorsifixed and versatile, partly or completely exserted; ovary 2-locular, ovule solitary in each locule, erect from a near- basal placenta, style slender, stigmas 2. Fruits drupa- ceous, dry or fleshy, laterally compressed and 2-parted or with 2 prominent rounded lobes, black at maturity.
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A genus of about 40 species in tropical America, with the largest number of species in Brazil. Our representative of this genus is recognized by the very short woody stems, stiff subsessile leaves, cy- mose branching of the short inflorescences, two- lobed fleshy fruits, and restriction to open grassy habitats in deciduous or partly deciduous forma- tions.
rocky or savanna-like habitats, the stiff narrow sessile leaves often three at a node, the short di- chotomously branched panicles, the small flowers, and the sessile fleshy deeply two-lobed or rounded fruit (laterally compressed when dry). Chacon (2258 CR, MO) stated that the flowers are white with blue-lavender stamens. Our material belongs to variety mexicana, which is distinguished by its narrow leaves and lack of pubescence.
Declieuxia fruticosa (Willd. ex Roem. & Schult.) Kuntze, Rev. gen. pi. 1: 279. 1891. Houstonia fruticosa Willd. ex Roem. & Schult., Syst. Veg. 3: 527. 1818. D. mexicana DC, Prodr. 4: 479. 1830. D. fruticosa var. mexicana (DC.) Standl., Field Mus. Nat. Hist., Bot. Ser. 12: 378. 1936. Figure 1.
Erect subshrubs or herbaceous 20-70 cm tall, often from a hard woody rootstock, with vertical simple or few-branched stems, leafy stems 0.7-3 mm thick, gla- brous (in variety mexicana) to pubescent, with 2 or 4 longitudinal ridges or wings (0.5 mm high), becoming terete; stipules 2-5 mm long, linear to linear-subulate, glabrous (in variety mexicana) or pubescent, deciduous. Leaves 2 or 3/node, sessile or subsessile (petiole to 1 mm long); leaf blades 20-^0(-50) mm long, 4-15(-22) mm broad, narrowly elliptic-oblong to linear-oblong or ob- long, apex acute or obtuse, base cuneate, drying stiffly chartaceous to subcoriaceous, glabrous and often lus- trous above, glabrous (in variety mexicana) or puberu- lent beneath, 2° veins (2-)3-6/side. Inflorescences ter- minal or axillary, 1-4 cm long, 1.5-5 cm broad, solitary or 3 at the ends of branchlets, open cymose paniculate with dichotomous nodes (and a sessile flower at the di- chotomy), primary peduncle 5-15(-40) mm long, bracts 1.5-3 mm long, linear, pedicels 0-1 mm long. Flowers heterostylous, hypanthium 0.3-1 mm long, calyx lobes 0.3-0.8 mm long, linear-oblong, glabrous (in variety mexicana) to villous, corolla white to blue or purple, 4.5-6 mm long, tube 3-4.5 mm long, cylindrical, lobes 4, ca. 2 mm long; anthers ca. 1 mm long, linear; style 3-4.5 mm long, stigmas 0.3 mm long. Fruits slightly fleshy and prominently 2-lobed (when both ovules de- velop), ca. 2 mm long and 3 mm broad, sessile, the lobes suborbicular, the fruit subglobose when only 1 ovule develops and ca. 2 mm diam., drying black, glabrous.
Small subshrubs of open grassy savanna-like sites in deciduous and semideciduous forest forma- tions, from ca. 20 to 1200(-1800?) m elevation. In Costa Rica the species is restricted to the Pacific slope and is common in Guanacaste and the Bue- nos Aires area of the General Valley. Flowering throughout the year (primarily in July-Septem- ber). The species ranges from southern Mexico to Brazil.
Declieuxia fruticosa is recognized by the short stature from a woody base, restriction to open
Deppea Chamisso & Schlechtendal
REFERENCE— D. H. Lorence and J. D. Dwyer, A revision of Deppea (Rubiaceae). Allertonia 4: 389- 436. 1988.
Slender shrubs and small trees, stems sparsely to densely puberulent; stipules interpetiolar, small, triangular, per- sistent. Leaves opposite (rarely in whorls of 3), opposing leaves of the same node often unequal, petiolate, leaf blades entire and pinnately veined, drying thin-charta- ceous, domatia sometimes present. Inflorescences axil- lary or terminal, scorpioid, umbellate or corymbiform to thyrsoid, the flowers in cymose groups (or rarely of solitary flowers), pedunculate, flowers pedicellate, brac- teolate. Flowers bisexual and radially symmetrical, hy- panthium hemispheric to turbinate or cylindrical, calyx lobes 4, minute or large, equal or unequal, usually with a small gland in each sinus; corolla short-funnelform to salverform, yellow (less often white, orange, or purple), corolla tube glabrous within, usually shorter than the lobes, corolla lobes 4, spreading or erect at anthesis, comoluatc in bud; stamens 4, inserted near the base of the tube, filaments short and linear, anthers dorsifixed, oblong to narrowly ellipsoid, exserted or partly included, a nectariferous disc present; ovary 2-locular, placentas elongate and peltate on the septum, ovules many in each locule and longitudinally imbricate, style slender, stigma entire or bilobed. Fruits a small dry capsule, turbinate to obovoid, coriaceous to chartaceous, usually with (6-)8 longitudinal costae, bisulcate and dehiscing loculicidally from apex, valves cleft, calyx persisting; seeds many and minute, angulate, testa foveolate and reticulate.
A genus of about 25 species, centered in Mexico and ranging through Central America ( 1 species) to southeastern Brazil ( 1 species). This account is based on the recent revision by Lorence and Dwyer (see reference above). Deppea can be confused with Hamelia and Hoffmannia, which have fleshy fruits.
Deppea grandiflora Schlcchtend., Linnaea 19: 748. 1847. D. costaricensis Polak., Linnaea 41: 566. 1877. D. floribunda Hemsl., Diagn. PI. Nov. Mexic. 3 1 . 1 879. D. longipes Standl., Contr. U.S. Natl. Herb. 18: 138. 1916. Figure 38.
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Shrubs or small trees, 1-4 m tall, leafy branchlets 1- 4 mm thick, hirtellous with short (0.3 mm) brownish hairs, at first with longitudinal ridges but becoming terete and glabresent; stipules 0.5-1 mm long, 2-6 mm wide, broadly deltoid, glabrate to densely hirtellous, the inner surface or margin with 4-8 dark brown digitate colleters. Leaves opposite and subequal or unequal at the same node (with 1 up to 2 times as long as the other), petioles 6-35(-50) mm long, 0.5-2 mm thick, glabrate or densely hirtellous on the adaxial side; leaf blades (3-)5-17(-21) cm long, (1.5-)2-7(-8.5) cm broad, elliptic to narrowly elliptic, narrowly ovate or lanceolate, apex tapering grad- ually and acuminate, base attenuate to acute (obtuse), drying chartaceous, glabrous to sparsely puberulent with minute (0.2 mm) thin whitish hairs above, minutely strigillose along the veins beneath, occasionally with tuft- ed domatia in the vein axils beneath, 2° veins 6-1 I/side and weakly loop-connected near the margin. Inflores- cence terminal (rarely axillary), 3-12 cm long and 3-15 cm broad, dichasial and corymbiform, with 15-100 flowers, peduncles 1.2-7 cm long, densely and minutely strigillose-hirtellous, primary branches 1-3 cm long and with up to 3 additional orders of branching, distal cy- mules of 2-6 flowers, pedicels 1-4 mm long, bracteoles present or absent. Flowers with hypanthium 1-2 mm long, 0.5-1 mm diam.. obconic or turbinate, with lon- gitudinal costae, calyx cup 0. 1-0.2 mm deep, lobes 0.3- 1 mm long, deltoid; corollas funnelform or rotate, yellow, glabrous, tube 1-2 mm long, corolla lobes 4, 4—7 mm long, 3-4 mm wide, obtuse; stamens 4, filaments 2 mm long, anthers 3-4 mm long, exserted, basally sagittate; style 3.5-4.5 mm long, stigmas 1.5-2 mm long, entire. Fruits 2-5 mm long, 2-4 mm diam., obconical to subglo- bose, with 6-8 prominent longitudinal costae, opening at the top; seeds 0.5-0.7 mm long, discoid and often angulate, testa foveolate.
Shrubs of evergreen montane forest formations, from 1 600 to 2700 m elevation. Flowering in Jan- uary-July, with a peak in April. Fruiting in Jan- uary-September. The species has been collected around Monteverde, Volcan Barva, and in the western Cordillera de Talamanca (to above San Isidro del General) in Costa Rica. The species ranges from central and eastern Mexico through the highlands of Guatemala and Honduras to the Chiriqui Highlands of Panama.
Deppea grandiflora is recognized by its restric- tion to higher montane forest formations, small shrubby habit, thin leaves unequal at a node and gradually tapering at both ends, bright yellow gla- brous corollas with very short tube and broad lobes, and small costate capsules opening at the top. Plants with pseudoaxillary inflorescences and unopened flowers may resemble species of Hoffmannia. Ma- terial of Chiococca is also similar.
Didymaea Hooker f.
Perennial herbs, scandent or procubent, stems brittle, usually much-branched and with long slender inter-
nodes, glabrous or rarely puberulent; stipules interpetio- lar and 2-lobed or apparently free (with 4/node), decid- uous or persisting and becoming recurved. Leaves petiolate; leaf blades entire and pinnately or subpal- mately veined, membranaceous to thin chartaceous, domatia absent. Inflorescences of solitary flowers in the axils of leaves, the pedicels long but not articulate, be- coming recurved in fruit. Flowers bisexual and radially symmetrical, hypanthium turbinate-globose, calyx en- tire; corolla campanulate to rotate, yellowish to greenish brown or purple, glabrous, corolla lobes 4, triangular and subacute, valvate in bud; stamens 4, inserted between the corolla lobes, filaments short and subulate, anthers dorsifixed; ovary 2-locular, ovules solitary in each locule and attached on the lower half of the septum. Fruits 2-parted or 2-lobed, somewhat fleshy, becoming dark blue or black and lustrous, the lobes rounded, 1 lobe usually smaller and lacking a fully developed seed; py- renes rounded.
A small genus of two to five species, ranging from Mexico to Panama. The plants are recog- nized by their herbaceous climbing Galium-like habit, small thin leaves, minute flowers solitary in the leaf axils, and fleshy, often 2-parted rounded fruit. The circumscription of species in Central America is not resolved (see below).
Didymaea alsinoides (Cham. & Schlechtend.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1291. 1938. Nertera alsinoides Cham. & Schlechtend., Linnaea 6: 413. 1831. D. alsi- noides var. australis Standl., loc. cit. 1 292. 1 938. D. alsinoides var. mollis Standl., loc. cit. 1292. 1938. D. australis (Standl.) L. O. Williams, Fieldiana Bot. 24, pt. 11:61. 1972. Figure 3.
Herbs or weak-stemmed climbers, leafy stems 0.3-2 mm thick when dry, quadrangular or with 2 or 4 lon- gitudinal ridges or wings ca. 0.5 mm high, sparsely and minutely (ca. 0. 3 mm) puberulent or glabrescent; stipules 0.5-2.5 mm long, narrowly triangular or linear, decid- uous. Leaves with petioles ( l-)2-8 mm long, 0.2-0.5 mm broad; leaf blades (5-) 7-30 mm long, (3-)4-14 mm broad, ovate to ovate-elliptic or narrowly ovate (rarely lanceo- late), apex gradually narrowed and acute with sharp tip, base abruptly narrowed or rounded and obtuse to trun- cate, decurrent on petiole, drying membranaceous (translucent), glabrous or minutely puberulent above the midvein on the upper surface, sparsely pubescent be- neath with thin hairs ca. 0.3 mm long, 2° veins 1-3/side, often with subpalmate venation in broadly ovate leaves. Inflorescence of solitary flowers in leaf axils, usually only 1 flower per node, pedicels 1-5 mm long, glabrous. Flow- ers ca. 3-4 mm long, hypanthium ca. 0.7 mm long, calyx to 0.2 mm long, truncate; corolla ca. 3 mm long, greenish purple, tube ca. 1.5 mm long. Fruits 4-6 mm long, sub- globose when 1 -seeded, deeply 2-parted and 6-8 mm broad when 2-seeded, becoming fleshy and blue at ma- turity, lustrous, drying black, usually glabrous; seeds of- ten curved and reticulate.
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FIELDIANA: BOTANY
Climbing plants of the shaded understory in montane evergreen wet forest formations, from 1500 to 2800(-3100) m elevation. Flowering and fruiting in September-June. The species (in a wide sense) ranges from Mexico to the Chiriqui High- lands of Panama.
Didymaea alsinoides is recognized in Costa Rica by its slender-stemmed clambering habit, thin, usually ovate leaves, minute flowers usually soli- tary at each node, and small fleshy blue-black fruit that are subglobose or deeply two-lobed. The pre- ceding description is based on Costa Rican and Panamanian collections referred to as Didymaea australis by Williams. They differ from the more northerly collections in having more ovate leaves often truncate at the base. Recognizing the various morphological and geographic forms of D. alsi- noides as separate species seems unwise, insofar as nearly all live in the same kind of montane habitats and there is considerable morphological variation in any one area; compare Williams' treatment (in Standley & Williams, 1975, pp. 60- 63). Compare Nertera granadensis with smaller leaves and orange fruit.
Diodia Linnaeus
Annual or perennial herbs or small shrubs, erect or scandent, usually woody at the base, stems often much- branched near the base, terete or 4-angled, glabrous or pubescent; stipules interpetiolarand united with the leaf bases to form a broad sheath, usually bearing 2-12 slen-
der awns from the truncated or rounded distal edge of the sheath, persisting. Leaves opposite or pseudoverti- cillate (with smaller axillary leaves), sessile or short-pet- iolate: leaf blades mostly narrow and often scabrous, entire or serrulate with minute scabrous hairs along the margin. Inflorescences axillary and sessile, capitulate or fasciculate (rarely of solitary axillary flowers) and often verticillate, long-spicate when the distal subtending leaves are reduced, subtended by the leaves and stipular sheaths, flowers usually subsessile. Flowers bisexual and radially symmetrical, small or minute, hypanthium obovoid to turbinate, calyx lobes 2^*; corolla funnelform to cam- panulate, white to pink or purplish, corolla tube short or long, throat glabrous to villous, corolla lobes 4(3, 5- 6), valvate in bud; stamens 4 (3, 5-6), inserted in the corolla throat, filaments filiform, anthers dorsifixed. lin- ear-oblong; ovary 2-locular (rarely 3- or 4-locular), ovule solitary and ascending in each locule, affixed to the center of the septum, style filiform and exserted, bilobed or with 2 short branches. Fruits splitting into 2 mericarps (cocci), crustaceous to slightly woody, septicidal from apex (rarely splitting at the base), without a central axis, each mericarp indehiscent or opening slightly at the base; seeds ellipsoid, longitudinally sulcate on the inner face, rounded abaxially.
A genus of about 35 species in the tropical and subtropical Americas and with a few species in Africa. The weedy growth habit, awned stipular sheaths, narrow leaves, small sessile flowers, and fruit of two one-seeded indehiscent mericarps help to distinguish this genus. Species of this genus may resemble species of Crusea and Spermacoce, but the leaves do not become pseudoverticillate. This treatment has benefited from the annotations and advice of C. D. Adams (pers. comm., 1991).
Key to the Species of Diodia
la. Plants essentially glabrous, often prostrate and restricted to the Caribbean seashore; leaves often
closely clustered on short lateral branches (1-4.5 cm long); rarely collected D. serrulata
Ib. Plants minutely to conspicuously pubescent, rarely prostrate and not restricted to the Caribbean
shore; leaves not closely clustered on short lateral branches 2
2a. Largest leaves usually less than 3 cm long, stems erect, stipular sheaths with usually glabrous awns
3
2b. Largest leaves more than 3 cm long, stems erect or clambering, stipular sheaths with awns with thin
whitish hairs distally or glabrous 5
3a. Leaves usually petiolate, to 2 cm long, thin-chartaceous and drying dark, often verticillate;
evergreen formations, 600-1200 m elevation D. brasiliensis
3b. Leaves sessile, to 3(-4) cm long, subcoriaceous and drying grayish, opposite; savannas in sea- sonally very dry deciduous forest areas, 0-300 m elevation 4
4a. Top of the fruit with erect stiff hairs 0.2-0.5 mm long, back of the fruit smooth; plants annual
(collections often with slender fibrous roots); corolla 3-4 mm long; a common species
D. teres
4b. Top of the fruit glabrous or with few minute white hairs, back of the fruit with 3 longitudinal ribs; plants mostly perennial (collections with thick taproots); corolla 8-10 mm long; rare . . .
D. apiculata
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
123
5a. Fruits dehiscent, rounded and indehiscent at the apex or opening slightly at the apex but mericarps then often separating from the base and opening slightly at the base; stipular awns mostly glabrous; leaves usually chartaceous; commonly collected Spermacoce ocymifolia
5b. Fruits (mericarps) indehiscent or opening slightly near the top; stipular awns with thin hairs; leaves usually drying stiffly chartaceous; rarely collected D. sarmentosa
Diodia apiculata (Willd. ex Roem. & Schult.) K. Schum. in Engler, Bot. Jahrb. 10: 313. 1889. Spermacoce apiculata Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. D. rigida (Willd. ex Roem. & Schult.) Schlechtend. & Cham., Lin- naea 3: 301. 1828. Spermacoce rigida Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818, not S. rigida Salisb. 1 796. Figure 6.
Herbs or subshrubs 9-40 cm tall, stems erect or pro- cumbent, perennial and usually woody and branched at the base, leafy stems 0.5-3 mm thick, hirsutulous to hispidulous with whitish hairs ca. 0.3 mm long or with sparse longer hairs to 1 mm long (rarely glabrous); stipule sheath 1-2 mm long, with 6-9 setae 3-10 mm long. Leaves opposite, often closely spaced, sessile; leaf blades 10-30 mm long, 1.5-7 mm broad, linear-lanceolate to narrowly linear-oblong, broadest near the base, apex gradually narrowed and acute with slender tip, base ob- tuse to subtruncate, margins usually revolute, drying subcoriaceous, hispidulous or hirsute above and below, 2° veins 2-3/side or obscure. Inflorescences fasciculate, ca. 5 mm broad, with 2-8 flowers at a node, flowers sessile. Flowers ca. 10 mm long, hypanthium 2-2.5 mm long, 1-1.5 mm diam. (3-4 mm at the mouth), calyx lobes 1.8-2.2 mm long, unequal, subulate-lanceolate, erect, green; corolla funnelform to campanulate, white to rose, glabrous on the exterior, tube 4-8 mm long and 1-1.5 mm diam., lobes 2-5 mm long, broadly ovate to triangular, 1.5-3 mm broad at the base; stamens with filaments 0.5-0.8 mm long, anthers 0.7-1.7 mm long; style 5-8 mm long. Fruits 2.5-3(-4) mm long, glabrous or puberulent, mericarps with 3 longitudinal costae (ribs) on the curved dorsal (abaxial) side, obovoid, flat on the inner (adaxial) side, 1.5-2.2 mm broad, calyx lobes to 2 mm long.
Plants of seasonally very dry deciduous for- mations, from near sea level to 300 m (to 1 600 m elevation in Honduras and to 2000 m in Guate- mala). Probably flowering throughout the year in northern Central America. It is primarily found in the region around Liberia, Guanacaste, and is collected in the wet season in Costa Rica. The species ranges from Mexico and the West Indies through Central America to Brazil.
Diodia apiculata is distinguished by the setose stipular sheaths, narrow sessile stiff scabrous leaves, the small axillary flowers, and the mericarps with three longitudinal ribs. The mericarps are often sparsely puberulent, in contrast to the very similar D. teres.
Diodia brasiliensis Spreng., Syst. Veg. 1 : 406. 1 824. D. polymorpha Cham. & Schlechtend., Linnaea 3: 344. 1828. D. brasiliensis var. angulata (Benth.) Standl., Carnegie Inst. Wash. Publ. 46 1 : 90. 1935. Triodon angulatum Benth., PI. Hartw. 70. 1840. Figure 1.
Small shrubs to 1 m tall, much-branched, leafy stems 0.3-2(-5) mm thick, glabrous (minutely puberulent), with 4 longitudinal ribs and 4-angled; stipule sheath small (0.5 mm), setae to 2 mm long. Leaves opposite or pseu- doverticillate with 4, 6, or 8 small leaves at a node on reduced axillary shoots (sometimes appearing to be an- isomorphous with pairs of leaves differing in size), pet- ioles 0-4 mm long; leaf blades 4-15(-20) mm long, 1- 4(-5) mm broad, oblong to elliptic, apex bluntly acute, base acute and decurrent on petiole, drying chartaceous and dark, scabrous along the margin, 2° veins 2/side or obscure. Inflorescences often spiciform with flowers ver- t id Hate in the axils of greatly reduced distal leaves, cap- itulae 3-5 mm broad, flowers sessile. Flowers 3-4 mm long, hypanthium 0.3-0.5 mm long, calyx lobes 4, ca. 0.5 mm long; corolla white, tube 1-2 mm long, lobes 4(-5), ca. 1 mm long. Fruits broadly turbinate, 2 mm long (including the calyx lobes), 2 mm diam., glabrous, the persistent calyx lobes ca. 0.7 mm long, the mericarps usually remaining attached to each other at the base, smooth on their abaxial surfaces.
Small plants of open or forested sites in ever- green formations, from 600 to 1000 m elevation in most of Central America (near sea level in Be- lize). Probably flowering and fruiting throughout the year. The species ranges from Mexico to Brazil.
Diodia brasiliensis is recognized by its many- branched erect stems and the small leaves often pseudoverticillate and drying black. This species is only known from collections by Brenes near San Ramon and it may not be native to Costa Rica. Some of this material had been annotated as C. polymorpha, now considered to be a synonym of D. brasiliensis.
Diodia sarmentosa Sw., Prodr. Veg. Ind. Occ. 30.
1788.
Herbs, stems procumbent to scandent, to 4 m long, sometimes forming tangles, leafy stems 0.7-4 mm thick, with 4 longitudinal ridges, hispidulous with hairs 0.3- 0.6 mm long; stipule sheaths 1-3 mm long, bearing more
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FIELDIANA: BOTANY
than 6 brown setae 4-8 mm long on each side, setae with minute thin hairs distally. Leaves opposite, petioles 0- 3 mm long; leaf blades 3-6 cm long, 0.8-2.5 cm broad, lanceolate to oblong-lanceolate, narrowly oblong-elliptic or narrowly ovate, apex acuminate, base obtuse, margins minutely aculeate-serrulate, drying subcoriaceous, sca- brous and hispid u 1 1 nis above and below, with short (0.1- 0.2) scabrous hairs and longer 0.2-0.5 mm hairs, 2° veins 3-5/side, deeply impressed above and prominent below, strongly arcuate-ascending. Inflorescences glomerules of 1-5 flowers in each axil, 5-1 5 mm broad, often becoming verticillate in fruit, bracts linear-lanceolate. Flowers with hypanthium ca. 2 mm long and 1 mm diam., calyx lobes usually 2 large and 2 small, to 2.5 mm long, persistent; corolla white, tube 1-1 .5 mm long, lobes 1-1 .5 mm long. Fruits splitting into 2 mericarps, 3.5-5 mm long, 2-2.8 mm broad, obovoid, persisting sepals 1-2 mm long, sparsely puberulent with short (0.2 mm) straight hairs, abaxial surface without ribs; seeds ca. 3 x 1.6 mm, dark brown, smooth.
Scandent plants of evergreen or partly decidu- ous forest formations, from near sea level to 900 m elevation (to 1 500 m in Guatemala). Flowering and fruiting in December-May. The species is rarely encountered in southern Central America, though it occurs on Cocos Island. The species rang- es from Mexico and the West Indies into northern South America, and it occurs in Africa.
Diodia sarmentosa is recognized by the thin stipular setae, stiff subsessile leaves with deeply impressed and strongly ascending secondary veins, very small axillary flowers, and small two-parted fruit.
Diodia serrulate (P. Beauv.) G. Taylor in Exell, Cat. S. Tome 220. 1940. Spermacoce serrulata P. Beauv., Fl. Oware 1: 39, t. 23. 1805. D. mar- itirna Thonning ex Schumacher, Beskr. Guin. PI. 75. 1827.
Prostate or clambering herbs, sometimes forming mats, stems to 1.5 m long, leafy stems 1.2-4 mm thick, gla- brous and brownish, at first with prominent wings but becoming quadrangular or terete; stipule sheath 1 .5-2.5 mm long, to 3 mm broad, with 3-5 linear awns 0.5-3 mm long, glabrous or with a few minute hairs. Leaves opposite or sometimes appearing verticillate (4), often crowded on short lateral branches, subsessile or with short (2 mm) winged petioles; leaf blades 10-45 mm long, 4-13 mm broad, narrowly elliptic to lanceolate, oblong-lanceolate or narrowly elliptic-oblong, apex acute, gradually narrowed to the cuneate base and decurrent on petiole, drying stiffly chartaceous and grayish brown above, glabrous above, slightly scabrous beneath, 2° veins 3-4/side. Inflorescences of solitary axillary flowers (2/ node), subsessile on pedicels ca. 1 mm long. Flowers glabrous externally, calyx lobes 4; corolla white, 6-7 mm long, lobes 4, 2 mm long; anthers 0.9 mm long. Fruits
5-6 mm long, ca. 3 mm wide, splitting into 2 indehiscent mericarps, glabrous and persisting calyx lobes to 2 mm long; seeds 2.1-2.3 mm long, 1.4-1.6 mm broad, dark reddish brown and smooth.
Rarely collected plants restricted to areas close to the Caribbean seashore. Probably flowering and fruiting throughout the year. The species ranges from British Honduras and the West Indies to Colombia; it also occurs on the west coast of Af- rica.
Diodia serrulata is distinguished by its ocean- side habitat, often prostrate habit, glabrous parts, awned stipular sheath, and solitary flowers. The leaf edges are entire and quite scabrous but not serrulate. We have seen only the following from Costa Rica: Gomez- Laurito 12109 CR, Playa Co- des near Pto. Viejo, and Shank & Molina 4336 F, Playa del Parismina, Limon.
Diodia teres Walt., Fl. Carol. 87. 1788. D. pros- trata Sw., Prodr. Veg. Ind. Occ. 30. 1788. Fig- ure 6.
Annual herbs to 40 cm tall, usually stiffly erect, stems simple or branched near the base, leafy stems 0.7-2.3 mm thick, with 4 longitudinal ribs, hispidulous and sca- brous with short (0.1-0.3 mm) and longer (1-2 mm) hairs, internodes usually 0.5-3 cm long; stipule sheaths 1-2 mm long, with 6-9 conspicuous glabrous awns 2-8 mm long. Leaves sessile, leaf blades(4-)10-30(-45) mm long, l-6(-8) mm broad, linear to linear-elliptic or lin- ear-oblong, broadest near the base, apex acute, tip 0.5- 1 .8 mm long, usually revolute along the thickened mar- gin, scabrous-hispid along the margin, drying subcoria- ceous, scabrous or hirsute above, scabrous and hispidu- lous beneath with thin white hairs 0.7-1.7 mm long, 2° veins usually obscure. Inflorescences of sessile solitary or clustered flowers in leaf axils, the glomerules 3-6 mm broad and with 2-4 flowers at each node. Flowers with hypanthium 1.5-2 mm long, calyx lobes often unequal, 0.5-3 mm long and 0.1-0.8 mm broad, green; corolla pink to purplish (white), tube 3-4.5 mm long, 0.5-0.7 mm diam. near the base, glabrous to sparsely puberulent, lobes 1 .5-2.5 mm long, 0.5-1 .5 mm broad; stamens with filaments 0.5 mm long, anthers 0.5-0.7 mm long, style 3-5 mm long. Fruits 2-5 mm long, with erect thin hairs at apex and persisting calyx ca. 1 mm long, mericarps (cocci) 1.8-2.5 mm broad, abaxial surface without lon- gitudinal ribs (in Central America), minutely pubescent.
Plants of seasonally very dry deciduous for- mations (especially sandy grass savannas) on the northern Pacific slope, 0-300 m elevation (to 1 400 m in Guatemala). Flowering and fruiting in June- January. The species ranges from the eastern Unit- ed States to South America.
Diodia teres is recognized by its short height,
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stipular setae, stiff narrow sessile scabrous leaves (drying grayish), very small axillary pink flowers, and distinctive mericarps (cocci).
Duroia Linnaeus f.
Small trees or shrubs, dioecious, branchlets tetrago- nous or terete, with expanded areas housing ants in a few species, glabrous or puberulent; stipules interpetiolar and intrapetiolar, forming a cap over the shoot apex, circumscissile and deciduous. Leaves opposite or ver- ticillate with 3-5 leaves at a node, sessile or petiolate, entire and pinnately veined, drying thin-chartaceous to coriaceous. Inflorescences terminal, 6 flowers subcapi- tate, umbellate, to corymbose or cymose, sessile or pe- dunculate, 2 flowers 1-3 at the tip of the stem, sessile or pedunculate, the flowers usually short-pedicellate. Flow- ers unisexual, radially symmetrical, usually large, hy- panthium oblong to hemispheric, calyx cupular to tu- bular, truncated and entire or with 5-9 lobes, corolla salverform, white to yellowish white, often thick or fleshy,
sericeous on the outer surfaces, glabrous or pilose in the tube within, corolla lobes 5-9(-12), convolute in bud; stamens 5-9, inserted in the corolla tube, subsessile or sessile, anthers dorsifixed, linear, acute, included; ovary 1-5-locular, with 5-6 parietal placentas sometimes joined in the center, ovules many and biseriate, stigmas 2. Fruits baccate, globose to oblong, large with a thick fleshy cor- tex, with 1-4 locules; seeds large and horizontal, flattened and suborbicular, immersed in pulp, testa thin.
A tropical American genus of about 25 species, with 2 or 3 species in Central America. Durroia hirsuta (Poepp. & Endl.) Schumann and D. petio- laris Hook. f. have swollen elongated areas of the stem with longitudinal slits, and D. saccifera Hook, f. has saccate ant domatia at the base of its leaf blades. Our species have no such structures and are not known to have an association with ants. Specimens may resemble species of Amaioua, Hippotis, and Randia.
Key to the Species of Duroia
la. Leaves subsessile, obovate, with hairs ca. 2 mm long, minor venation lineolate; calyx ca. 4 mm long, male corolla lobes 5-8 mm long D. costaricensis
Ib. Leaves petiolate, broadly oblong to elliptic-obovate, with hairs ca. 0.5 mm long, minor venation not lineolate; calyx ca. 8 mm long, male corolla lobes ca. 10 mm long D. utleyorum
Duroia costaricensis Standl., Contr. U.S. Natl. Herb. 20: 208. 1919. Figure 25.
diam., oblong to ovoid, covered with long hairs but with the surface visible beneath the hairs.
Shrubs or small trees, 5-10 m tall, leafy branchlets 2- 6 mm thick, hirsutulous with straight thin ascending hairs ca. 2 mm long, becoming reddish brown and gla- brescent; stipules 1 5—45 mm long, caducous, densely hir- sute-sericeous on the outer surfaces. Leaves closely crowded at the tips of branchlets, subsessile with petioles 2-6(-10) mm long and ca. 2 mm thick, densely hirsute; leaf blades (7-)9-19 cm long, (3-)4-7.5 cm broad, ob- long-obovate to narrowly obovate, widest at or above the middle, apex abruptly narrowed and slender acu- minate or caudate-acuminate, the narrow (ca. 2 mm) tip 5-13 mm long, gradually narrowed to the cuneate base, drying thin-chartaceous, often brown, with long ( 1 .5-2.5 mm) thin straight or slightly crooked hairs on upper and lower surfaces, 2° veins 5-8/side, 3° veins subparallel between the secondaries. Male flowers 8-15 and sub- capitate or fasciculate-cymose (tightly clustered at the tips of stems), 1 5-20 mm long, outer surfaces of calyx and corolla densely sericeous with ascending hairs, calyx 4-5 mm long, calyx lobes 6-7 and equalling the tube; corolla 11-15 mm long, white, corolla lobes 6-8, 5-8 mm long, equalling or longer than the tube. Female flow- ers l(-3) subsessile at apex of branchlets, hypanthium ca. 8 mm long, densely sericeous, with hairs 2-3 mm long, calyx lobes ca. 4 mm long, linear. Fruits 1-3 at the tips of branchlets, subsessile, ca. 22 mm long, 1 2 mm
Small trees of lowland rain forest formations of the Golfo Dulce region, from 10 to 200 m ele- vation. Flowering in March and May-June; fruit- ing in July-August and October. This species is known only from southernmost Costa Rica.
Duroia costaricensis is recognized by the sub- sessile thin hirsute obovate leaves clustered at the ends of stems, the terminal clusters of subsessile sericeous flowers, and the hirsute subsessile fruits at the tips of branches. The type collection (Pittier 6803 us) came from Sierpe, but most of the other collections come from near Rincon de Osa. Duroia hirsuta (Poepp. & Endl.) Schumann of South America has inflated stems, longer petioles, gen- erally longer leaves, pedunculate male flowers, and larger fruit. Duroia genipifolia, now Randia geni- pifolia (Standl. & Steyerm.) Lorence, of Guatemala has less hirsute leaves with better denned petioles and more elliptic blades. All three species are sim- ilar in appearance. This species also resembles Costa Rican material of Hippotis.
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FIELDIANA: BOTANY
Duroia utleyorum Dwyer, sp. nov.
Frutices vel arbores 1.5-7 m altae. Foliae lamina ob- longa vel oblongo-elliptica, 10-23 cm longa, 6-14 cm lata; venis lateralibus 7-1 1, hirsutulis; petiolis 6-23 mm longis. Flores lobis calycinis ca. 1 mm longis; corollae tubo ca. 12 mm longo, lobis ca. 10 mm longis. Fructus immaturi solitarii terminales.
TYPUS— Liesner /774(holotypuscR, isotypus MO), from Rincon de Osa, Costa Rica.
Shrubs or small trees, (1.5-)4-7 m tall, leafy stems 2- 5 mm thick, densely pubescent with thin straight yel- lowish hairs 0.3-0.7 mm long; stipules ca. 9 mm long and 4 mm broad, ovate-lanceolate with a sharp acu- minate apex, united (intrapetiolar) to form a short (1 mm) basal tube, with thin brown margins and hirsute. Leaves with petioles 6-23 mm long, 1-3 mm thick, hir- sutulous with yellowish hairs; leaf blades 10-23 cm long, 6-14 cm broad, broadly oblong to broadly obovate or elliptic-oblong, apex short-acuminate with tip 5-1 5 mm long, base obtuse to rounded and subtruncate, drying chartaceous, brown, pubescent on the major veins above, densely hirsute on the veins beneath with yellowish hairs ca. 0.7 mm long, sparsely hirsutulous between the veins, 2° veins 7- 11 /side. Inflorescences of solitary terminal female flowers, on pedicels ca. 8 mm long; male inflo- rescences of 3-7 terminal sessile flowers. Flowers whitish pilosulous to sericeous; 3 flowers with calyx tube 5-6 mm long, calyx teeth ca. 1 mm long, remote; corolla (preanthesis) white and probably salverform, tube 12 mm long, lobes 10 mm long, convolute and apparently lanceolate. Fruits solitary and terminal, the immature fruit ca. 1 5 mm diam. and globose with persisting calyx 10 mm high and 8 mm diam., densely hirsutulous with hairs ca. 0.5 mm long.
A species of lowland (10-300 m) rain forest for- mations on both the Caribbean and Pacific slopes of Costa Rica. Young flowers were collected in February on the Osa Peninsula (Jimenez et al. 650 CR, Liesner 1774 & 1853 CR, MO); young fruits were collected along the Rio Sarapiqui in May (Hartshorn 1486 CR).
Duroia utleyorum is recognized by its larger broad leaves, sessile terminal flowers, and lowland
evergreen forest habitat. This poorly known spe- cies is more likely to be confused with species of Randia than with its local cogener. Many impor- tant collections have been made in Costa Rica by Kathleen and John Utley.
Elaeagia Weddel
Trees or large shrubs, branchlets terete and puberulent; stipules interpetiolar (also intrapetiolar in some species), often covering the shoot apex, caducous or deciduous. Leaves opposite, petiolate or subsessile; leaf blades entire and pinnately veined, usually puberulent, thin-charta- ceous to coriaceous, domatia present or absent. Inflo- rescences terminal, paniculate (rarely racemose), many- flowered, pedunculate. Flowers bisexual, radially symmetrical, usually small, hypanthium hemispheric to turbinate, often sulcate on opposite sides, calyx tube cu- pular and spreading or inflated, calyx lobes 5 or none, short or inconspicuous; corolla campanulate to funnel- form, corolla white to yellow-white, tube usually short and broad, corolla lobes 5, oblong, rotate to reflexed, convolute to slightly imbricate in bud; stamens 5, in- serted between the corolla lobes, filament densely pu- berulent at the base, anthers dorsifixed, oblong, exserted; ovary 2-locular, placentas peltate and attached to the septum, ovules many in each locule, crowded and ver- tical, style short, stigmas 2 with obtuse tips. Fruits small woody capsules, crowned or encircled by the persistent calyx, loculicidally and basipetally dehiscent into 2 valves, the valves splitting at the top; seeds many, minute, elon- gate with membranous or winged testa.
A genus of about 1 2 species in Mexico, Central America, Cuba, and tropical South America. Elae- agia utilis Weddel of Colombia is the source of a protective lacquer. The genus was misspelled as Elaeagnia in the Flora of Panama (1980). Three mesoamerican species of Elaeagia were compared by Lorence (in Bol. Soc. Bot. Mexico 45: 65-69. 1 983). The small rounded capsules resemble those of Chimarrhis.
Key to the Species of Elaeagia in Costa Rica
la. Leaf blades usually 9-25 cm wide, often subsessile and sometimes auriculate at the base, with 1 1- 19 pairs of secondary veins; inflorescences usually much-branched panicles E. auriculata
Ib. Leaf blades usually 3-9 cm wide, clearly petiolate and gradually narrowed to the base, with 6-12 pairs of secondary veins; inflorescences with few lateral branches (paniculate with many branches in E. myriantha) 2
2a. Calyx cup entire distally; flowers separate and borne on long (4 mm) slender pedicels
E. nitidifolia
2b. Calyx cup with undulate or minutely lobed margin; flowers crowded and subsessile in small groups of2-5 . . 3
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3a. Corolla tubes 1-1.5 mm long, lobes ca. 1.7 mm long; leaf blades with 7-9 major 2° veins per side, without tufted domatia along the 2° veins; Cordillera de Tilaran E. uxpanapensis
3b. Corolla tubes 2.2-3.4 mm long, lobes 0.4-1 mm long; leaf blades with 9-12 major 2° veins per side, often with tufted domatia along the 2° veins; General Valley E. myriantha
Elaeagia auriculata Hem si.. Diagn. PI. Nov. Mex- ic. 32. 1879. Figure 39.
Shrubs or small trees, 3-10 m tall, leafy branchlets 3- 8 mm thick, puberulent or glabrescent; stipules 20-50 mm long, to 20 mm broad, oblong and rounded at apex, reddish brown and with subparallcl venation. Leaves with petioles 0-10 mm long, 1.5-3 mm thick, often dif- ficult to see on auriculate leaves; leaf blades ( 1 2-) 1 5-40 cm long, (6-)9-25 cm broad, elliptic-obovate to broadly elliptic or obovate, apex abruptly narrowed or rounded and obtuse to short-acuminate, gradually narrowed to the cuneate or auriculate base, drying chartaceous to subcoriaceous, dark brown above, upper surface gla- brous to short (0.2-0.5 mm) pubescent, lower surfaces sparsely to densely pubescent with short stiff hairs, 2° veins 1 l-16(-19)/side, 3° veins usually subparallel and the 4° veins raised beneath. Inflorescences 12-20 cm long, to ca. 20 cm broad, many flowered and much- branched panicles, solitary or 3 from the terminal leaf- bearing node, primary peduncles 3-5 cm long, with 3- 4 pairs of 1° opposite lateral branches, puberulent, flow- ers in racemose or cymose distal branches, pedicels 1-5 mm long. Flowers 5-7 mm long, hypanthium ca. 1 mm long, calyx cup ca. 1 mm long (including the lobes), calyx lobes 0.5-1 mm long and 1.5 mm broad, glabrous; co- rolla white, tube 1-2 mm long, lobes 1-2 mm long and becoming reflexed; stamens with filaments 2 mm long, glabrous distally, anthers ca. 1.5 mm long. Fruits 3-5 mm long and 3-4.5 mm broad, short-ovoid and broadly rounded, calyx persistent and visible on the lower part of the capsule, opening at the top (but not splitting to the base), becoming dark brown or black; seeds 0.6-1 mm long, body of the seed 0.2-0.4 mm long, with a thin membranous wing at opposite ends.
Trees of evergreen lower montane cloud forest and rain forest formations, along the central high- lands, from Volcan Tenorio in the Cordillera de Guanacaste to San Vito de Goto Brus, and on the Caribbean slope, ranging from 350 to 1700 m el- evation. Flowering in November-March; fruiting in February-June. The species ranges from Gua- temala and Honduras to eastern Panama.
Elaeagia auriculata is recognized by the large oblong stipules, short petioles, large broad leaves often auriculate at the base, large terminal pani- cles, small flowers with truncated calyx lobes and very short corolla tubes, and short rounded cap- sules opening only at the top. The very large leaves with an auriculate base obscuring the petiole are very distinctive, but not all specimens have this kind of leaf base (see below). The small capsules make this species look like species ofChimmarhis.
Elaeagia karstenii Standley was recorded for Costa Rica's flora (Standley, 1938) on the basis of a single sterile collection: Valeria 1665 F, from Tapanti at 1300 m. The longer (10-14 mm) and slender (1.5-2 mm) petioles and the smaller sparsely puberulent leaves do resemble some ex- amples ofE. karstenii from South America. How- ever, longer petioles with cuneate (not auriculate) lamina bases do occur in specimens that otherwise appear to be typical of E. auriculata: Lent 3734 and Molina et al. 17346 (note: both of the latter collections come from below 1000 m elevation). Thus, it is possible that Valeria 1665 is an aberrant juvenile shoot of an E. auriculata plant. Alter- natively, expanding the concept of E. auriculata to include material with longer petioles, cuneate leaf bases, and smaller more glabrous leaves may require the inclusion of South American speci- mens and synonymizing E. karstenii.
Elaeagia myriantha (Standl.) C. M. Taylor & Hammel, Novon (in press). Sickingia myrian- tha Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 27. 1930. Holtonia myriantha (Standl.) Standl., Trop. Woods 30: 37. 1932. Simira myr- iantha (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 306. 1972. Deppea panamensis Dwyer, Ann. Missouri Bot. Gard. 67: 145. 1980.
Trees, 8-25 m tall, leafy stems 2-7 mm thick, gla- brous, drying brownish or grayish; stipules 3-5 mm long, short-tubular with rounded lobes, often splitting between the leaf bases and remaining entire above the petioles (ligulate), translucent, persisting. Leaves with petioles 5- 20(-30) mm long, 0.8-2 mm thick, glabrous, drying red- dish brown to dark brown; leaf blades (5-)9-20 cm long, (2-)3.5-7(-8) cm broad, elliptic-oblong to elliptic or el- liptic-obovate, apex acuminate with tip 10-15 mm long, base gradually narrowed and acute or cuneate and de- current on petiole, drying chartaceous, greenish to dark brown, glabrous above, glabrous or minutely (0.05 mm) papillate-puberulent on the veins beneath, 2° veins 9- 12/side, domatia with short hairs present in the vein axils. Inflorescences 10-22 cm long, 12-18 cm broad, pyramidal with usually opposite 1° branches, peduncles 8-40 mm long, 1.5-2.5 mm thick, minutely puberulent, larger (2 cm) narrow leaf-like bracts present or absent, with smaller (0.5-3 mm) bracts and bracteoles subtend- ing branches and flowers, flowers usually subsessile in groups of 2-5. Flowers glabrous externally, hypanthium 1-1.5 mm long, obconic, calyx cup 0.1-0.3 mm long.
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FIELDIANA: BOTANY
1.3 mm diam., calyx lobes 0.2-0.4 mm long, broadly triangular; corolla tubular-campanulate, cream white, tube 2.2-3.4 mm long, ca. 2 mm diam., lobes 5, 0.4-1 mm long, triangular, slightly imbricate in bud; filaments exserted 0.5-2 mm, anthers ca. 1.5 mm long; stigmas 1- 2 mm long. Fruits ca. 3 mm long, 3 mm broad at the top when opened, dark brown; seeds 0.3-0.5 mm long, broad, reticulate.
Collected with flowers in January in the General Valley at 975 m elevation (Skutch 2387 us, the only Costa Rican collection). This species is also known from the Andes of Colombia and Vene- zuela.
Elaeagia myriantha is distinguished by the large pyramidal terminal panicles of small white flowers with broad cylindrical corolla tube and minute lobes and the almost glabrous leaves with pubes- cent domatia in the vein axils. These are unusual in that they are found in the axils of some 3° veins as well as along the midvein. The Colombian type (H. H. Smith 1810F) has smaller leaves than most other collections.
Elaeagia nitidifolia Dwyer, Ann. Missouri Hot. Gard. 67: 157. 1980. Chiococca jefensis Dwyer, loc. cit. 67: 88, f. 19. 1980. Figure 39.
Trees to 1 2 m tall, leafy stems 2-4 m thick, glabrous, becoming terete; stipules 3-5 mm long, to 4 mm broad, cylindrical at first but splitting, rounded above the pet- iole and deeply split between the petioles (becoming lig- ulate), glabrous. Leaves closely clustered distally, petioles 6-17(-25) mm long, 1-2 mm thick, glabrous; leaf blades 10-23 cm long, 3.5-8 cm broad, oblanceolate to nar- rowly oblong-obovate or elliptic-oblong, apex short-acu- minate, tip 5-8 mm long, gradually narrowed to the cuneate base and decurrent on petiole, drying charta- ceous, subglabrous above, minutely (0.05-0. 1 mm) pa- pillate-puberulent on the veins beneath, 2° veins 8-12/ side. Inflorescences solitary or 3, terminal, 8-1 6 cm long, open paniculate with distant cymose flower groups, pe- duncles 3-5 cm long, glabrous, pedicels 3-10 mm long, slender. Flowers with hy pan t h i urn and calyx continuous, ca. 3 mm long and 3-4 mm broad distally, calyx cup entire distally; corolla white, tube 1-2 mm long, lobes 5, 3-4 mm long, 1.3-2 mm broad, oblong; anthers 1.5- 2 mm long; stigma lobes 2-2.5 mm long. Fruits to 6 mm long and 10 mm broad, turbinate, truncated distally.
Trees of wet evergreen forest formations of the Caribbean slope at 400-800 m elevation. The spe- cies is only known from the Rara Avis site in Heredia and was collected flowering in September (O. Vargas 128 CR, MO). The species is also known from central and eastern Panama.
Elaeagia nitidifolia is distinguished by its un- usual ligulate stipules, the open few-flowered in-
florescences, long pedicles, and cupulate calyx with entire margin. Specimens may resemble Rustia oc- cidentalis (with porate anthers) and Simira myri- antha (with smaller corolla lobes and domatia).
Elaeagia uxpanapensis D. Lorence, Bol. Soc. Bot. Mex. 45: 66. 1983 (1984). Figure 39.
Trees, 15-40 m tall, 55-80 cm dbh, leafy stems 3-9 mm thick, flattened in early stages, glabrous and becom- ing terete; stipules 8-16 mm long. 3-7 mm broad, ovate lanceolate with overlapping margins, acute at the apex, glabrous and drying dark, deciduous. Leaves with peti- oles 3-12 mm long, ca. 1.5 mm thick, glabrous, drying dark; leaf blades 6-15(-20) cm long, 3-7(-9) cm broad, broadly elliptic to broadly oblong or slightly elliptic- obovate, apex abruptly narrowed and bluntly obtuse, base cuneate and slightly decurrent on petiole, drying stiffly chartaceous to subcoriaceous, dark grayish brown above, glabrous above, with a few thin yellowish hairs 0.3-1 mm long along the side of the midvein beneath and in leaf axils (= domatia?), 2° veins 7-9/side. Inflo- rescences solitary and terminal or axillary to distal leaves, 6-14 cm long (perhaps enlarging in fruit), narrowly pa- niculate, peduncles 2.5-5 cm long, glabrous, proximal lateral branches 1-6 cm long, usually minutely ap- pressed-puberulent, flowers sessile in opposite or ter- minal glomerules of 2-5 flowers, bracts 0.5-3 mm long. Flowers ca. 4 mm long, yellowish green, hypanthium 1- 1.8 mm long, minutely puberulent at the base, calyx lobes 0.5-1 mm long; corolla white, salverform to short - funnelform, glabrous externally, tube 1-1.5 mm long, scarcely exceeding the calyx lobes, lobes 4-5, ca. 1 .7 mm long and 1 mm wide, bluntly rounded distally, with long hairs at the mouth of the tube and base of lobes within; stamens 4-5, filaments 2-4 mm long, anthers 0.6-0.7 mm long, oblong; style and stigma 3-4 mm long, stig- matic lobes 1 mm long. Fruits not seen (probably ca. 2 mm long and 2.5 mm broad).
Only known from the wet Caribbean slopes of the Cordillera de Tilaran at ca. 900 m elevation. Immature flowers were collected in July (Haber & Bella 1928 CR, MO) and mature flowers in May (Herrera 600 CR, MO). This species is known only from the Rio Penas Blancas valley below Mon- teverde and the Reserva Forestal de San Ramon (Gomez- Laurito 12065 CR), in Alajuela Province. The species is also known from southern Mexico.
Elaeagia uxpanapensis is recognized by the op- posite subsessile flower clusters along the branches of the inflorescence, the very small flowers, and the two-locular ovule with placentas borne on the septum. The large size of the trees, flattened young leafy stems, and occasional domatia along the midvein are additional distinctive characteristics. When first discovered in Costa Rica, specimens were thought to be a species related to Warszewic- zia schwackei Schum. of South America.
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Exostema L. Richard
Shrubs or trees, branchlets usually terete, glabrous or puberulent; stipules interpetiolar, small, entire or bifid, deciduous or persisting. Leaves petiolate or subsessile; leaf blades entire, drying membranaceous to subcoria- ceous, domatia sometimes present. Inflorescences ter- minal or axillary panicles with the distal flowers usually in groups of 3, or of solitary axillary flowers, with or without bracts, pedicels present. Flowers bisexual, ra- dially symmetrical (or somewhat bilaterally symmetrical by curvature of the corolla tube), small to large, hypan- thium cylindrical or obovoid, calyx with (4-)5(-6) lobes, lobes broad or narrow; corolla short or long, salverform, tube often long and narrow, throat glabrous or barbate, corolla lobes (4-)5, oblong or linear, imbricate in bud with 2 exterior; stamens 5 inserted at the base of the corolla tube, filaments essentially free, elongate and fi-
liform, anthers basifixed, linear, usually exserted; ovary 2-locular, placentas peltate on the septum with many ascending ovules, style filiform, usually exserted, stigma capitate or bilobed. Fruits capsular, woody, 2-locular, ellipsoid to cylindrical, septicidal from apex, bivalved, valves entire or bipartite from the apex; seeds many, imbricate, flattened and oblong, testa forming a thin marginal wing around the body of the seed.
A genus of ca. 35 species of the American trop- ics, especially abundant in drier vegetation of the West Indies. The flowers with narrow corolla lobes almost equalling the length of the slender corolla tube and the long linear exserted anthers make the flowers quite distinctive. Our species are confined to deciduous or partly deciduous vegetation.
Key to the Species of Exostema
la. Flowers solitary in the axils of leaves, 6-8 cm long, corolla lobes 4-5; fruit ca. 1 cm broad, erect; leaf blades usually clearly decurrent on the petiole, domatia absent E. caribaeum
Ib. Flowers more than 1, borne in small panicles axillary to distal leaves or terminal, 2-3 cm long; corolla lobes 6; fruit 3—4 mm broad, pendulous; leaf blades very slightly decurrent on the petiole, domatia often present E. mexicanum
Exostema caribaeum (Jacq.) Roem. & Schult., Syst. Veg. 5: 19. 1819. Cinchona caribaeum Jacq., Enum. PI. Carib. 16. 1760. E. longicitspeOeTsl., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhaven 1852: 48. 1853. Figure 31.
Shrubs or small trees to 8 m tall, leafy branchlets 1 .2- 3.5 mm thick, glabrous; stipules l-3(-5) mm long, 2-3 mm broad at the base, triangular to subulate, glabrous on the abaxial surface but slightly ciliate along the distal edge, persistent. Leaves evenly spaced along the stems, petioles 6-1 2(- 16) mm long, 0.4-0.8 mm thick; leaf blades 4-1 1 cm long, 1.5-5 cm broad, elliptic-ovate to ovate, apex gradually narrowed and acuminate, tip 1-2 cm long, base obtuse to acute and decurrent on petiole, drying thin-chartaceous and dark, glabrous above, glabrous or pubescent with thin curved whitish hairs ca. 0.4 mm long beneath, often with short (0.5 mm) hairs (domatia) in the vein axils beneath, 2° veins 4-5/side. Inflores- cences absent, the flowers solitary in distal leaf axils, bracts ca. 1 mm long, pedicels 4-10 mm long, ca. 0.5 mm thick (dry), glabrous. Flowers 6-8 cm long, hypan- thium 3-5 mm long, 2-3 mm diam., glabrous, calyx lobes 0.5-1 mm long, broadly triangular, corolla white, glabrous, tube 23-45 mm long, 2-3 mm diam., often curved, lobes 25-40 mm long, ca. 2 broad and linear, becoming recurved; stamens 5. long-exserted, anthers 16-26 mm long, 0.2-0.3 mm broad (dry); stigma 2 mm long. Fruits 7-15 mm long, 6-14 mm broad, oblong- ellipsoid, smooth and dark brown externally; seeds 3-6 mm long and 3-4 mm broad, with a thin brownish wing on all sides and slightly longer at the 2 ends, body of the seed ca. 5 mm long and 3 mm broad.
Trees of deciduous and partly deciduous forests of the Pacific slope, from near sea level to 300 m elevation (to 1300 m in Guatemala). Rarely col- lected in Costa Rica (Guanacaste and adjacent Puntarenas provinces). In Central America flow- ering in June-August; fruiting in October-January and April. The species ranges from central Mexico to northern Costa Rica and the West Indies.
Exostema caribaeum is recognized by its re- striction to seasonally very dry vegetation, the long flowers solitary in leaf axils, the long narrow co- rolla tube, long corolla lobes, and long slender ex- serted anthers. The woody bivalved fruit splitting down the broader side and the seeds surrounded by a short thin wing are also distinctive. We have not seen Oersted's type of E. longicuspe from Pun- tarenas (cf. Standley, 1938, p. 1295). This species is called "caribee bark tree" and "princewood" in the Caribbean, where the wood is used for making handles. The bark is used for treating fevers and malaria in Mexico.
Exostema mexicanum A. Gray, Proc. Amer. Acad. Sci. 5: 180. 1 86 1 . Figure 40.
Small to medium-sized trees to 20 m tall with trunks to 40 cm dbh, leafy branchlets 1.3-4 mm thick, glabrous
130
FIELDIANA: BOTANY
to sparsely puberulent, lenticels conspicuous, stems be- coming grayish; stipules l-3(-4) mm long, triangular and acuminate to short-tubular and cuspidate, deciduous. Leaves well spaced along the stems, petioles 4-10(-15) mm long; leaf blades (5-)6-15(-18) cm long, (2-)3- 7.5(-10) cm broad, ovate-oblong to elliptic-oblong, apex long-acuminate, base obtuse to rounded and subtrun- cate, usually glabrous above and below except for the minute (0.2 mm) hairs (domatia) in the vein axils be- neath, 2° veins 6-9/side. Inflorescences 3-6(-l 2) cm long, 4-7(-10) cm broad, terminal or axillary to distal leaves, primary peduncles 1.5-4 cm long, glabrous or minutely puberulent, trichotomous, lenticellate, bracts 1-2 mm long, distal flowers in groups of 3, often crowded, ped- icles 2-4 mm long, minutely (0. 1 mm) puberulent. Flow- ers ca. 2 cm long, hypanthium 1 .5-2.4 mm long, glabrous or sparsely and minutely puberulent, calyx lobes 5-6, very small (0.5 mm) and deltoid/acute; corolla white to yellowish white or yellow, subglabrous or with thin curved whitish hairs, tube 7-10 mm long, 0.7-1.3 mm diam., lobes 6, ca. 9-12 mm long, 1 .5 mm broad, oblanceolate- linear; stamens 6, filaments to 1 6 mm long, anther 3-4 mm long, 0.2-0.3 mm broad; style to 2 cm long, stigmas clavate, 0.6 mm long. Fruits 10-14 mm long, 3-4 mm broad, obovoid-clavate to narrowly oblong-obovoid, surface glabrous and dark brown with whitish lenticels, the 2 valves each splitting in 2 at the top; seeds 6-9 mm long, 2-3 mm broad, body of the seed ca. 2.5 x 1 .5 mm, oblong, wing often lobed on 1 end.
Trees of deciduous and partly deciduous forests of the Pacific slope, from near sea level to 900 m elevation (to 1 300 m in Guatemala). Flowering in July-September; fruiting in July and September- February. The species ranges from Tamaulipas, Mexico, to central Panama.
Exostema mexicanum is distinguished by its de- ciduous habitat and thin leaves, close clusters of narrow-tubed flowers in short axillary or terminal panicles, the long narrow corolla lobes, exserted linear anthers, and woody bivalved capsules with winged seeds. The bark has been boiled to make a treatment for malaria and fevers. This species is rarely collected in Costa Rica.
trapetiolar, often forming a tubular sheath above the leaf base, short-triangular to long-aristate at apex (with 2 awns per node), persistent or deciduous. Leaves often held in a single plane, petiolate (rarely subsessile); leaf blades entire, usually oblong to lanceolate, usually gla- brous, lacking domatia, pinnately veined. Inflorescences terminal (rarely axillary), 1 -many-flowered, corymbose to umbellate (rarely capitate), pedunculate, peduncle and branches often with a pale blue or purple color like the flowers, pedicels usually present. Flowers bisexual and radially symmetrical, monomorphic or distylous, hy- panthium small, ovoid to turbinate, terete or angular, calyx cupular to short-tubular, truncate and entire to 4-lobed, persistent; corolla usually salverform, white to blue or lavender, tube short or long, usually narrow, glabrous, corolla lobes 4(-5), linear to lanceolate, valvate in bud, spreading or reflexed; stamens 4 (5. 6), inserted on the tube or at the throat, anthers linear, dorsifixed. linear, included or exserted; ovary 1-locular (rarely 2-locular in early stages), ovules 2 (less often 1), erect from a basal placenta, style filiform, with 2 short branch- es. Fruit baccate or drupaceous, transversely oblate (reni- form) to subglobose, often broader than long, smooth or costate, deep blue to blue-black, 1-locular and 1- (rarely 2-) seeded; seed (pyrene) horizontal, transversely in- dented (excavated) and somewhat reniform, with thin testa.
A large genus of about 1 30 species, ranging from Mexico, Central America, and the West Indies through tropical South America. The unusual fruit (usually broader than long) and single seed help to distinguish this genus. A striking sky blue to lavender color of both the flowers and inflores- cences characterizes a number of our species. The usually four-parted flowers, short-tubular stipules with only a single apex on each side, and a well- developed submarginal vein are additional char- acters found in many species. The fruits tend to have a leathery exocarp in Faramea, in contrast to the succulent exocarps of Coussarea and Psy- chotria. Despite these unusual traits, specimens of some species may look very similar to some spe- cies of Psychotria, Coussarea, and Rudgea.
Faramea Aublet
Shrubs or small trees, branchlets terete, 4-angled or flattened, usually glabrous; stipules interpetiolar and in-
Key to the Species of Faramea
la. Flowers solitary or the inflorescence with 2-3 flowers (l^t flowers per node); rarely collected .
Ib. Flowers few to many in pedunculate open inflorescences; commonly collected species 4
2a. Leaf blades 12-20 cm long; peduncles 2-5 cm long, calyx and hypanthium ca. 12 mm long
[Caribbean slope at 20-1 100 m elevation in Costa Rica] F. pauciflora
2b. Leaf blades 1-4 cm long; peduncles less than 2 cm long, calyx and hypanthium less than 5 mm long 3
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
131
3a. Inflorescences terminal; shrubs 3-4 m tall; Pacific slope, 1200-1700 m elevation
F. capulifolia
3b. Inflorescences axillary; herbaceous subshrubs ca. 1 m tall; Caribbean slope, ca. 300 m ele- vation F. myrtidfolia
4a. Leaves sessile or subsessile and the leaf blade usually rounded at the petiole, petioles 0-5 mm long 5
4b. Leaves definitely petiolate or if subsessile the leaf blades not rounded at the petiole, petioles 4-20
mm long 7
5a. Leaf blades narrowly oblanceolate, 10-16 cm long and 1.5-3.5 cm broad; Chiriqui Highlands
at ca. 2000 m elevation F. scalaris
5b. Leaf blades usually oblong, 15-27 cm long and 6-17 cm broad; 0-500 m elevation .... 6 6a. Leaves strongly tripliveined, secondary veins united by a linear (melastome-like) lateral
submarginal vein; southern Caribbean lowlands F. trinervia
6b. Leaves not strongly tripliveined, secondary veins loop-connected near margin; Golfo Dulce area F. sessifolia
7a. Leaves strongly tripliveined (like that of Melastomaceae); flowers and fruit bright blue; 0-800(-1000) m elevation F. suerrensis
7b. Leaves not strongly tripliveined and lacking strong lateral veins near the margin (sometimes present in F. eurycarpa), the 2° veins often loop-connected near the margin with the submarginal vein arcuate; flowers blue or white 8
8a. Inflorescences umbellate or fasciculate in the leaf axils, without lateral branches; flowers usually white 9
8b. Inflorescences paniculate, corymbose or cymose, with conspicuous lateral branches from the pri- mary peduncle or central rachis; flowers blue or white 10
9a. Inflorescences fasciculate, some flowers subtended by ovate bracts 5-1 1 mm long; flowers
not drying black; leaf blades to 1 7 cm long, usually oblong; 0-300 m elevation
F. parvibracteata
9b. Inflorescences umbellate, lacking conspicuous bracts; flowers drying black; leaf blades to 1 1 cm long, elliptic; 1000-2300 m elevation F. ovalis
lOa. Flowers with the corolla tube 12-22 mm long and corolla lobes 8-18 mm long, white and usually drying black; fruit slightly broader than long, often with persisting calyx tube; stipular awns 4-18 mm long [0-800 m elevation] F. occidentalis
lOb. Flowers with corolla tubes 4-13 mm long, corolla lobes 3-7 (-10) mm long, blue or white and rarely drying black; fruit distinctly broader than long, a persisting calyx tube rarely present; stipular awns 1-6 mm long 11
1 1 a. Leaf blades usually drying yellowish green beneath, with a prominent submarginal vein; inflores- cences robust with branches ca. 1 mm thick when dry; stipules early deciduous; corolla tube 6- 1 0 mm long; 500-1 700 m elevation F. eurycarpa
1 1 b. Leaf blades usually drying grayish or pale green beneath; inflorescences delicate with primary branches ca. 0.5 mm thick when dry; 0-1600 m elevation 13
1 2a. Stipules clearly tubular and persisting on the leafy stems; corolla tubes 6-12 mm long, calyx lobes 0.2-0.4 mm long F. multiflora
\ 2b. Stipules quickly deciduous, tubes short and inconspicuous; corolla tubes 2-6 mm long, calyx lobes 0-0.7 mm long 13
13a. Corolla tube slender, 0.7-1.5 mm diam., blue; 0-400 m elevation F. stenura
13b. Corolla tube broad, 1.5-2 mm diam., white; 1300-1400 m elevation F. hondurae
Faramea capulifolia Dwyer, sp. nov. fructum solitarium gerenti. Floras non visi. Fructus glo-
bosus, 6-9 mm longus, glaber, niger.
TYPUS— Folsom 4262 (holotypus MO, isotypus ?PMA),
Frutices ad 4 m alti. Foliae lamina lanccolata. 2-6 cm from Cerro Pirre, ridgetop at 1200 m elevation, Darien, longa, 0.5-1.6 cm lata, acumine ad 10 mm longo, venis Panama, lateralibus 10-16; petiolis 2-4 mm longis; stipulis 2-8
mm longis, vagina 1-5 mm longa. Inflorescentiae ter- Shrubs, 1-4 m tall, leafy stems 0.5^4 mm thick, gla-
minales, pedunculo nullo vel 1 5-20 mm longo, glabro, brous with 2 or 4 barely distinct longitudinal ridges;
132 FIELDIANA: BOTANY
stipules 2-8 mm long, sheathing tube 1-5 mm long, truncated distally with 1 small narrow lobe (2/node), glabrous, deciduous. Leaves with petioles 2—4 mm long, ca. 0.5 mm thick, glabrous, sulcate above; leaf blades 2- 6 cm long, 0.5-1.6 cm broad, lanceolate to narrowly ovate-elliptic, apex acuminate, base acute and slightly decurrent on petioles, drying chartaceous, olive green above, glabrous above and below, 2° veins 5-9/side and loop-connected ca. 1 mm from the leaf margin (major and intermediate secondaries often difficult to distin- guish). Inflorescences terminal and solitary, sessile or with peduncles to 2 cm long in fruit (after the loss of the distal leaf pair), peduncles ca. 0.4 mm thick and glabrous, pedicels 0-2 mm long, fruit solitary. Flowers solitary, ca. 10 mm long, glabrous, corolla ca. 8 mm long, rose- colored, tube ca. 5 mm long, ca. 0.9 mm diam. Fruits 6-9 mm diam., glabrous, persisting calyx 0.5-1 mm high, surfaces smooth, brilliant blue or becoming black.
Understory plants in wet cloud forests from 1 000 to 1700 m elevation. Flowering in May; fruiting in June-July and December. This species is known only from the Cordillera de Tilaran area in Costa Rica and eastern Panama.
Faramea capulifolia is recognized by its small leaves, glabrous parts, and solitary terminal flow- ers and fruits. The Costa Rican material has some- what smaller leaves (to 4.5 cm long) and blue fruits, whereas the type has leaves to 6 cm long and fruits becoming black. The fruits are subtended by a node where the leaves have fallen, with the distal internode appearing to be a peduncle. Flowering data was added in proof, based on Gomez- La urito 12431 F. Also placed here are Barringer et al. 4194 CR, F, Dryer 1711 F, and Zamora et al. 647 CR, F.
Faramea eurycarpa J. D. Smith, Bot. Gaz. 44: 1 1 3. 1907. F. bocaturensis Dwyer, Ann. Missouri Bot. Gard. 67: 163. 1980. Figure 45.
Shrubs or small trees to 4(-8) m tall, leafy branchlets 1.5-5 mm thick, glabrous and drying yellowish green; stipules to 1 6 mm long, united to form a tubular sheath 3-8 mm long, 2-4 mm broad, with awns 1-3 mm long, deciduous. Leaves with petioles 3-7(-10) mm long, 1- 1.5 mm thick, glabrous; leaf blades (5-)9-16(-19) cm long, (2-)3-5.5 cm broad, narrowly oblong to elliptic- oblong, apex obtuse or rounded and acuminate to cau- date-acuminate, tip 10-14(-20) mm long and ca. 2 mm broad, base cuneate (or slightly rounded at the petiole), drying chartaceous, usually yellowish green below with the midvein yellow, glabrous above and below, 2° veins 9-14/side and loop-connected near the margin to form an arcuate submarginal vein 2-4 mm from the edge, with less distinct secondary veins paralleling the major sec- ondaries. Inflorescences terminal and solitary, to 14 cm long and 1 2 cm broad, pyramidal panicles with opposite bracteolate branches, glabrous, peduncles 2-5 cm long, stout and 1.3-3 mm thick (dried), the slender pedicels
4-10 mm long. Flowers with hypanthium 0.7-1.5 mm long, calyx cup 0.5-1.5 mm long, calyx teeth 4 or 5, 0.3- 1 mm long, narrow distally; corolla salverform, blue, purple, or white with purple apex, tube 6-10 mm long, 1-1.5 mm diam., lobes 2-4 mm long, narrowly ovate; stamens 4, attached near the middle of the tube and subsessile, anthers 3-4 mm long; style 4-8 mm long, stigmas ca. 0.7 mm long. Fruits 6-8 mm long and 9-13 mm broad, transverse-reniform to subglobose, blue, pur- ple, or black, drying dark, smooth or slightly rugose when dried, persisting calyx less than 1 mm high.
Plants of the very wet evergreen cloud forests of the Caribbean slope, from (300-)500 to 1200(-1700) m elevation. Flowering in March- June, September, and December; probably fruit- ing throughout the year. The species ranges from Costa Rica and Panama to Colombia and Ecua- dor.
Faramea eurycarpa is recognized by the oblong leaves with slender "drip tips," greenish yellow color of the veins beneath when dried, deciduous stipules, arcuate submarginal vein, and small blue flowers with small corolla lobes. The inflorescence branches and deciduous tubular stipules are much thicker in texture than those of/", multiflora, which shares many of the same habitats as F. eurycarpa. Collections from the Chiriqui Highlands have a greater range of variation than those found in Cos- ta Rica, but they do not appear to be specifically distinct. The name F. bocaturensis Dwyer applies to the distinctive Chiriqui Highland material.
Faramea hondurae Standl., J. Wash. Acad. Sci. 18: 169. 1928. Figure 45.
Shrubs, 3-4.5 m tall, leafy branchlets 1 .3-3 mm thick, glabrous; stipules forming a short (3-4 mm) tube, ca- ducous and leaving a transverse whitish scar. Leaves with petioles 6-16 mm long, 0.8-1.4 mm broad, glabrous; leaf blades (6-) 1 0- 1 9 cm long, ( 1 . 5-)3-5 cm broad, nar- rowly oblong to narrowly elliptic-oblong, apex long-acu- minate to caudate-acuminate, tip 12-20 mm long, base acute, drying chartaceous and dark green above, glabrous above and below, 2° veins ca. 14/side (but with less prominent 2° veins between the major), an arcuate sub- marginal vein present near (ca. 2 mm) the margin and connecting the secondaries. Inflorescences terminal or axillary to distal leaves, solitary or 3, 5-8 cm long, to 6 cm broad, paniculate with 2 or 3 pairs of opposite branches, primary peduncles 20-30 mm long, bracts 2- 6 mm long and mucronate, distal bracteoles 0.5-3 mm long, pedicels 2-5 mm long, slender, glabrous. Flowers glabrous, hypanthium ca. 1 mm long, turbinate, calyx tube 0.5-1 .5 mm long, campanulatc. with 4 minute lobes; corolla salverform, white, tube 2-4 mm long, 1 .5-2 mm diam.. lobes 1-2 mm long, obtuse; anthers ca. 1.5 mm long, style and stigma ca. 2.3 mm long. Fruits unknown.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
133
A problematic species known only from two collections (Standley 36534 us, 37890 us the type) collected between 1 300 and 1 700 m elevation near Bajo La Hondura, San Jose, in March 1924. The short, relatively broad white corolla tubes make this species unique among our species of Faramea and make the generic placement doubtful. How- ever, a dissection of an ovary (Standley 36534) showed two ovules in a single locule, consistent with Standley's placement of this species in Fara- mea.
pearance to F. stenura but the tubular stipules of F. multiflora are persisting, the calyx lobes are less well developed, and distal inflorescence branches lack the small bracts characteristic of F. stenura. Central American collections differ in minor ways (generally smaller thinner leaves and inflores- cences) from South American collections, but all appear to be part of the same wide-ranging species.
Faramea myrticifolia Dwyer, sp. nov.
Faramea multiflora A. Rich., ex DC., Prodr. 4: 497. 1830. F. talamancarum Standl., Publ. Field Columb. Mus., Hot. Ser. 4: 332. 1929. Figure 45.
Shrubs or small trees, (l-)2-5(-6) m tall, leafy branch- lets 1-3.5 mm thick, terete and glabrous; stipules 4- 8(-l 3) mm long, 2-3 mm broad but broader beneath the inflorescence, the tubular basal sheath 2-5 mm long, awns l-5(-8) mm long, persisting. Leaves with petioles 3-7(-l 1) mm long, 0.7-1.8 mm thick; leaf blades (6-)8- 13(-17) cm long, 1.5-4(-7) cm wide, oblong to oblong- obovate or narrowly elliptic-oblong, apex gradually or abruptly narrowed and caudate-acuminate or acuminate, tip 7-15 mm long and straight or falcate, base acute or obtuse, leaves drying thin-chartaceous to chartaceous and dark greenish above, 2° veins 6-12/side (and lesser parallel 2° veins between the major), united near the margin by a slender arcuate submarginal vein 2-4 mm from the leaf edge. Inflorescences terminal, solitary (or 3), 5-1 4 cm long, to 9 cm broad, paniculate with opposite branches 1-2 cm long, bright blue, glabrous, primary peduncles 2-3(-6) cm long, basal branches subtended by small leaves or broad bracts 8-22 mm long, distal bracts absent, pedicels 3-7(-10) mm long, slender. Flowers gla- brous externally, hypanthium 0.5-1 mm long, calyx tube 0.2-0.4 mm long, calyx lobes 4, 0. 1-0.4 mm long; corolla bright sky blue, salverform, tube 6-12 mm long, 1.2-2 mm diam.. lobes 4, 4-9 mm long, 1.3-2.5 mm broad, ovate-oblong; anthers 2-3 mm long. Fruits 6-8 mm long, (8-) 10- 13 mm broad, oblate or transversely reniform, laterally compressed (oblong in cross-section), surface smooth and without costae, blue-black at maturity; py- renes solitary.
Understory shrubs of wet evergreen forest in- teriors and forest edges, from 20 to 1600 m ele- vation. Flowering in every month but October (flowering is mainly in May-June at La Selva); fruiting throughout the year. The species ranges from northern Costa Rica to Brazil and Bolivia.
Faramea multiflora is recognized by the thin smaller leaves (often drying grayish green be- neath), narrow acuminate apices, persisting tu- bular stipule sheaths, and bright blue flowers and inflorescences. This species is very similar in ap-
Suffrutices 0.3-0.4 m alti; ramulis multis glabris. Foli- ae lamina elliptica 2-4 cm longa, 8-15 mm lata, glabra, venis lateralibus 6-8; petiolis 3-6 mm longis; stipulis 4- 5 mm longis. Inflorescentiae axillares, flores 2-5 geren- tes, pedunculis 2—4 mm longis; pedicellis 1-2 mm longis. Flores glabri; calycis cupula ca. 0.5 mm longa, lobis ca. 0.5 mm longis, lilacinis; corolla viridi-alba, ca. 4 mm longa. Fructus ca. 8 mm lati, globosus, azureus, glaber.
TYPUS— L. D. Gomez et al. 23401 (holotypus CR, iso- typus MO), from Las Brisas de Pacuarito, Limon, Costa Rica.
Herbaceous subshrubs, ca. 40 cm tall, main stem un- branched for 20 cm and with many slender distal op- posite horizontal branches, leafy stems 0.5-1 .3 mm thick, glabrous, prominently 2- or 4-ridged; stipules 4-5 mm long, with a short (0.5-1 mm) truncated tube and single filiform central awn 3-5 mm long, persisting. Leaves with petioles 3-6 mm long, 0.3-0.9 mm thick, glabrous; leaf blades 2-4 cm long, 8-15 mm broad, elliptic to elliptic-oblong, apex acute or short-acuminate, base acute and decurrent on petiole, drying chartaceous, dark green above, glabrous above and below, 2° veins 6-8/side and loop-connected 1-1.5 mm from the margin. Inflores- cences axillary to leaves, 1 or 2/node, cymose umbellate with 2-5 flowers on a short (2-4 mm) glabrous peduncle drying dark, glabrous, bracts ca. 2 mm long and linear, pedicels 1-2 mm long. Flowers glabrous and drying blackish, hypanthium 0.5-1 mm long, calyx tube ca. 0.5 mm long and truncated with linear lobes 0.5 mm long; corolla greenish white, ca. 4 mm long and 0.7 mm diam. in bud. Fruit globose, ca. 8 mm diam., intense blue but drying black, usually I/node.
Distinctive little plants with many (ca. 1 2) distal lateral horizontal branches, small leaves, unusual stipules, and minute flowers. Presently known only from the type, collected near Siquerres, Limon, 1 8 April 1985, at about 300 m elevation. It appears to be related to F. cobana J. D. Smith of Honduras, but that species has larger (8- 1 0 cm) leaves, longer peduncles (to 1 5 mm), and a higher-elevation hab- itat.
Faramea occidentalis (L.) A. Rich., Mem. Fam. Rubiac. 96. 1830. Ixora occidentalis L., Syst. Nat. ed. 2: 893. 1759. F. zeteki Standl., Contr.
134
FIELDIANA: BOTANY
Arnold Arbor. 5: 147. 1933. F. belizensis Standl., Carnegie Inst. Wash. Publ. 461: 90. 1935. F. standleyana L. O. Williams, Phytologia 26: 490. 1973. Figure 44.
Shrubs or small trees, 2-6(-10) m tall, to 20 cm dbh, leafy branchlets 1.5-5 mm thick, glabrous, terete, often dichotomously or trichotomously branched; stipule sheath 2-8 mm long, 5-8 mm wide at the base, with awns 4-12(-18) mm long, deciduous. Leaves with pet- ioles 6-15 mm long, 1-2 mm thick, glabrate; leaf blades 8-18(-21)cm long, (2.5-)3.5-9.5(-l l)cm broad, oblong or elliptic to narrowly elliptic-obovate (less often ovate- oblong), apex rounded to obtuse and acuminate to cau- date-acuminate, the narrow (2 mm) tip 4-17 mm long, base acute to obtuse or rounded and subtruncate, drying chartaceous to stiffly chartaceous, glabrous above, gla- brous or minutely puberulent beneath, 2° veins 6- 107 side and only loosely loop-connected near the margin (a definite submarginal vein absent), with 1-3 more weakly defined secondaries between the major secondary veins. Inflorescences terminal or less often axillary, solitary or several, 5-12 cm long, equally broad, umbelliform or trichotomous. few-branched and with 3-9 flowers, often drying black, peduncles 1.5-6 cm long, lateral branches opposite and few, distal flowers usually in groups of 3, bracts 4 mm long, linear, pedicels 3-12(-20) mm long. Flowers white but drying black, sweet scented in life, probably nocturnal, hypanthium ca. 2 mm long, oblong, calyx cup 1.5-3 mm long, teeth absent or minute; corolla salverform, tube 10-19(-22) mm long, 2-3 m diam., narrowly cylindrical, lobes 4, 8-16(-25) mm long, 1.3- 3 mm wide near the base, lanceolate to narrowly ovate; stamens 4, filaments 0.5 mm long, anthers ca. 8 mm long, slightly exserted; style as long as the corolla tube or 'Ath as long, stigma 2.5-5.5 mm long. Fruits 6-9 mm long, 9-14 mm diam., globose to subglobose-oblate. per- sisting calyx 1-2 mm long and 2-3 mm diam., drying black.
Trees and shrubs of evergreen forest formations, from near sea level to 1 000 m elevation. Flowering in February-July and October in southern Central America (primarily in June in Costa Rica); prob- ably fruiting throughout the year. The species ranges from southern Mexico, Central America, and the West Indies through tropical South Amer- ica.
Faramea occidentalis is recognized by the large slender flowers drying black, relatively few- branched and few-flowered inflorescences also drying black, and rounded fruit slightly shorter than broad. This species varies greatly in leaf form and it does not have a well-developed submarginal vein. Though wide-ranging, it has not been col- lected very often in Costa Rica. This species is closely related to F. luteovirens with larger flowers. Fruiting and sterile material may be difficult to separate from F. eurycarpa.
Faramea ovalis Standl., Contr. U.S. Natl. Herb. 18: 138. 1916. F. quercetorum Standl., J. Wash. Acad. Sci. 18: 168. 1928. Figure 45.
Small trees or shrubs, 2-6(-13) m tall, to 20 cm dbh, leafy branchlets 1-4 mm thick, glabrous, becoming gray- ish; stipules 3-13 mm long, the basal cupulate tube 1- 2 mm high, with a single slender awn 3-10 mm long, often persisting. leaves not closely congested, petioles 3-10 mm long. 0.7-1.8 mm thick, glabrous; leaf blades 3-1 1 cm long, 1.5-4.8 cm broad, elliptic-oblong to ob- long or narrowly elliptic, apex abruptly narrowed and acuminate or caudate-acuminate, the tip 6-1 3 mm long, base acute to obtuse, drying stiffly chartaceous and gray- ish green to dark green above (slightly paler beneath), glabrous above and below, 2° veins (3-)5-8/side and loop-connected distally, with a vein-like edge along the leaf margin. Inflorescences terminal (axillary), 3-7 cm long, to 4 cm broad, umbellate with 3-5(-9) flowers, peduncles 8-24 mm long, 0.5 mm diam., glabrous and drying dark, pedicles 4-18 mm long, ca. 0.5 mm thick. Flowers glabrous externally, drying dark, hypanthium 1-
2 mm long, calyx tube 0.5-2 mm high, teeth to 0.5 mm long; corolla salverform to somewhat funnelform, white or tinted with pink-purple, tube (7-)9-14 mm long, 1 .2-
3 mm diam., lobes 4-6 mm long; stamens 4. Fruit 6-10 mm diam., globose, blue drying black and smooth, per- sisting calyx tube 1-2.5 mm long and ca. 2 mm diam.
Plants of moist evergreen cloud forests, 1 000- 2300 m elevation (400-600 m on Volcan Orosi). Flowering in March-June and December; prob- ably fruiting throughout the year. The species is found in the northwestern cordilleras, eastern parts of the Cordillera de Talamanca, and the Chiriqui Highlands.
Faramea ovalis is recognized by its cloud forest habitat, lack of pubescence, smaller often caudate- acuminate leaves, umbellate inflorescences with relatively few large flowers, and globose black fruits. The species resembles smaller-leaved specimens of F. occidentalis and some species of Coussarea.
Faramea parvibractea Steyerm., Mem. New York Bot. Gard. 17: 376. 1967. Figure 45.
Shrubs or small trees 2-8 m tall, leafy branchlets 1-4 mm thick, glabrous and drying greenish, strongly qua- drangular; stipules 6-15 mm long, 0.5-1 mm broad, encircling the shoot apex, quickly caducous. Leaves with petioles 5-14(-20) mm long, ca. 1 mm thick, glabrous; leaf blades 7-17 cm long. 1.5-6(-7.5)cm broad, elliptic- oblong, ovate-oblong, to oblong-lanceolate, apex acu- minate to caudate-acuminate, tip 6-12 mm long, base acute to obtuse, drying chartaceous, glabrous above and below, 2° veins 6-10/side. weakly loop-connected 3-5 mm from the margin. Inflorescences fasciculate or pseu- doumbellate from distal nodes, to 6(-10) cm long and equally broad, with 6-ll(-19) primary peduncles 11-
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20(-30) mm long, flowers usually borne in distal groups of (l-)2, 3, or 5 at apex of the primary peduncles, a few peduncles with broadly ovate leaf-like bracts 5.5-16 mm long and 3-12 mm broad at their apex, bracts subcordate and with petiole-like base ca. 1 mm long, white in life, pedicles 1-3 mm long. Flowers with hypanthium 0.5- 1.5 mm long, calyx tube 0.5-1 mm long, calyx lobes small and usually unequal (ca. 0.1 mm and 0.5 mm), corolla funnelform, white, 7-12(-16) mm long, tube 5- 8 mm long, 1.5-3 mm diam., lobes 4, 5-10 mm long, 2-3.5 mm broad, lanceolate to narrowly oblong, obtuse; stamens exserted, filaments 3-3.5 mm long, inserted at or below the middle of the tube, anthers 3.5-4 mm long, linear, bluntly rounded; style 4-6 mm long. Fruits 5-7 mm long, 8-12 mm broad, oblate and rounded in cross- section, smooth, the ribs obscure or prominent when dried; pyrene solitary.
Plants of wet lowland rain forest formations of the Caribbean slope, from 4 to 600 m elevation. Flowering in January-June and October; fruiting in February and May-August. The Costa Rican collections are mostly from between Tortugero and Limon, with a few from the Osa Peninsula. This species is also known from Panama and Vene- zuela.
Faramea parvibractea is recognized by the clus- ters of few-flowered peduncles at distal nodes, the ovate-subcordate bracts subtending some flower groups, the elliptic-oblong leaves with "drip tips," the narrow stipules enclosing the shoot apices, the glabrous white flowers, and the fruit often in um- bellate groups at the ends of leafy stems. The un- usual bracts do not appear to be developed be- neath all the flower groups, but they are large and contradict the specific name.
Faramea pauciflora Dwyer, Ann. Missouri Bot. Gard. 67: 172. 1980.
Understory shrubs or small trees, 2-5(-10) m tall, branches at right angles to main stem, leafy stems 1.5- 4 mm thick, glabrous; stipules 8-15 mm long and 4 mm broad, triangular to tubular with 2 slender awns 3-5 mm long, covering the terminal bud, glabrous, drying black, caducous. Leaves held in a single plane, petioles 4-7(-l 2) mm long, 0.9-1.5 mm thick, glabrous, drying dark; leaf blades 11-22 cm long, 4-11 cm broad, elliptic-oblong to narrowly oblong or elliptic-obovate, apex acuminate or caudate-acuminate, tip 7-17 mm long, base acute, drying stiffly chartaceous, dark brown or blackish above, glabrous above, glabrous or sparsely and minutely pa- pillate-puberulent beneath, 2° veins 8-12/side. Inflores- cences ca. 10 cm long with 1-3 flowers, terminal or ax- illary, usually I/node, peduncles 2-5 cm long, 0.7-1.5 mm thick, often with 1-2 pairs of stipule-like bracts near the base and terminated by several minute bracts, ped- icels 1 5-55 mm long, 0.5 mm thick, glabrous and drying
black, merging with the hypanthium. Flowers glabrous externally and drying black, hypanthium and calyx 10- 1 5 mm long, 2.5-4 mm diam., tube ca. 4 mm long, lobes few or unequal, 1-8 mm long; corolla salverform, white, tube 10-22 mm long, lobes 4, 13-30 mm long, rotate and with a narrow tip 4-9 mm long often held at 90°. Fruits 12-14 mm long, 10-14 mm diam., with a per- sisting calyx tube 7-9 mm long and 3-4 mm diam., dark blue drying black, pendulous.
Plants of the wet Caribbean slope at elevations from 20 to 1100 m. Flowering in March-April; fruiting in March-April and August-September in Panama. The species ranges from near Nuevo Ar- enal southward to Colombia.
Faramea pauciflora is unusual because of its glabrous parts, usually axillary inflorescences with one to three long white flowers and long pedicels. Two Costa Rican collections have been seen: Her- rera 2556 and A. Smith 1644. This species is close- ly related to F. luteovirens Dwyer of Panama with thicker leaves, fewer 2° veins, and shorter calyx. Compare F. occidentalis and the pubescent but similarly few-flowered Coussarea enneantha (fig. 46).
Faramea scalaris Standl., Contr. U.S. Natl. Herb. 13: 139. 1916.
Shrubs or small trees, 1.5-6 m tall, leafy branchlets 1.5-4 mm thick, terete or quadrangular, drying brown- ish, leaves distant (3-6 cm) along the stems; stipules 4- 8 mm long, with a short-tubular sheath, truncated or deltoid at apex, with or without a short (1-2 mm) awn, persistent. Leaves subsessile with petioles l-3(-5) mm long; leaf blades (5-) 7- 15 cm long, (1-) 1.3-4 cm broad, narrowly elliptic-oblong to narrowly oblanceolate, apex acuminate, tip 7-15 mm long and straight or falcate, base gradually narrowed but often slightly rounded and subauriculate at the petiole, drying chartaceous and greenish, glabrous above and below, 2° veins 9-13/side, these arising at almost 90° angles and united by a linear or slightly arcuate lateral vein 2-5 mm from the leaf edge, a slender submarginal vein also present 0.3-1 mm from the margin in fully developed leaves. Inflorescences terminal and solitary, 3-6 cm long, paniculate with short opposite branches, primary peduncles 1 5-20 mm long, glabrous, bracts caducous, pedicels 2-4 mm long. Flow- ers glabrous, hypanthium ca. 1.5 mm long, calyx tube ca. 0.5 mm long, lobes 0.1-0.3 mm; corolla salverform, blue becoming white, tube 7-10 mm long, 1-1.5 mm diam., lobes 3-4 mm long, ca. 3 mm broad, ovate, disc 1 mm long; style ca. 7 mm long. Fruits unknown.
Plants of montane wet forest formations, from 1800 to 2300 m elevation. Flowering in January and March. The species is known only from the Chiriqui Highlands in western Panama.
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Faramea scalaris is distinguished by its narrow subsessile leaves with prominent lateral (mela- stome-like) venation, the small inflorescences, and the restricted high-elevation habitat. The unusual leaf venation suggests that this species is closely related to F. trinervia and F. suerrensis of lowland formations.
Faramea sessifolia P. Allen, Rain Forests of Golfo Dulce 409. 1956. Figure 44.
Shrubs or small trees, 3-7 m tall, leafy branchlets 2- 7 mm thick, the nodes to 10 mm thick, rectangular in cross-section (flattened and 4-angular), glabrous, drying yellowish; stipules 10-25 mm long, 3-5 mm broad at the base, triangular-subulate with a narrow distal tip, caducous. Leaves somewhat dimorphic with smaller and narrower leaves often subtending the inflorescences, ses- sile or subsessile, petioles 0-^4(-6) mm long, ca. 2 mm thick; larger leaf blades 17-27 cm long, 6-17 cm broad, narrowly oblong to ovate-oblong, smaller leaf blades 8- 1 7(-l 9) cm long and 2-4(-6) cm broad, narrowly oblong- lanceolate to narrowly oblong, apex abruptly narrowed (in larger leaves) or gradually narrowed (in smaller leaves) and acuminate, tips 8-17 mm long, base rounded and subtruncate, drying chartaceous and yellowish green, gla- brous above and below, 2° veins 12-16/side and loop- connected 2-5 mm from the margin. Inflorescences ter- minal, solitary or 3, 10- 15 cm long and 10-1 2 cm broad, paniculate with opposite many-flowered branches, pri- mary peduncles (2-)6-7 cm long, 0.7-2 mm thick, gla- brous, whitish in life and yellowish when dried, bracts 0.5-1 mm long, pedicels 1-2 mm long above small (0.5 mm) bracteoles. Flowers glabrous, hypanthium 0.5-1 mm long, calyx tube 0.2-0.5 mm long, lobes 0.2-0.4 mm long; corolla salverform, bright blue or bluish pur- ple, tube ca. 6-7 mm long and 1 mm diam., lobes 3-4 mm long, to 2 mm broad at base. Fruits 4-5 mm high, 7-9 mm broad, oblate, circular or oblong in cross-sec- tion, with 8 longitudinal costae, pale grayish brown be- coming black.
Plants of the lowland rain forests of the Osa Peninsula region, 20-400 m elevation. Flowering in April-May (Allen 5539 F, us type); fruiting in March, August-September, and November. The species is known only from southwestern Costa Rica.
Faramea sessifolia is recognized by its large sub- sessile leaves with many secondary veins and ar- cuate submarginal vein, leafy stems rectangular in cross-section, lack of pubescence, blue flowers, and restricted distribution. The veins on the lower leaf surface and inflorescences dry yellowish. This name should not be confused with F. sessilifolia (H.B.K.) A. DC. of South America.
Faramea stenura Standl., Publ. Field Columb. Mus., Hot. Ser. 4: 331. 1929. Figure 45.
Small trees to 7 m tall, leafy branchlets 1.7-4 mm thick, glabrous, drying greenish; stipules 6-10 mm long, 3-4 mm broad at the base, triangular to ovate, acute or mucronate at apex, caducous. leaves with petioles 5-10 mm long, 0.6-1.5 mm thick; leaf blades 9-19 cm long, 2.5-5.5 cm wide, narrowly oblong to narrowly oblong- obovate, apex acuminate, tip 5-23 mm long, base acute to cuneate, drying chartaceous, dark grayish green above, glabrous above and below, 2° veins 9-12/side, arising at almost 90° and weakly loop-connected near the margin, with thinner secondaries present and parallel with the larger. Inflorescences usually terminal and solitary, pa- niculate pyramidal, 3-5.5 cm long, 4-7 cm broad, blue, primary peduncles 1.5-5(-8) cm long and 1-1.5 mm thick, bracts 3-5 mm long, linear or triangular, purplish, pedicels 2-4 mm long. Flowers glabrous externally, hy- panthium ca. 0.6 mm long, calyx tube 0.2-0.3 mm long, lobes 4, 0.3-0.7 mm long; corolla salverform, pale to deep blue, glabrous, tube 5-8 mm long, lobes 4, 3-5 mm long, 1.2-1.5 mm broad, elliptic to ovate, acute at apex. Fruits 4-8 mm long, 12-15 mm broad, transversely reni- form (oblong in cross-section), rounded basally and flat- tened or depressed above, smooth and usually drying green.
Plants of lowland rain forest formations, 30- 600 m elevation (but see below). Flowering in Jan- uary-September; probably fruiting throughout the year. It ranges along the Caribbean coast, from Veracruz, Mexico, to western Panama, and in the Golfo Dulce area.
Faramea stenura is distinguished by its small blue corollas, caducous stipules, oblate fruit, and narrowly oblong leaves with long "drip tips." This species is very similar to F. multiflora, but their stipules help differentiate most collections. At La Selva F. stenura grows in low swales, whereas F. multiflora grows on the ridgetops. There are very similar plants growing at higher elevations (ca. 1200 m) in Chiriqui with larger (10 x 14 mm) ribbed fruit that dry whitish. It is not clear whether they are a high-elevation subspecies or an unde- scribed closely related species.
Faramea suerrensis (J. D. Smith) J. D. Smith, Bot. Gaz. 44: 1 1 2. 1 907. Faramea trinervia K. Schum. & J. D. Smith var. suerrensis J. D. Smith, Bot. Gaz. 31: 115. 1901. Figure 44.
Shrubs or small treelets, 2-6 m tall, leafy branchlets 2-6 mm thick (to 12 mm broad at the node), usually quadrangular in cross-section, glabrous; stipules 5-10 mm long, 3-8(-12) mm wide at the base, often united to form a short tube, entire or rounded with a small ( 1
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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mm) mucronate tip. Leaves with petioles 4-18 mm long, 1-2 mm thick; leaf blades 9-20(-28) cm long, 3-8(-12) cm broad, narrowly elliptic-oblong to oblong or narrowly oblong-oblanceolate, apex short-acuminate to long-cus- pidate, tip to 1 8 mm long in some specimens, base acute to rounded and subtruncate, drying chartaceous to sub- coriaceous, pale grayish green to yellowish green, gla- brous above and below, the major veins often becoming impressed above, 2° veins 8-16(-25)/side and with less prominent parallel 2° veins, 2° veins arising at nearly 90° from the midvein and joined near the margin by a prom- inent linear lateral vein (2-)5-7(-14) mm from the leaf edge, with a smaller submarginal vein 0.5-2 mm from the leaf edge. Inflorescences terminal and solitary (or 3), corymbose panicles with opposite branching, 9-12(-20) cm long, to 16 cm broad, bright blue, many-flowered, peduncles 2-5(-9) cm long, 2-3 mm thick, glabrous, ped- icels 2.5-9 mm long. Flowers distylous, glabrous, hy- panthium ca. 1.2 mm long, calyx tube a small (0.3 mm) rim, lobes minute or 0.5 mm long and triangular; corolla salverform, brilliant pale blue to deep blue, tube 6-10 mm long, 0.8-1.5 mm diam., lobes 4, 3-5 mm long, ca. 1.7 mm broad; stamens 4, filaments attached near the middle of the tube, anthers 2.2 mm long, included. Fruits 6-1 1 mm long, 12-16 mm broad, transversely reniform and rounded-oblong in cross-section (somewhat flat- tened laterally and flat or slightly depressed distally), longitudinal ribs slightly developed or obscure, exocarp spongy and deep blue in life; pyrenes solitary.
Plants of lowland rain forest formations on both the Pacific and Caribbean slopes, from near sea level to 800(-1000) m elevation. Flowering in Jan- uary-August and November; fruiting in every month except December. The species ranges from southern Nicaragua through Costa Rica and Pan- ama to northwestern Colombia.
Faramea suerrensis is distinguished by its larger leaves with prominent melastome-like venation, brilliant bluish inflorescences and flowers, lack of pubescence, and lowland rain forest habitat. The unusual quality of the blue coloring of the inflo- rescence and the leaf venation make this one of our most distinctive species of Rubiaceae. This species is very closely related to F. trinervia (q.v.); it is possible that the two may be conspecific as Schumann and Smith originally thought. The breeding system was studied by Bawa and Beach (1983).
Faramea trinervia K. Schum. & J. D. Smith, Bot. Gaz. 31: 115. 1901. F. bullata Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 294. 1929. Figure 44.
Shrubs or small trees, leafy stems 2-6 mm thick, gla- brous; stipules 6-10 mm long, the basal tube 2-4 mm long with rounded or acute apex and narrow awn 3-5 mm long. Leaves sessile or subsessile with petioles 2-4
mm long; leaf blades 14-31 cm long, 4-12 cm broad, narrowly oblong to narrowly oblong-obovate, apex acu- minate or caudate-acuminate, somewhat narrowed be- low the middle, base abruptly rounded and subcordate, drying stiffly chartaceous to subcoriaceous, grayish green, glabrous above and below, 2° veins 10-1 7(-22)/side (dif- ficult to separate from the less prominent secondaries), united near the margin by a linear lateral vein 5-10 mm from the leaf edge, a smaller submarginal vein also pres- ent 1-3 mm from the edge. Inflorescence solitary and terminal (or 3 with 2 axillary), 4-7 cm long, to 12 cm broad, paniculate with opposite or trichotomous branch- ing at the apex of the peduncle, basal bracts (small leaves) to 2 cm long, peduncles 3-6 cm long, 2.2-3.4 mm thick. Flowers glabrous externally, hypanthium ca. 1 mm long, obconic, calyx limb ca. 0.3 mm long with minute (0.2 mm) lobes; corolla color not known, salverform, tube 6-7 mm long, 0.5-0.9 mm diam., lobes 4, ca. 4 mm long and 1 mm broad. Fruits 7-8 mm long and 12-16 mm broad, oblate-reniform, flat or depressed centrally above, drying smooth and black.
A species of lowland Caribbean rain forest for- mations, 10-600 m elevation. Flowering in March (Tonduz 8690 us) and April (Tonduz 9583 us the type); fruiting in April (Barringer et al. 2632 CR, F) and July (/. Chacon 209 CR). This species is known only from the Talamanca valley region, southeastern Costa Rica, and Bocas del Toro Province in Panama.
Faramea trinervia is recognized by the stiff nar- rowly oblong leaves with melastome-like vena- tion, coupled with the short petioles and rounded auriculate leaf bases. This species is very similar to and may prove to be conspecific with material placed under F. suerrensis (a later name). The thicker subsessile leaves rounded at the base are a unique character combination within the more common and more widely ranging F. suerrensis, and we treat the two as distinct species, although they appear identical in most other respects. The type of F. bullata (Cooper 507 F from Panama) has short petioles but with a rounded lamina base, and we interpret it as an aberrant form of F. tri- nervia.
Ferdinandusa Pohl
Trees or shrubs, stems glabrous or puberulent; stipules interpetiolar, triangular, deciduous or caducous. Leaves opposite or verticillate, decussate or distichous, petio- late, glabrous or puberulent, entire, pinnately veined, without domatia. Inflorescences terminal or axillary to the distal leaf pair, paniculate with opposite branching and cymose to corymbose in form (rarely fasciculate or umbellate), bracteate, flowers pedicillate. Flowers bisex- ual and usually radially symmetrical (corolla tube some- times curved), monomorphic, calyx cupular, calyx teeth
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FIELDIANA: BOTANY
short, deciduous or persistent; corolla salverform to fun- nelform, white to reddish or greenish, corolla lobes 4(-5), convolute or broadly imbricate in bud, rounded distally; stamens 4(-5), filaments attached in the upper part of the tube, anthers versatile, exserted or included; ovary 2-locular with few to many ovules vertical on the axile placentas, stigmas subcapitate. Fruits a cylindrical to oblong (subglobose) woody capsule with septicidal de- hiscence from apex; seeds few to many, elliptic and flat- tened, with entire to lacerate marginal wings.
A genus of 20-25 species with 1 species in Costa Rica and Panama and the others in South Amer- ica. The elongate capsular fruit with small winged seeds and broadly overlapping corolla lobes dis- tinguish this genus. Compare this genus with ma- terial placed in Ladenbergia (with valvate corolla lobes) and Macrocnemum.
Ferdinandusa panamensis Standl. & L. O. Wil- liams, Ceiba 3: 34. 1952. Figure 40.
Trees to 20 m tall, major branches held at 90° angles or drooping, leafy stems 2-5 mm thick, terete or slightly quadrangular, glabrous or with erect hairs 0.5-1 mm long, stems slightly resinous where the hairs are broken off; stipules 5-20 mm long, narrowly triangular, acu- minate, glabrous, caducous. Leaves opposite or 3/node, smaller in size beneath the inflorescence, petioles 4-10 mm long, 1.5-3 mm thick, glabrous or pubescent; leaf blades (6-)10-18(-22) cm long, (4-)6-10 cm broad, ob- long to elliptic-oblong or ovate-oblong, apex abruptly short-acuminate, tip 3-10 mm long, base obtuse to rounded or truncate (subcordate), leaves drying stiffly chartaceous, glabrous above and below or with erect brownish hairs 0.5-1 mm long on the veins beneath and more sparsely on the surfaces beneath, 2° veins 6-9/side, arcuate-ascending near the margin but loop-connected only near apex, domatia absent. Inflorescences terminal (apparently axillary when leaves develop at the first branching node of the panicle), solitary or 3, 6-15 cm long, 6-22 cm broad, primary peduncles 2-5 cm long, bracteolate, bracts ca. 1 mm long, pedicels 5-12 mm long and merging with the flower base. Flowers glabrous, hypanthium ca. 2 mm long, calyx cup 0.5-1 mm long, 2-4 mm diam., calyx teeth 4 or 5, ca. 0.5 mm long; corolla funnelform, yellowish green or white, fleshy, tube (4-)6-18(-25) mm long, 2-3 mm diam., slightly ex- panded at the base, and much expanded distally, lobes 5, 4-6 mm long, 3-6 mm broad at the base; stamens 5, filaments linear, attached near the middle of the tube, anthers sometimes dimorphic (long and short); style to 11 mm long, stigmas ca. 1.5 mm long. Fruits (2-)3-6 cm long, 6-1 2 mm thick, narrowly oblong or cylindrical, abruptly rounded at the base and apex, with a short (0.5- 1 mm) persistent calyx, surface smooth and with obscure longitudinal ribs, brown; seeds 1-2 cm long, elliptic.
Trees of lowland Caribbean rain forest forma- tions, from near sea level to 200 m elevation. Rowers have been collected in January-May and
November-December; fruits were collected in February-July. The species is known only from Costa Rica and near Chiriqui Lagoon in Panama (von Wedel 2232 F the type).
Ferdinandusa panamensis is recognized by the elongate woody capsules with small winged seeds, fleshy flowers with broadly overlapping corolla lobes, broad leaves often rounded at the base, and the occasional presence of stiffbrownish hairs. The names cafe macho and cafecillo have been used for this species in Costa Rica. The wood is very hard and young trees are used for boat poles.
Galium Linnaeus
REFERENCES— L. Dempster, The genus Galium (Rubiaceae) in Mexico and Central America. Univ. Calif. Publ. Bot. 73: 1-33. 1978. The genus Galium (Rubiaceae) in South America, IV. Allertonia 5: 283-345. 1990. F. Ehrendorfer, Revision of the genus Relbunium (Endl.) Benth. & Hook. (Rubi- aceae-Galieae). Bot. Jahrb. Syst. 76: 516-553. 1955.
Herbs or slender vines, annual or perennial, often woody at the base (rarely shrubs), erect, decumbent or climbing, monoecious or dioecious, stems usually slen- der and with 4 prominent longitudinal ridges (square in cross section), glabrous or puberulent with thin hairs; stipules apparently absent but represented by leaf-like parts, indistinguishable from the true leaves (except that they lack axillary buds). Leaves in whorls of (3-)4-8(-10 to many) at each node, comprising the true leaves and the transformed stipules, sessile or subsessile, often nar- rowly oblong, entire or with sharp retrorse trichomes along the edge (serrulate), with 1 or 3 major veins, with- out domatia. Inflorescences terminal or axillary, often paniculate with dichasial or trichotomous branching, or with 3 terminal flowers from the distal node, inflores- cences often with bract-like reduced leaves and thin- divaricate branching (with a 4-parted involucre in spe- cies formerly placed in Relbunium), pedicels articulate beneath the flower. Flowers bisexual or unisexual, ra- dially symmetrical, usually very small, calyx tube minute or lacking; corolla rotate to campanulate or urceolate, white to yellow, green, pink, or red, corolla lobes (3-)4(-5), valvate in bud; stamens (2-)3-4(-5), filaments attached to the short tube of the corolla, anthers versatile and exserted; ovary 2-locular with 1 ovule borne on the sep- tum in each locule, styles 2 (sometimes united at the base), stigmas capitate. Fruits usually 2-lobed or 2-parted (singular when 1 ovule fails to develop), dry or fleshy, smooth and glabrous to tuberculate or densely hispidu- lous, small, finally separating into 2 1 -seeded mericarps, the mericarps rounded and indehiscent; seeds convex dorsally, attached to the pericarp, testa membranous.
A large genus of some 300-400 species, es- pecially well represented in the North Temperate
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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zone and at higher elevations in the tropics. Dempster listed 4 1 species in Mexico and Central America (3 of which are probably early introduc- tions); the genus is especially well represented in Baja California and the central highlands of Mex- ico. Galium, as a genus, is easy to recognize with its small whorled leaves on slender herbaceous
clambering 4-angled stems, minute flowers on slender pedicles, and 2-lobed little fruit. Material of Galium may be mistaken for species ofDidym- aea and Nertera. This treatment is based in large part on the annotations and publications of Laura- may Dempster.
Key to the Species of Galium
la. Leaves in whorls of 6 or 8(-10); fruit covered with ascending curved hairs G. mexicanum
1 b. Leaves in whorl of 4 at each node; fruit glabrous or with uncinate (hooked at the tip) hairs ... 2 2a. Flowers solitary in the leaf axils and subtended by a whorl of 4 leaf-like or calyx-like bracts
G. hypocarpium
2b. Flowers rarely solitary in leaf axils, rarely subtended by 4 bracts 3
3a. Fruits glabrous, leaves with 1 primary vein and no lateral veins [usually glabrous; stems with minute
(0. 1-0.2 mm) hairs] G. aschenbornii
3b. Fruits covered with ascending uncinate hairs; leaves with 1 primary vein and 2 lateral veins usually
visible 4
4a. Leaf blades usually ovate to elliptic; stems and leaves with few to many longer (0.4 mm) hairs;
inflorescences usually with few (ca. 5) flowers; corollas often hispidulous G. uncinulatum
4b. Leaf blades usually oblong-lanceolate to elliptic, stems and leaves sparsely short-puberulent (ca. 0.2
mm); inflorescences usually with more than 5 flowers; corollas glabrous G. orizabense
Galium aschenbornii Schauer, Linnaea 20: 701. 1 847. Relbunium aschenbornii (Schauer) Hemsl., Biol. centr. amer. Bot. 2: 62. 1881. Figure 3.
Creeping or climbing herbs to 1 .2 m long, with slender woody or herbaceous stems arising from a small root- stock, sometimes rooting from distal nodes, with both long (5 cm) and short (4 mm) internodes, leafy stems 0.3-1 mm thick, glabrous or with short retrorse hairs 0. 1-0.2 mm long; stipules leaf-like. Leaves usually 4/node, petioles 0. 1-0.5 mm long; leaf blades 4-8(-14) mm long, 1 .2-3(-4) mm broad, narrowly oblong or elliptic-oblong (to lanceolate or ovate-oblong), apex obtuse to acute with a short (0.3 mm) tip, base acute to obtuse (rarely round- ed), margin entire or with a few retrorse aculeolate tri- chomes, midvein prominent, secondary and lateral veins obscure or weakly developed. Inflorescences often of 3 terminal flowers subtended by a whorl of 4 reduced leaves, or of several flowers in compound dichasia, pedicels 1- 5 mm long, filiform. Flowers ca. 1.5 mm long, greenish yellow, said to be <J, 2, and bisexual on the same plant or on different plants, hypanthium 0.5-0.7 mm long, calyx ca. 0.3 mm long; corolla rotate, lobes 0.5-1 mm long, white to yellowish or red. Fruits 3-4 mm long, 4- 7 mm broad when dry, 2-lobed distally, orange becoming black, glabrous and wrinkled when dried, borne on slen- der pedicels 2-5 mm long.
Plants of evergreen montane forest formations, from 1 200 to 2900(-3400) m in Central America. Probably flowering and fruiting throughout the
year. The species ranges from the states of Jalisco and San Luis Potosi in Mexico to western Panama. Galium aschenbornii is recognized by the usu- ally narrow leaves in whorls of four, short petioles, and glabrous fruit. It is infrequently collected in Costa Rica.
Galium hypocarpium (L.) Clos in Gay, Fl. Chil. 3: 186. 1847. Vaillantia hypocarpia L., Syst. Nat. ed. 10: 1307. 17 59. Relbunium hypocarpium (L.) Hemsl., Biol. cent. amer. Bot. 2: 63. 1881. G. hypocarpium (L.) Fosberg, Sida 2: 386. 1966. Figure 3.
Herbs to 60 cm high, prostrate to procumbent or climbing over low objects, leafy stems 0.4-1 .3 mm thick, with 4 prominent longitudinal ridges, nodes usually well spaced, puberulent with thin whitish hairs 0.3-1 mm long; stipules leaf-like. Leaves 4/node, sessile; leaf blades 3-1 3(-22) mm long, 2-4(-8) mm broad, oblong to ovate- oblong or broadly oblong-obovate, apex obtuse to rounded, with a minutely apiculate tip, base obtuse, dry- ing chartaceous to subcoriaceous, both surfaces with stiff ascending hairs to 0.7 mm long or the surfaces glabrous and the margins hirsutulous. 2° veins 2-3/side (or the venation obscure). Inflorescences of solitary flowers in the leaf axils, usually with 2 flowers per node, peduncles 5-15 mm long, with an involucre of 4 bracteoles sub- tending each flower, bracteoles subequal, 2-4 mm long,
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ovate to oblong or lanceolate, hirsutulous. Flowers small, hypanthium rounded, calyx tube and teeth undeveloped; corolla campanulate, white, 1.5-2.5 mm long, tube cy- lindrical, lobes 4, shorter than the bracteoles and alter- nating with them, surfaces glabrous and with marginal hairs; stamens 4, filaments short, anthers small. Fruits 2-3 mm long, to 3.5 mm broad, broadly 2- or 3-lobed, or with only 1 seed and globose, orange or reddish or- ange, glabrous to puberulent.
Plants of moist evergreen montane forest for- mations and high elevation paramos, from 1800 to 3400 m elevation. Probably flowering and fruit- ing throughout the year. The species ranges from Veracruz, Mexico, southward through the higher elevations of Central America into the Andes mountains of South America as far south as north- ern Chile and Argentina.
Galium hypocarpium is recognized by its slender stems with four leaves at each node, creeping or clambering habit, the solitary axillary flowers sub- tended by an involucre of four leaf-like bracteoles. These plants closely resemble our other species of Galium, but the other species rarely have single flowers subtended by a four-parted involucre.
Galium mexicanum H.B.K., Nov. gen. sp. 3: 337 (quarto). 1818. Figure 3.
Trailing or climbing herbs to l(-2.4?) m long, leafy internodes 0.3-2.8 mm thick, with minute (0. 1-0.3 mm) retrorse aculeolate hairs, often with dense longer (0.5 mm) whitish hairs at the node; stipules leaf-like. Leaves usually 6 or 8(-12)/node, essentially sessile; leaf blades (4-)8-20(-25) mm long, 1-3 mm broad, narrowly ob- lanceolate to narrowly oblong, apex obtuse (rounded), usually with an apiculate tip ca. 0.5 mm long, base grad- ually narrowed, with curved aculeolate retrorse hairs (ca. 0.2 mm long) along the margin and midvein beneath, mostly glabrous above and on the flat surfaces beneath, primary vein prominent, 2 weakly denned lateral veins often present. Inflorescences usually terminal, paniculate arrangements of small cymose groups of 3-7 flowers sub- tended by reduced leaf-like bracts, with divaricate branching and slender pedicels 1-3 mm long. Flowers 1.5-2 mm long, bisexual, hypanthium 0.5-0.7 mm long, turbinate, densely hirsutulous with minute (0. 1 mm) as- cending hairs, calyx reduced; corolla 1-1 .5 m long, white to pink or red, campanulate or rotate, glabrous exter- nally, puberulent within. Fruits dry, ca. 3 mm broad, covered with ascending slightly curved hairs or with a few uncinate hairs at the tip (in Costa Rica).
Plants of evergreen montane forest formations, from ( 1 200-) 1 600 to 3 1 00 m elevation. Flowering in April-July and December-January in southern Central America. Rarely collected in Costa Rica but apparently common in the Chiriqui High-
lands. The species ranges from the southwestern United States to Panama.
Galium mexicanum is recognized by the slender clambering stems with whorls of six or eight nar- rowly oblong or oblanceolate leaves and the small fruit with dense pubescence of curved hairs. The aculeolate trichomes on stems and leaves help the plants to climb and makes them adhesive and very difficult to disentangle from shrubbery or clothing.
Galium orizabense Hemsley, Diagn. PI. Nov. Mexic. 3: 54. 1878. Figure 3.
Erect or spreading herbs, 20-75 cm long, with several to many stems from a small root stock, leafy stems 0.3- 1 mm thick, with thin white curves hairs 0.1-0.3 mm long; stipules leaf-like. Leaves 4/node, subsessile or short- petiolate, petioles to 2 mm long; leaf blades 6-18(-25) mm long, 2-4(-5) mm broad, narrowly oblong to nar- rowly elliptic-oblong or lanceolate, apex obtuse or short- apiculate, drying thin-chartaceous or membranaceous, with thin hairs ca. 0.3 mm long, on the upper surface, margin and major veins beneath, with a prominent 1° vein and 2 straight lateral veins (usually readily appar- ent). Inflorescences paniculate, with divaricate branches subtended by slightly reduced leaves, often with 5-15 flowers on open lateral branches, pedicels 1-10 mm long. Flowers 1-1.5 mm long, hypanthium ca. 0.5 mm long, covered with minute hairs; corolla rotate, lobes ca. 0.4 mm long and equally broad, white or greenish yellow. Fruits dry, 1-2 mm broad, covered with minute uncinate hairs, borne on thin (to 0.5 mm) pedicels.
Plants of evergreen montane forest formations, from 1 500 to 2500 m in southern Central Amer- ica. Flowering in March, July-August, and De- cember-January in southern Central America. The species ranges from eastern and central Mexico to Panama.
Galium orizabense is recognized by the uncinate hairs on the fruit and the narrow verticillate leaves. This species is very similar to G. uncinulatum and the two may be conspecific. However, most spec- imens can be differentiated by the key, and we follow Dempster's treatment.
Galium uncinulatum DC., Prodr. 4: 600. 1830. Figure 3.
Prostrate or procumbent herbs 1 5-90 cm long, leafy stems 0.3-1 .5 mm thick, with thin whitish hairs 0.3-0.6 mm long, slightly scabrous; stipules leaf-like. Leaves 4/ node, sessile or subsessile with petioles to 1 (-2) mm long; leaf blades (4-)6-12(-22) mm long, (2-)2.5-6(-13) mm broad, ovate to narrowly ovate or narrowly oblong, apex obtuse (rounded) with a short (0.5 mm) tip, base obtuse,
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141
drying membranaceous or chartaceous, margin and sur- faces with thin ascending or spreading hairs 0.2-0.4 mm long, with 1° vein and 2 well-defined lateral veins, the lateral veins 0.5-1 mm from the margin in larger leaves. Inflorescences terminal or axillary, with opposite or cy- mose branching, to 3 cm long, flowers usually few (3-5) on the slender peduncles, often subtended by whorls of reduced leaves, pedicels 1-10 mm long, 0.1-0.2 mm thick when dried. Flowers 1-1.5 mm long, hypanthium/ ovary ca. 0.5 mm long, covered by minute hairs that will expand in fruit, calyx reduced; corolla campanulate to rotate, white to greenish or yellow, usually puberulent externally. Fruits ca. 1.5 mm long and (l-)2-3 mm broad, usually 2-lobed and rounded, dry and covered by pale yellowish or whitish hooked (uncinate) hairs ca. 0.3-0.4 mm long.
Plants of evergreen montane forest formations from 1000 to 2800(-3300) m elevation. Flowering collections have been made in all months of the year except May and October-November in Cen- tral America. The species has been little collected in Costa Rica. The species ranges from southern Arizona and Texas (U.S.A.) through Mexico and highland Central America to Panama.
Galium uncinulatum is recognized by having four, often broad, little leaves at each node, small few-flowered inflorescences with thin peduncles and pedicels, and fruits densely covered with thin uncinate hairs. This species is very similar to G. orizabense(q.v.), which appears to live in the same habitats but tends to have narrower leaves and shorter puberulence. Specimens referred to as Ga- lium obovatum H.B.K. by Standley, both in the herbarium and in his flora (1938), are G. uncin- ulatum.
Gardenia Linnaeus
Trees or shrubs, branches terete, glabrous or puber- ulent; stipules interpetiolar and intrapetiolar, triangular, apex acute to acuminate, often forming a short sheath at the base. Leaves opposite or in whorls of 3, subsessile to short-petiolate, coriaceous to chartaceous, entire, pin- nately veined, domatia often present. Inflorescences ter- minal or axillary, of 1 or 3 flowers (rarely more and corymbose), sessile or short-pedicellate. Flowers radially symmetrical, bisexual, usually large, hypanthium ovoid to ellipsoid or obconic, calyx tube short (sometimes spathe-like), calyx lobes 5-8 when present; corolla sal- verform to campanulate or funnelform, white or yellow, corolla tube glabrous or puberulent, corolla lobes 5-11, convolute in bud, spreading or recurved; stamens 5-9, inserted on the upper half of the tube, filaments short or absent, anthers dorsifixed, linear to linear-oblong, in- cluded or partly exserted, disc annular to crenate; ovary 1 -locular (rarely 2- or 6-locular at apex), ovules many and horizontal on parietal placentas, style linear and
terete, stigma linear to clavate, 1 - or 2-lobed. Fruits ob- long to ovoid, pyriform or globose, terete or costate, the outer wall fleshy to leathery or woody, rupturing irregu- larly or the endocarp breaking into 2-5 valves; seeds very many, imbedded in a fleshy pulp, horizontal, an- gulate, embryo small.
A genus of about 200 species in the tropics and subtropics of the Old World. A few species are important as ornamental trees in warm climates; they are also grown under glass in cold climates for their large aromatic flowers, which are often used for corsages. The genus is similar to Genipa. The genus has not become naturalized in Central America, where one species is commonly seen in gardens.
Gardenia augusta (L.) Merr., Interpr. Herb. Am- boin. 485. 1917. Varneria augusta L., Amoen, Acad. 4: 136. 1759. G. jasminoides Ellis, Phil. Trans. 51, pt. 2: 935. 1761. G.florida L., Sp. PI. ed. 2: 305. 1762.
Shrubs or many-branched small trees to 5 m tall, leafy branchlets 1.5-5 mm thick, sparsely and minutely (0.2 mm) puberulent, glabrescent; stipules 5-10 mm long, at first enclosing the apex and splitting down one side to become spathe-like, with a short tube 2-4 mm long at the base, persisting. Leaves 2(-3)/node, petioles 1—4 mm long and little differentiated from the leaf base; leaf blades 3-12 cm long, 1.5-5 cm broad, elliptic-obovate to ellip- tic-oblong or broadly elliptic (in smaller leaves), apex acuminate, base acute and decurrent on petiole, glabrous above and below (but sometimes with pit domatia and a few hairs in the leaf axils), 2° veins 6-9 /side. Inflores- cences often of 3 terminal flowers, or solitary flowers in the axils of near-terminal leaves, pedicels ca. 1 0 mm long. Flowers large (6-10 cm long), apparently differing in size in different cultivars, sweetly aromatic, hypan- thium ca. 10 mm long, calyx lobes 8-30 mm long, nar- rowly oblong, spur-like; corolla white, tube 2-5 cm long, corolla lobes usually 6 (some cultivars with 2 series), ca. 25 mm long and 18 mm broad, obovate; anthers ca. 18 mm long.
Plants of parks and gardens cultivated for their large white sweetly aromatic flowers. These plants, native to Asia, are called jazmin, jazmin delcabo, and "gardenia."
Genipa Linnaeus
Trees, branchlets usually thick, terete, glabrous or pu- berulent; stipules interpetiolar and intrapetiolar, connate to form a short tube, caducous or deciduous with the leaves. Leaves opposite and decussate, subsessile or pet-
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iolate, often large, pinnately veined, chartaceous to co- riaceous, without domatia. Inflorescences terminal or axillary, flowers solitary or in few-flowered cymes or subcapitate, pedicels present and continuous with the hypanthium base. Flowers radially symmetrical, bisex- ual (rarely unisexual and dioecious), 5- or 6-parted, hy- panthium turbinate to campanulate, calyx tube truncat- ed and entire or with 5-6 short lobes; corolla salverform to funnelform, carnose, white to yellowish white, tube short to long, glabrous or puberulent externally, barbate in the throat within and at the base of the lobes, corolla lobes 5-6, convolute in bud, spreading; stamens 5-6, inserted in the upper part of the tube, anthers subsessile, dorsifixed, linear, partly exserted; ovary 1-locular or be- coming 2-locular, style thick, stigmas fusiform, placen- tation parietal, ovules many and horizontal in vertical files. Fruits baccate, large, ovoid to subglobose or ob- ovoid, calyx tube persistent at apex of the fruit, pericarp thick, fleshy to coriaceous; seeds many, large, com-
pressed (with two parallel flattened sides), the testa slight- ly fibrous.
A genus of 5-10 species, ranging from southern Florida (U.S.A.) and Mexico through Central America into tropical South America. The genus is distinguished by its few-flowered terminal and subterminal inflorescences, large flowers with thick corolla lobes, parietal placentation, and many large horizontal seeds in vertical files within the large, often solitary fruit. Cenipa vulcanicola Stand!, of Mexico and Guatemala has been transferred to Glossostipula concinna (Standl.) Lorence; it has axile placentation.
Key to the Species of Genipa
la. Corolla densely sericeous distally, peduncles to 25 mm long; stipules acute at apex, persisting with the leaves; leaves glabrous or pubescent, with 7-18 major secondary veins on each side; widespread G. americana
Ib. Corolla glabrous on the outer surfaces, peduncles to 10 mm long; stipules broadly ellipsoid and rounded at apex, usually caducous; leaves glabrous above and with appressed hairs on the veins beneath, with 6—9 major secondary veins on each side; not recorded north of southern Costa Rica
. G. williamsii
Genipa americana L., Syst. Nat. ed. 10. 2: 931. 1759. G. oblongifolia Ruiz & Pav., Fl. Peruv. Chil. 2: 67, pi. 220. 1798. G. caruto H.B.K., Nov. gen. sp. 3: 407 (quarto). 1820. G, ameri- cana var. caruto (H.B.K.) Schum. in Mart., Fl. Bras. 6(6): 352. 1889. G. codonocalyx Standl., Contr. U.S. Natl. Herb. 1 7: 446. 1 9 1 4. G. venosa Standl., J. Wash. Acad. Sci. 18: 168. 1928. Fig- ure 26.
Small to large trees 4-27 m tall, often with a spreading hemispheric crown, trunk to 50 cm thick, bark smooth and lenticellate, leafy branchlets 4-9 mm thick, densely pubescent in early stages or glabrous; stipules 10-25 mm long, triangular, the basal sheathing tube 1-3 mm long, acute, deciduous with the leaves. Leaves with petioles 2-13 mm long, 2-3 mm thick, glabrous or pubescent; leaf blades 1 2-42 cm long, (4-)6-19 cm broad, obovate to elliptic-obovate or broadly oblanceolate, apex acu- minate to obtuse or rounded, gradually narrowed to a cuneate or slightly decurrent base, drying chartaceous and often very dark above, glabrous and lustrous above, glabrous to densely pubescent beneath with thin soft hairs ca. 0.5 mm long, 2° veins 9-1 8/side. Inflorescences terminal or subterminal, 4-10 cm long and with 1-9 flowers, cymose, peduncles to 25 mm long, glabrous, pedicels 4-12 mm long. Flowers 2.5-4 cm long, appar-
ently bisexual but perhaps functionally unisexual, hy- panthium difficult to distinguish from the calyx tube and together 7-17 mm long, calyx tube to 10 mm diam. distally, truncate or with broad short lobes, glabrous on the exterior and puberulent within; corolla ca. 4 cm broad, salverform, carnose, densely descending-sericeous ex- ternally (except at the base of the tube), white or yellow- ish white, darkening with age, tube 5-15 mm long, 4-7 mm diam. (to 10 mm at the lobes), lobes 5-6, 11-28 mm long, 5-12 mm broad, obovate and rounded at apex; anthers 6-14 mm long, becoming recurved between the lobes; stigmas ca. 5 mm long. Fruits 4-1 1 cm long, 3- 1 1 cm diam., obovoid to subglobose, smooth and grayish brown or yellowish brown, the persisting calyx 3-6 mm long, crateriform on apex of the fruit and 8-10 mm diam., pedicels up to 5 cm long in fruit; seeds 6-12 mm long, 4-7 mm broad, ca. 2.3 mm thick.
Trees of both wet evergreen rain forests and seasonally very dry deciduous forest formations in the Caribbean and Pacific lowlands, from near sea level to 900 m elevation. Flowering occurs primarily in March-August; fruiting throughout the year. This species ranges from southern Flor- ida and the West Indies, through Mexico and Cen- tral America through tropical South America to Paraguay.
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Genipa atnericana is recognized by the larger leaves with short petioles clustered at the ends of stems, usually solitary large fruit with many hor- izontal seeds in vertical files, and large sericeous flowers with short corolla tubes and large lobes. This is a common and distinctive tree, especially conspicuous in deciduous forest formations in the dry season because of its large terminal fruit (but compare Alibertia edulis). This species may be confused with species of Borojoa, but those tend to have sessile terminal flowers and stipules with parallel venation. Guaitil, caruto, jagua, andjagua negro are common names for this species. The juice of the young pulpy fruit turns black or dark blue and is used by Native Americans as a dye or body paint. The species is sometimes cultivated, and the fruit is eaten. The wood is easy to work but strong and resistant; it is used for making fur- niture and carts and in building construction.
Genipa americana is here interpreted to be a very variable species, following Dwyer (1980) and Steyermark (1974). The types of Standley's G. co- donocalyx(Pittier 12085 us) and G. venosa (Stand- ley & Valeria 45269 us) appear to represent no more than unusual forms of G. americana and were described when the full pattern of variation in G. ameriana was not apparent. The type of G. venosa has prominent petioles (3.5—4 cm long) and unusually long fruit, but it seems better to treat it as a variant of G. americana rather than as a dis- tinct species. Collections with the leaves densely pilose beneath have been referred to variety caruto (H.B.K.) K. Schum.
Genipa williamsii Standl., J. Wash. Acad. Sci. 8: 642. 1918. Figure 26.
Small to medium-sized trees, 4-20 m tall, with boles ca. 25 cm dbh, leafy stems 3-7 mm thick, glabrescent, becoming pale brown; stipules 10-32 mm long, 6-18 mm broad, ovate from a narrowed base, flattened, ap- pressed-sericeous. Leaves with petioles 7-22(-60) mm long, 1 .5-2.7(-4) mm thick, glabrous, often drying black- ish and lustrous; leaf blades 8-17(-33)cm long, 5-10(-16) cm broad, obovate-oblong, to broadly elliptic or elliptic- oblong, apex rounded and lacking a narrowed tip, base obtuse to acute and slightly decurrent on petiole, drying stiffly chartaceous to subcoriaceous and dark reddish brown above, glabrous above, appressed-sericeous on the major veins beneath with hairs ca. 0.3 mm long, 2° veins 7-9/side and weakly loop-connected distally, 3° venation obscure. Inflorescences of ca. 3 (4-7) terminal flowers subtended by 2 ovate-lanceolate bracts (stipules) ca. 1 2 mm long with glabrous surfaces but ciliolate along the edge, peduncles 5-10 mm long, pedicels 8-10 mm long and continuous with the hypanthium, drying black,
bracteoles 3 mm long or reduced to ridges. Flowers gla- brous externally, drying black, hypanthium 3-4 mm long, 4-5 mm broad at apex, obconic, calyx tube 0.5-1 mm high, entire or slightly undulate; corolla salverform, white and carnose, corolla tube 1.8-3 cm long, 4-5 mm diam., lobes 5, 18 mm long, 8-12 mm broad distally, obovate- oblong and rounded distally; anthers sessile, stigmas 4, to 4 mm long, unequal. Fruits subglobose, ca. 7 cm diam., drying black; seeds ca. 10 mm long, 5-6 mm broad, imbedded in white pulp.
This species has been collected only in south- ernmost Limon Province at 450-650 m elevation (Hammel et al. 1 7597 CR, MO, Herrera 3208 CR.MO) in Costa Rica. Flowering in July; fruiting in April- June and October in Panama. The species ranges to Colombia.
Genipa williamsii is recognized by the leaves rounded distally, clavate flower buds, and both the fleshy flowers and the large fruits that dry black. It is similar to species of Ladenbergia, but those have domatia and valvate corolla lobes.
Geophila D. Don
REFERENCE— L. O. Williams, Geophila (Rubi- aceae) in North America. Phytologia 26: 263-264. 1973.
Creeping perennial herbs, stems slender and puberu- lent or glabrous, rooting at the nodes; stipules interpetio- lar, small, rounded-ovate to triangular, entire to shal- lowly bilobed, persisting. Leaves usually with long petioles; leaf blades rounded and often cordate at the base, membranaceous to thin-chartaceous, venation pin- nate, domatia absent. Inflorescences terminal or pseu- doaxillary, few-flowered heads or cymes, peduncles short or long, flowers subtended by an involucre of small bracts, pedicels short or absent. Flowers radially symmetrical and bisexual, usually 5-parted (less often 4-7-parted), calyx tube with 4-7 narrow lobes, persistent; corolla fun- nelform to salverform, white, corolla tube narrow, pilose in the throat, corolla lobes 4-7, valvate in bud, spreading or recurved; stamens 4-7, filaments filiform and inserted in the floral tube, anthers dorsifixed, linear, half exserted; ovary 2-locular, ovules solitary in each locule and basal, style slender with 2 stigmas. Fruits a juicy berry, usually containing 2 1 -seeded pyrenes (nutlets); pyrenes plano- convex and smooth or costate on the dorsal surface, with a ventral sulcus.
A genus of 20-30 species native to the American tropics, Africa, and Asia. The slender creeping stems, long-petiolate leaves with rounded blades, few-flowered inflorescences, and fleshy, two-seed- ed fruits characterize this genus. These plants may be mistaken for species ofCoccocypselum, but that genus has many-seeded fruit.
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Key to the Species of Geophila
la. Fruit black at maturity, pyrenes weakly costate; peduncles 2-10 cm long; basal lobes of the leaf
blades separated by a sinus (cordate with non-overlapping lobes) G. macropoda
Ib. Fruit red at maturity (if fruit are blue go to the genus Coccocypselum), pyrenes strongly costate;
peduncles 0.2-10 cm long; basal lobes of the leaf blades separate to overlapping 2
2a. Ovary, fruit and leaves conspicuously pilose with thin hairs 0.5-2 mm long [leaf blades subcordate
with a small basal sinus; peduncles to 7 cm long in fruit] G. cordifolia
2b. Ovary and fruit glabrous, leaves glabrous or puberulent with short (0.1-0.3 mm) hairs; peduncles
to ca. 2 cm long 3a
3a. Leaf blades ovate and slightly longer than broad, often narrowed at apex, cordate but usually without
a visible basal sinus (because the lobes overlap slightly) G. repens
3b. Leaf blades ovate-triangular and distinctly longer than broad, usually acute at apex, cordate to
subcordate at the base and with a small sinus G. gracilis
Geophila cordifolia Miq., Stirp. Surin. Sel. 176. 1 850. Mapouria trichogyne Muell.-Arg. in Mart., Fl. Bras. 6(5): 426. 1881. Geophila trichogyne (Muell.-Arg.) Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 423. 1931. Figure 2.
Creeping herbs, leafy stems 0.7-1 .5 mm thick, densely pubescent with slender pale straight or crooked hairs 0. 5- 2 mm long; stipules 2-4(-6) mm long, 1.5-4 mm broad (broadest beneath the inflorescences), usually glabrous, persisting. Leaves with petioles 3-7(-l 3) cm long (short- er on leaves subtending the inflorescences), conspicu- ously pubescent with slender crooked or straight mul- licellular hairs 0.7-1.5 mm long; leaf blades 3-7(-ll) cm long, 2-6(-8.5) cm broad, broadly ovate to oblong- ovate or narrowly ovate (triangular-ovate), apex obtuse to short-acuminate, base cordate with rounded lobes 8- 35 mm broad, basal sinus 3-15 mm deep, drying thin- chartaceous and brownish, both surfaces covered with thin usually crooked hairs 0.7-2 mm long, 2° veins 3- 5/side, not usually loop-connected near the margin. In- florescences terminal, capitate, 1-3 cm long (to 6 cm in fruit), with 5-1 7 flowers, peduncles 5-1 5 mm long, elon- gating in fruit, densely pubescent, bracts 3-10 mm long, lanceolate and pubescent, pedicels 0.5-3 mm long. Flow- ers with an urceolate hypanthium, calyx tube 0.5 mm long and 1 .5 mm diam.. lobes ca. 3 mm long, with slen- der hairs ca. 1 mm long; corolla 5-6 mm long, white often tinged with pink distally, tube 2-4.5 mm long, glabrous externally, with a short collar of hairs at the point of filament attachment within, lobes 5, 1 .5-2.5 mm long; stamens with anthers 0.8-1 mm long, included. Fruits ca. 8 mm long, ovoid-globose, orange to red, with scattered slender hairs, pyrenes ca. 4 mm long and 3 mm broad, with 3-5 prominent longitudinal dorsal ribs.
Plants of evergreen forest formations, from near sea level to 500(-1100) m elevation. Probably flowering and fruiting primarily in the wet season (May-December). The species ranges from Belize along the Atlantic slope of Central America to Colombia, Venezuela, and the Amazon basin in Brazil and Peru.
Geophila cordifolia is recognized by the long, often crooked, thin multicellular hairs that cover almost all parts of the plants. The long-petiolate leaves with cordate bases, short inflorescences elongating in fruit, and glabrous stipules are ad- dition distinctions. In Costa Rica the species, is known only from the La Selva area and from Vol- can Rincon de la Vieja.
Geophila gracilis (Ruiz & Pav.) DC, Prodr. 4: 537. 1 830. Psychotria gracilis Ruiz & Pav., Fl. Peruv. 2: 63, pi. 211, f. C. 1799. G. croatii Steyerm., Phytologia35:401. 1977.
Creeping herbs to 20 cm tall or slender-stemmed vines to 1.5 m long, leafy stems 0.3-1.3 mm thick, glabrous, horizontal internodes 3-7 cm long, with adventious roots near the nodes; stipules 2-4 mm long. 1-2 mm broad, usually rounded at apex, curving outward in age and persisting. Leaves with petioles 2-8 cm long (but shorter below the inflorescences), 0.3-1 mm thick, glabrous abaxially and with 2 rows of short (0.2-0.5 mm) stiff retrorse or erect hairs along either side of the adaxial sulcus; leaf blades 1.8-5 cm long, 1-3.5 cm broad, tri- angular-ovate to ovate, apex gradually narrowed and acute (or obtuse), base cordate to subcordate, sinus 3-8 mm deep, the basal lobes usually separate but occasion- ally overlapping and the sinus obscured, drying thin- chartaceous, glabrous above or with few thin hairs 0.3- 0.8 mm long, usually glabrous beneath, 2° veins 3-47 side, usually loop-connected near the margin. Inflores- cences terminal, 12-15 mm long, capitate with ca. 3-7 flowers, peduncles 2-8 mm long (apparently longer when the leaves of the subtending node are reduced and bract- like, bracts 4-7 mm long, 1-1.5 mm broad, united at the base and persisting, pedicels 0-1 mm long. Flowers with hypanthium ca. 2 mm long, calyx lobes ca. 3 mm long and 0.5 mm broad, narrowly oblong, persistent and enlarging in fruit; corolla white. Fruits red at maturity, 3-5 mm long, subglobose. pyrenes ca. 3.5 mm long, with 3 prominent longitudinal dorsal ridges.
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Plants of evergreen forest formations, from near sea level to ca. 500 m elevation. Flowering in May- December (primarily in June and July in central Panama; Croat, 1978). The species is known from southeastern Nicaragua, the Canal area of Pana- ma, and the upper Amazon basin of Brazil, Peru, and Bolivia.
Geophila gracilis is recognized by its slender stems rooting at most nodes, triangular-ovate leaf blades with little or no puberulence, and short terminal capitate inflorescences subtended by bracts fused at the base. Variation in specimens from South America clearly encompass the dis- tinctions used to separate G. croatii. This species is apparently common on Barro Colorado Island, Panama, and has been collected in Nicaragua, but it has yet to be collected in Costa Rica.
Geophila macropoda (Ruiz & Pav.) DC., Prodr. 4: 537. 1830. Psychotria macropoda Ruiz & Pav., Fl. Peruv. 2: 63, pi. 21 1, f.6. 1799. Figure 2.
Creeping herbs, leafy stems 0.8-2 mm thick, glabrous or very minutely (0.1 mm) puberulent, often with 2 prominent longitudinal ridges; stipules 2-6 mm long, ovate-oblong, glabrous, deciduous or obscured by the adventitous roots. Leaves with petioles (2-) 3-1 4 cm long (shorter below the inflorescences), 0.7-1.2 mm thick, glabrous abaxially but with 2 adaxial ridges with short (0.1-0.4 mm) dense hairs; leaf blades 3-9 cm long, 2.5- 8 cm broad, broadly ovate to ovate-orbicular, apex rounded to broadly obtuse, base cordate with lobes 1-4 cm broad, basal sinus 2-15 mm deep, drying membra- naceous or thin-chartaceous and often grayish green, gla- brous above, glabrous beneath except for the minute puberulence on the major veins near the base, 2° veins 3-5/side and weakly loop-connected near the margin. Inflorescences usually axillary, 2-4 cm long and elon- gating in fruit, capitate with 3-7 flowers, peduncle 1.5- 5(-7) cm long, minutely puberulent with whitish hairs ca. 0. 1 mm long, bracts 3-6 mm long, united at the base, pedicels to 2 mm long. Flowers with hypanthium ca. 1.5 mm long, essentially glabrous, calyx lobes 2-3 mm long; corolla 3-7 mm long, tube 3-4 mm long, corolla lobes 5, 2-3 mm long. Fruits black or blue, sessile, 5-10 mm long, 3-7 mm diam.. ellipsoid to ovoid; pyrenes 4-7 mm long, 2-3 mm broad, without raised longitudinal ribs (costae) on the convex surface.
Plants of the lowland Caribbean rain forest for- mations, from near sea level to 600 m. Flowering in April-November. The species ranges from southern Mexico through Central America to Bo- livia and Paraguay.
Geophila macropoda is recognized by the axil- lary and long-pedunculate inflorescences, closely clustered flowers, black fruit, and pyrenes without prominent longitudinal costae.
Geophila repens (L.) I. M. Johnston, Sargentia 8: 281. 1949. Rondeletia repens L., Syst. ed. 10: 928. 1759. Psychotria herbaceajacq., Enum. PI. Carib. 16. 1760. Geophila herbacea (Jacq.) Schumann in Engl. & Prantl., Nat. Pflanzenfam. 4,4: 119. 1891. Figure 2.
Creeping herbs to ca. 10 cm high, leafy stems 0.5-1 mm thick, glabrous or very sparsely and minutely pu- berulent; stipules 0.5-2 mm long, 1-2 mm broad, broad- ly ovate, glabrous, persisting or deciduous. Leaves with petioles l-6(-8.5) cm long (sometimes shorter in leaves subtending the inflorescences), 0.4-1.3 mm thick, gla- brous on the abaxial surface but with short (0.2-0.5 mm) retrorse or crooked hairs along the adaxial (upper) side; leaf blades 1.2-5.5 cm long, 1-5 cm broad, broadly ovate to ovate-suborbicular, apex bluntly obtuse or rounded- obtuse, base cordate with lobes 5-25 mm broad, basal sinus 1-7 mm deep and usually obscured by the over- lapping basal lobes, drying membranaceous, upper sur- face glabrous or with a few short hairs near the margins, glabrous beneath or with a few short (0. 1-0.3 mm) hairs on the veins near the petiole, 2° veins 3-5/side, cystoliths visible (as short whitish lines) or obscure on the lower surface. Inflorescences solitary and terminal on short leafy shoots (apparently 2-3 and axillary when directly subtended by small leaves), with (l-)2-5 flowers, pe- duncles 5-35 mm long, puberulent with short retrorse hairs, subtending bracts 3-6 mm long, united at the base and lanceolate distally, pedicels 0-2 mm long. Flowers white or becoming pink in age, hypanthium ca. 1 mm long, calyx tube 1-1.5 mm long, glabrous, calyx lobes 1-3 mm long, 0.3-0.7 mm broad, glabrous; corolla 8- 14 mm long, glabrous or puberulent, funnelform, tube 6-9 mm long and 1-1.5 mm diam., lobes 5, 3-5 mm long, 1.5-2.5 mm broad, ovate and obtuse to acute; sta- mens with filaments ca. 0.5 mm long, attached near the middle of the tube, anthers ca. 2 mm long; style 5-7 mm long. Fruits 8-10 mm long, ovoid or globose, bright red at maturity, subsessile or short (1-2.5 mm) pedicellate; pyrenes 3.5-5 mm long, 2.5-3 mm broad, with 3 slightly raised longitudinal ribs (costae) on the convex face.
Plants of the shaded forest floor in evergreen or partly deciduous forest formations of both the Ca- ribbean and Pacific lowlands of Costa Rica, from near sea level to 800 m elevation. Flowering in June-October; fruiting in July-November. The species ranges from Mexico and the West Indies to Peru and Bolivia in the New World; it is also found in West Africa, the Philippines, and the western Pacific.
Geophila repens is recognized by its creeping habit and short stature, small cordate leaves with the sinus obscured, petioles with puberulence along one side, bright red fruit, and pyrenes with weakly developed costae. This is our most commonly en- countered species of Geophila; it has been col- lected at La Selva and from near Canas in Guana- caste Province to the Golfo Dulce area along the
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Pacific. Lechuga is a name used for this species in the Golfo Dulce region. Breeding biology was stud- ied by Bawa and Beach (1983).
Gonzalagunia Ruiz Lopez & Pavon
Shrubs or small trees, distal branches often curved, pendant or scandent, leafy stems usually slender and pubescent, terete, nodes usually thickened; stipules in- terpetiolar, usually broad at the base and triangular with a narrow distal awn (rarely intrapetiolar and tubular). Leaves distichous, petiolate or subsessile; leaf blades of- ten thin-chartaceous, usually narrow and with ascending secondary veins, domatia absent or obscure. Inflores- cences solitary, usually terminal and narrowly long-ra- cemiform, spiciform or thyrsiform, the flowers solitary, cymose or fasciculate on short lateral branches of the central axis, bracteoles present, flowers sessile or pedi- cellate. Flowers radially symmetrical, bisexual, mono- morphic or distylous, small, 4- (less often 5-) parted, hypanthium urceolate to rounded, calyx tube very short, calyx lobes 4 or 5, small, equal or unequal, persisting in fruit; corolla salverform to funnelform, white or pink, corolla tube narrow, villose in the throat, corolla lobes
4(-5), imbricate or valvate; stamens 4(-5), filaments short or absent, anthers dorsifixed, 2-lobed at the base, ex- serted or partly exserted; ovary 2-or 4-locular, ovules numerous, placentation peltate on the septum, style with 2 or 4 stigmatic lobes. Fruits baccate, fleshy or spongy, subglobose, 2- or 4-locular and usually with 2 or 4 lobes or sulci, with 2 or 4 hard cocci; seeds 4-many within the cocci, minute.
A genus of 25-35 species, ranging from Mexico, Central America, and the West Indies through tropical South America. The long narrow spike- like distal inflorescences, slender drooping distal stems, small narrow-tubed flowers and baccate fruits with 2-4 pyrenes make this a very distinctive genus. Some species of Rondeletia with long-nar- row inflorescences may be confused with species of Gonzalagunia in the absence of fruit; Rondeletia has capsular fruit. Several of our species are weedy shrubs of open secondary growth and closely re- lated; they can be difficult to distinguish in the absence of mature flowers or fruit. A few species resemble species of Buddleia (Loganiaceae).
Key to the Species of Gonzalagunia
la. Stipules united or overlapping above the petioles to form a short tubular sheath or broad tube-like
base 4-12 mm long 2
Ib. Stipules not forming a tube above the node, or the broad margins not overlapping at the base, stipule
only 1-3 mm long before being narrowed into the awn-like apex 3
2a. Stipular sheath to 1 cm long; flower clusters and lateral branches of the inflorescences subtended
by caducous bracts 4-12 mm long and 1-2 mm broad, corolla 3—4 mm long; leaf blades with
5-8 pairs of major secondary veins; small treelets of wet forest understory .... G. bracteosa 2b. Stipular sheath 0-5 mm long or the stipule margins slightly overlapping; bracts less than 2 mm
long, corolla 4-6 mm long; with 10-14 pairs of secondary veins; trees to 18 m tall
Rondeletia brenesii
3a. Petioles usually less than 4 mm long; corolla tube usually less than 7 mm long 4
3b. Petioles usually more than 4 mm long: corolla tube usually more than 7 mm long 7
4a. Leaf blades with 4-7 pairs of major secondary veins, laminae thin-textured; flowers thin-textured
and often solitary, corolla glabrous externally G. rudis
4b. Leaf blades with 7-1 3 pairs of major secondary veins; thinly to stiff-chartaceous; flowers thick- textured, corolla densely sericeous externally 5
5a. Leaf blades subsessile, narrowly lanceolate, 9-26 cm long, with 9-15 pairs of secondary veins
arising at angles of about 30—40° [only known from the Golfo Dulce area] G. brenesii
5b. Leaf blades subsessile or short petiolate, ovate to elliptic and rarely lanceolate, to 1 6(-30?) cm
long, with 4-12 pairs of secondary veins arising at angles of 40-60° 5
6a. Corolla lobes ca. 1.5 mm long; leaf blades with 4-1 1 pairs of major secondary veins, ovate to
ovate-elliptic, thinly chartaceous; Caribbean and Pacific slope, 0-1200 m elevation
G. ovatifolia
6b. Corolla lobes 3-4 mm long; leaf blades with 7-12 pairs of major secondary veins, ovate-elliptic
to lanceolate, stiffly chartaceous; wet Caribbean slope 300-1200 m elevation
G. stenostachya
7a. Mature fruit becoming blue-black; corollas white or white tinged with pink, corolla tubes 8-13 mm long; inflorescences with the flower clusters sessile; 0-1200(-1500) m elevation . . G. panamensis
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7b. Mature fruit white; corollas reddish to pink, corolla tubes 6-10 mm long; inflorescences with the flower clusters on short (1-3 mm) peduncles (but note that some inflorescences may only have solitary flowers and no apparent secondary peduncles); 900-2200 m elevation G. rosea
Gonzalagunia bracteosa (J. D. Smith) B. L. Rob- inson, Proc. Amer. Acad. Arts 45: 405. 1910. Gonzalea bracteosa J. D. Smith, Hot. Gaz. 33: 252. 1902. Figure 21.
rescences appear to be restricted to northeastern Costa Rica. This species differs greatly from our other members of the genus; it resembles Psy- chotria pilosa.
Shrubs or small treelets, 1.5-4 m tall, leafy branchlets 2-4(-6) mm thick, terete, with many ascending strigose hairs 1-1.5 mm long; stipules 8-18(-24) mm long, 3-6 mm broad, with a tubular sheath to 12 mm long, dark brown and pubescent along the midvein, acute to acu- minate and with a slender tip 1-7 mm long. Leaves with petioles 2-10 mm long, 1-2 mm thick, broad, pubescent; leaf blades 6-19(-22) cm long, 2.5-7.5(-8.5) cm broad, narrowly elliptic-obovate to obovate-oblong, oblong or elliptic, usually broadest above the middle, apex acu- minate (acute), tip to 1 5 mm long, gradually narrowed to the acute or obtuse base, leaves usually drying char- taceous and dark brown above (rarely subcoriaceous), sparsely pubescent above with thin appressed hairs 0.5- 1 mm long (densely strigulose on the midvein), more densely appressed-pubescent beneath with brownish hairs 0.5-1.3 mm long, 2° veins 5-7 /side. Inflorescences sol- itary, axillary or terminal, 6-25 cm long, spike-like or thyrsiform panicles with small (5-15 mm) alternate or opposite flower clusters 2-14 mm distant along the rachis (rarely with lateral branches to 4 cm long), primary pe- duncles 2-8 cm long, with ascending hairs to 2 mm long, several bracts 5-12 mm long and 1-2 mm broad sub- tending the flower clusters, flowers sessile. Flowers with hypanthium ca. 1 mm long, densely hirtellous, calyx tube minute, calyx lobes ca. 0.5 mm long, difficult to see among the hairs, glabrous on the inner surface; corolla funnelform, greenish white to white, with few straight hairs 0.5 mm long externally, tube 2-3 mm long, corolla lobes 5, 1-2 mm long; stamens 5, anthers ca. 1 mm long; style ca. 2.5 mm long, stigma 0.4 mm long. Fruits 3-5 mm long, 3-5 mm broad, depressed globose, becoming blue or blue-black, pubescent, sessile; pyrenes 2-4.
Plants of lowland rain forest formations on the Caribbean and Pacific slopes in Costa Rica, from 10 to 850 m elevation. Flowering in January-Sep- tember; fruiting throughout the year. The species ranges from northeastern Costa Rica to Colombia.
Gonzalagunia bracteosa is characterized by the hirsutulous pubescence on many parts, obovate- oblanceolate leaves drying dark above, narrow in- florescences with short flower clusters subtended by conspicuous bracts, small sessile flowers, and tubular stipules. Two collections are noteworthy because the inflorescences have lateral branches 2-4 cm long and with many bracts along their length: Folsom 9778 (DUKE, F) and Zamora & San- chez 469 (CR, F). Specimens exhibiting such un- usual variation in bract development and inflo-
Gonzalagunia brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1302. 1938. Figure 20.
Shrubs, 1.5-3 m tall, leaf branchlets 1.5-6(-8) mm thick, rounded-quadrangular in cross-section, sparsely pubescent with appressed hairs ca. 0.5 mm long or gla- brous, drying reddish brown; stipules 4-8 mm long, 2.5- 7 mm broad at the base, with a very short (1-2 mm) broadly triangular base and a slender awn 4-6 mm long. Leaves subsessile, petioles 0-3(-5) mm long; leaf blades 9-26 cm long, 2.7-6 cm broad, narrowly lanceolate to narrowly elliptic-oblong or elliptic-oblanceolate, apex ta- pering gradually and long-acuminate, base obtuse, drying chartaceous and dark brown above, lustrous above in life, glabrous or sparsely pubescent above, with thin whitish ascending hairs beneath, the hairs longer (ca. 0.4 mm) on the major veins beneath, 2° veins 9-1 3/side and weakly loop-connected distally, 3° veins subparallel. In- florescences solitary and terminal, 20-50 cm long, spi- ciform with short (1-5 mm) lateral branches bearing 2- 6 flowers, peduncles 0-6 cm long, 1.5-3 mm thick, with ascending appressed hairs, bracts 1-3 mm long, linear, pedicels 1-2 mm long. Flowers 6-12 mm long, hypan- thium 0.5-1 mm long, sericeous at the base, calyx tube ca. 0.5 mm long, calyx lobes 0.2-0.3 mm long, glabrous; corolla white or pink, tube 7-10 mm long and 0.7-1 mm diam., pubescent, lobes 3 mm long and 1.8 mm broad, obtuse; ovary 4-locular. Fruits 3-5 mm long, 3-6 mm broad, usually 4-lobed, becoming white, minutely and sparsely puberulent.
Plants of rain forest formations in Costa Rica's southern Pacific lowlands, from near sea level to 500 m elevation. Flowering in April-December; fruiting in January and August-September. This species is endemic to Costa Rica, ranging from the forests of the Pacific slope above Quepos eastward to the Osa Peninsula.
Gonzalagunia brenesii is distinguished by its re- stricted range, long narrow subsessile leaves, long spicate inflorescences, and short narrowly tubular white or pink flowers. This species appears to be related to the G. panamensis-G. rosea complex.
Gonzalagunia ovatifolia (J. D. Smith) B. L. Rob- inson, Proc. Amer. Acad. Arts 45: 405. 1910.
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FIELDIANA: BOTANY
Gonzalea ovatifolium J. D. Smith, Bot. Gaz. 27: 336. 1899. Figure 20.
Shrubs to 3(-4) m tall, leafy branchlets 1.3-4 mm thick, with thin ascending brownish hairs ca. 0.5 mm long, terete, glabrescent; stipules 6-10 mm long, subulate with a short (1-2 mm) base and long (4-7 mm) awn-like apex. Leaves subsessile with petioles l-4(-5) mm long, ca. 1.3 mm thick, pubescent; leaf blades 5-12(-16) cm long, 2-6(-7) cm broad, ovate to ovate-elliptic, apex short-or long-acuminate, tip 3-20 mm long, abruptly narrowed to rounded at the obtuse base, unequal at the base, the leaves drying thin-chartaceous and dark above, pale grayish to pale greenish beneath, essentially glabrous above, appressed-pubescent on the veins beneath with thin short (ca. 0.5 mm) hairs, 2° veins 6-1 I/side. Inflo- rescence solitary and terminal, 12-45 cm long, narrowly spike-like with flowers in distant (3-10 mm) clusters, rachis slender 0.5-1 mm thick, with thin whitish as- cending hairs ca. 0.5 mm long, flowers in sessile or sub- sessile groups of 1-3, subtended by linear bracts 2-4 mm long, flowers usually solitary in the distal half of the inflorescence, pedicels to 1 mm long. Flowers with hy- panthium ca. 1 mm long, densely sericeous, calyx lobes 1-2 mm long, glabrous; corolla white, tube 4-5 mm long, lobes 1-2 mm long; anthers ca. 1 mm long. Fruits 2-3 mm long and 3-4 mm broad (dried), white, usually 4-lobed, pubescent with thin hairs ca. 0.3 mm long.
Shrubs of wet evergreen forest formations of both the Caribbean and Pacific slopes in Costa Rica, from near sea level to 1200 m elevation. Flowering in January-April and July-August; fruiting in the same months and in October and December. The species ranges from Nicaragua to Colombia.
Gonzalagunia ovatifolia is recognized by its sub- sessile ovate acuminate leaves, the long inflores- cences with few-flowered cymules or solitary flow- ers along its length, and the small corollas. This species may be difficult to separate from some specimens of G. rosea (q.v.).
Gonzalagunia panamensis (Cav.) K. Schum. In Mart., Fl. Bras. 6(6): 292. 1889. Buena pana- mensis Cav., Anales Hist. Nat. 2: 279. 1800. Gonzalea panamensis (Cav .) Spreng., Syst. Veg. 1:417. 1825. Figure 20.
Shrubs to 3(-5) m tall, erect or scandent, leafy branch- lets 0.8-4 mm thick, terete or slightly quadrangular, sparsely to densely sericeous with thin whitish ascending hairs ca. 0.5 mm long; stipules 4-7 mm long, the broad base 0.5-2 mm long, subulate with a narrow awn-like tip, pubescent on the midrib and edges. Leaves with petioles 6-27 mm long (shorter on young axillary shoots), ca. 1 mm thick, pubescent, with lateral margins contin- uous with the decurrent leaf margin; leaf blades 5-15 cm long, 1-6 cm broad, narrowly lanceolate to lanceo-
late-elliptic or ovate-lanceolate, apex tapering gradually and acute or acuminate, base acute and often decurrent on petiole, drying thin-chartaceous, densely pubescent on the veins and more sparsely between the veins with appressed hairs ca. 0.3 mm long above and below, 2° veins 5-7/side, ascending. Inflorescences solitary and terminal (or axillary by later lateral growth of side shoots), 6-22(-40) cm long, peduncles 1-5 cm long, 1-2 mm thick, pubescent, the flower clusters ca. 5 mm diam., essentially sessile (but solitary flowers pedicellate), bracts to 4 mm long and linear, pedicels 0-2 mm long. Flowers 4-parted, hypanthium 0.7-1 mm long, sparsely pubes- cent, calyx ca. 1 mm long, calyx lobes 0.5-1 mm long and 0.5 mm broad at the base; corolla white or white tinged with pink, tube (8-)10-13 mm long, 0.7-1 mm diam., sparsely pubescent, lobes 2-3 mm long, ca. 2 mm broad at the base, puberulent within; stamens 4, anthers 2-2.2 mm long; style 7-10 mm long, stigma ca. 0.7 mm long. Fruits 2.5-4 mm long, 3-8 mm broad, depressed globose to 4-lobed, glabrous or sparsely puberulent, red becoming purple black or black.
Shrubs of evergreen or partly deciduous forest formations of the Pacific slope in Costa Rica, from near sea level to 1 200 m elevation. Flowering in all months except March-April and November; fruiting in January, February, September-Octo- ber, and December. The species ranges from Mex- ico to Colombia and in the West Indies.
Gonzalagunia panamensis is recognized by the usually lanceolate leaves on slender well-devel- oped petioles, the flower clusters sessile on the inflorescence rachis (or the flowers solitary and pedicellate), the long narrow white corolla tube, and the fruit turning red or black. Collections from higher elevations have broader leaves and may represent introgression from another species; com- pare G. ovatifolia and G. rosea.
Gonzalagunia rosea Standl., Ann. Missouri Bot. Gard. 25: 836. 1938. G. longithyrsa Fosberg, Sida 2: 387. 1966. Figure 20.
Shrubs or small trees, 2-4(-6) m tall, branches erect or scandent, leafy branchlets 1-4 mm thick, terete, with stiff appressed-ascending pale yellowish or grayish hairs 0.2-0.5 mm long; stipules 3-7 mm long, ca. 3-4 mm wide at base, broadly triangular with a short (1-4 mm) narrow tip, pubescent on the edge and midrib. Leaves with petioles 4-20 mm long, ca. 1 mm thick, densely pubescent; leaf blades 7-18(-22) cm long, 2-6(-8) cm broad, narrowly ovate-elliptic to lanceolate, lanceolate- oblong or narrowly elliptic-oblong, apex gradually or abruptly acuminate (acute) with tip 5-1 5 mm long, base acute to obtuse, drying thin-chartaceous to chartaceous, dark grayish brown above, puberulent on the upper sur- face (denser on the major veins) with short (0.2-0.3 mm) straight hairs, more densely puberulent beneath with hairs to 0.5 mm beneath, 2° veins 5-9(-l l)/side, strongly as- cending (and not loop-connected near the margin). In-
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florescences solitary, terminal (axillary by further growth of lateral branches), 12-35 cm long, flowering part ca. 2.5 cm broad, peduncles 5-35 mm long, 1-2.5 mm thick, densely short hirsute, flowers in groups of l-3(-5) and borne on secondary peduncles 1-5 mm long, alternate or opposite on the rachis and 2-10 mm distant, bracts 1-3 mm long and linear, pedicels l-3(-4) mm long. Flowers with hypanthium 0.5-1.5 mm long, ca. 1 mm diam., densely pubescent, calyx tube ca. 0.5 mm long, calyx lobes 4 or 5, 0.5 mm high, triangular; corolla red in early bud, becoming rose red or pink, tube (6-)8-10 mm long, 0.7-1.6 mm diam., slender and widening be- low the lobes, sparsely to densely puberulent, lobes 2-3 mm long, ca. 2 mm wide at the base, glabrous distally on the inner surface and villous near the mouth; stamens 4, anthers 2-2.5 mm long, anthers partly exserted; style 9-1 1 mm long, stigma 0.3-0.7 mm long. Fruits 3-5 mm long, 3-6 mm broad, white, usually 4-lobed, glabrous or sparsely puberulent.
Common shrubby plants of lower montane ev- ergreen forest formations, from (900-)1 100-2200 m elevation. Probably flowering and fruiting throughout the year, but flowering primarily in February-September and fruiting primarily in June-September. The species ranges from the Cor- dillera de Tilaran in Costa Rica to eastern Panama.
Gonzalagunia rosea is recognized by the thin- petiolate narrowly ovate to lanceolate leaves, the long inflorescences with pedunculate flower clus- ters, the pink corollas, and the spongy white fruit. This is a common weedy shrub of open habitats. Some specimens of this species may be difficult to distinguish from G. panamensis and G. ovatifolia, and it is possible that hybridization occurs.
Gonzalagunia rudis (Standl.) Stand 1.. J. Wash. Acad. Sci. 17: 170. 1927. Duggenia rudis Standl., Contr. U.S. Natl. Herb. 18: 125. 1916.
Shrubs, 1.5-4 m tall, leafy stems 0.7-3 mm thick, with minute (0.2 mm) thin appressed-ascending whitish hairs, terete and glabrescent; stipules 4-7 mm long, with a short (1-2 mm) broad base and long narrow awn. Leaves with petioles 1-3 mm long, ca. 1 mm thick, hertellous; leaf blades 2.5-10 cm long, 1-4 cm broad, narrowly ovate to ovate-oblong or lanceolate, apex tapering gradually and acute to long-acuminate, base acute to obtuse, drying thin-chartaceous, with scattered thin appressed hairs ca. 0.5 mm long on both surfaces, with denser pubescence on the veins beneath, 2° veins 4-7/side. Inflorescences terminal or axillary, solitary, 4-14 cm long, spicate, pe- duncles 1-3 cm long, 0.3-0.7 mm thick, densely pubes- cent with ascending hairs, flowers usually solitary or in groups of 2-3 ca. 5 mm diam., subtended by linear bracts 2-3(-5) mm long, pedicels 1-2 mm long. Flowers 5-parted, hypanthium ca. 1 mm long, urceolate, densely puberulent, calyx lobes 1-2 mm long, linear, corolla white, tube ca. 6 mm long and 0.7 mm diam., lobes ca. 5 mm long and 1 mm broad, glabrous externally, puberulent
within; stamens included. Fruits 2-4 mm long, 2.5-5 mm broad, white, depressed globose, 4-lobed, with thin erect hairs 0.5 mm long.
Plants of evergreen forest formation on the Pa- cific slope of southern Costa Rica, from near sea level to 500 m elevation (to 1 000 m in Panama). Flowering primarily in the wet season (June-Sep- tember). This species ranges from about 84°W in Costa Rica to eastern Panama.
Gonzalagunia rudis is recognized by its smaller thin lanceolate leaves, the slender spike-like inflo- rescences with mostly solitary subsessile flowers, and the unusual calyx and corolla with long narrow lobes. Costa Rican collections differ somewhat in their narrower more lanceolate leaves, but collec- tions with such leaves are also found in Panama.
Gonzalagunia stenostachya (Standl.) W. Burger, comb. nov. Rondeletia stenostachya Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1372. 1938. Arachnothryx stenostachya (Standl.) Bor- hidi, Acta Bot. Hung. 33: 303. 1987.
Shrubs to 3 m tall, leafy stems 2.3-6 mm thick, with short dense yellowish or reddish brown hairs ca. 0.5 mm long; stipules 4-15 mm long, triangular, acute, with yel- lowish hairs along the midrib. Leaves with petioles 2-4 mm long, ca. 1.5 mm thick; leaf blades 7-16 cm long, 2-6 cm broad, elliptic, elliptic-oblong to lanceolate or oblanceolate, apex acute or short-acuminate, base grad- ually narrowed and cuneate base (rounded in Dryer 1274), drying stiffly chartaceous and dark reddish brown above (grayish green beneath), with thin hairs 0.4 mm long or glabrescent above, densely puberulent on the midvein beneath, 2° veins 7-12/side, ascending. Inflorescences (7-) 15-25 cm long, 1.3-2.5 cm broad, spiciform thyrsoid panicles, peduncles 1.3-6 cm long, 1.2-2 mm thick, stri- gose, lateral cymules sessile and separate along the rachis, of (l-)2-5 flowers, bracts ca. 3 mm long, linear, pedicels 0-4 mm long. Flowers 4-parted, hypanthium ca. 1.5 mm long and 1.5 mm diam., densely sericeous, calyx lobes 1-2 mm long, narrowly triangular; corolla white, densely sericeous with hairs 0.5-0.9 mm long, tube 4-9 mm long, slender, lobes 3-4.5 mm long, oblong. Fruits ca. 3 mm long and 4 mm broad (?immature) with sericeous hairs ca. 0.5 mm long.
Plants of wet evergreen forest formations of the Caribbean slope, 300-1200 m elevation. Flower- ing in February-March, July, and October; fruit- ing in October. This endemic species is known from near Monteverde and the La Selva-Braulio Carillo area.
Gonzalagunia stenostachya is recognized by its subsessile leaves with many ascending secondary veins, the long slender spikes with sessile and well-
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separated cymules, and the sericeous flowers. The Panamanian G. kallunkii Dwyer (?= G. veraguen- sis Dwyer) has rather similar inflorescences but the petioles of that species are well developed and the leaves have fewer secondary veins.
Guettarda Linnaeus
Trees or shrubs, branchlets terete, puberulent or gla- brous, occasionally with spines; stipules interpetiolar, simple or slightly connate above the petioles (intrapetio- lar), often slightly overlapping above the node, triangular and acuminate to rounded distally, persisting or decid- uous. Leaves opposite (rarely 3 or 4/node), petioles short to long; leaf blades entire, with pinnate venation, the distal secondaries often strongly ascending and the 3° veins often subparallel, domatia sometimes present. In- florescences solitary and axillary (1 or 2/node), usually pedunculate and with cymose branching, branches of the inflorescences often dichotomous (bifurcate) and with sessile flowers along 1 side, bracts and bracteoles present or reduced. Flowers bisexual (rarely unisexual), radially symmetrical, 4-9-parted (usually 5- or 6-parted), hy- panthium ovoid to globose or tubular, calyx tube cu- pulate to campanulate or short-tubular, calyx entire or with 2-9 poorly developed lobes/teeth; corolla funnel- form or salverform, white, yellowish, purple or bluish,
corolla lobes 4-9, imbricate or subvalvate, the margins often undulate; stamens 4-9, anthers narrow, sessile or subsessile, dorsifixed, included; ovary 2-9-locular, with 1 pendulous ovule from apex of each elongate-tubular locule, stigma capitate or lobed. Fruits drupaceous, glo- bose to elongate, rounded or angulate in cross-section, the exocarp fleshy but thin, endocarp woody to stony, 2-9-locular.
A genus of 60-80 species in the New World tropics, with a few species in the southwest Pacific and a species widespread on tropical coasts (G. speciosa L.). Some species of Guettarda have in- florescences with two equal cincinnoid branches; these are scorpioid cymes (cincinni) in which the sessile flowers are all in a close line along a single side of the rachis. The leaves of Guettarda are generally thin, often clustered at the ends of branchlets, and with the distal secondaries strong- ly ascending. Domatia are often present, and the 3° veins are usually subparallel. The inflorescences are always axillary, and the flowers are all salver- form in Costa Rican species. This treatment ben- efited from the annotations made by Alfredo Gri- jalva in 1982.
Key to the Species of Guettarda
la. Inflorescences subsessile; plants of Cocos Island and the evergreen Pacific lowlands . . .G. conferta
Ib. Inflorescences short-to long-pedunculate; plants of mainland Central America 2
2a. Bracts subtending the flowers 3-6 mm long, thin-brownish, narrowly ovate-oblong; Pacific slope
and lowlands 3
2b. Bracts subtending the flower absent or less than 3 mm long and caducous 4
3a. Inflorescences with peduncles 2.5-6 cm long; leaf blades usually rounded at the base; Gulf of Nicoya G. brenesii
3b. Inflorescences with peduncles 1-3 cm long; leaf blades acute at the base; western Costa Rica
G. foliacea
4a. Inflorescences with short (to 1 cm) branches, the branches not cincinnoid (scorpioid-cymose) in
appearance 5
4b. Inflorescences with conspicuous lateral cincinnoid (scorpioid) branches more than 1 cm long (with
the flowers all along 1 side 6
5a. Corolla tubes 8-12 mm long, peduncles less than 3 cm long; petioles 4-30(-40) mm long; fruit ca. 12 mm long, subglobose; Pacific and Caribbean lowlands G. macrosperma
5b. Corolla tubes 13-18 mm long, peduncles more than 3 cm long; petioles 20-70 mm long;
fruit ca. 20 mm long, oblong, truncated distally; Golfo Dulce and Panama
G. sanblasenesis
6a. Trees to 30 m tall; leaves and twigs glabrous; flowers becoming 2-4 mm distant on the rachis
[corolla tubes 20-30 mm long] G. turrialbana
6b. Trees to 10(-25) m tall; leaves and stems densely to sparsely puberulent; flowers 0-2 mm distant
on the rachis 7
7a. Stipules glabrous, often broadly overlapping; midvein sparsely appressed strigose along the sides
beneath [corolla tube 16-20 mm long; 1500-2500 m elevation] G. poasana
7b. Stipules pubescent (at least along the midrib); midvein densely pubescent over the entire surface
on the lower side of the leaf blade . 8
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
151
8a. Leaf blades narrowly oblong and coriaceous, densely tomentulose beneath; corolla tube 4-7 mm
long [1200-1900 m elevation] G. tornefortiopsis
8b. Leaf blades elliptic to broadly ovate, drying chartaceous, lacking a densely matted tomentum
beneath; corolla tubes 6-18 mm long 9
9a. Leaf blades rounded and often truncate at the base, to 30 cm long; peduncles to 10 cm long [corolla
tubes 15-18 mm long]; northern Caribbean lowlands G. combsii
9b. Leaf blades usually acute to obtuse at the base, to 22 cm long; peduncles to 4 cm long; evergreen
formations 300-1700 m elevation 10
lOa. Leaf blades with 6-10 pairs of major secondary veins, 3° veins clearly differentiated from the
smaller 4° veins; corolla tubes 13-18 mm long; Central Highlands and evergreen Pacific lowlands
G. crispiflora
lOb. Leaf blades with 3-5 pairs of major secondary veins, 3° and 4° veins little differentiated and parallel
with each other; corolla tubes 6-9 mm long; Caribbean slope Chomelia venulosa
Guettarda brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1303. 1938. Figure 33.
Trees to 10 m tall, leafy branchlets 2-4 mm thick, at first with straight ascending hairs 0.5-1 mm long, terete, soon glabrescent and becoming very dark with numerous narrow lenticels ca. 0.5 mm long; stipules 7-12 mm long, narrowly ovate-triangular, acute, sericeous along the midrib and base, deciduous. Leaves clustered at the end of branchlets, petioles 5-12 mm long, ca. 1 mm thick, with thin ascending hairs 0.5-1.3 mm long; leaf blades (4-)6.5-2 1 cm long, (3-)4.5-l 2 cm broad, broadly ovate to broadly elliptic, apex obtuse or subrotund, base obtuse (and rounded at the petiole) to broadly rounded and subcordate, drying chartaceous and dark above, sparsely hispidulous above with hairs ca. 0.5 mm long, more densely pubescent beneath with thin whitish hairs to 1 mm long, 2° veins 6-8/side, with denser tufts of hairs (domatia) in the vein axils, 3° veins subparallel basally and joining with the opposing tertiary veins at an angle. Inflorescences 3-10 cm long, equally wide, axillary, pe- duncle 2-6.5 cm long, ca. 0.8 mm thick, pubescent, usu- ally bifurcate with 2 main branches and further cymose branching, bracts and bracteoles 4-8 mm long, lanceo- late, pubescent along the midrib, distal axes to 4 mm long often bearing single flowers and resembling pedicels (but with bracteoles at their apex beneath the flowers). Flowers with hypanthium ca. 1 mm long and 1 mm broad, short-tubular and sericeous, calyx tube ca. 1 mm long, entire and more sparsely pubescent distally; corolla white, tube (9-) 14-1 8 mm long, 0.5-0.8 mm diam., with minute appressed-ascending whitish hairs externally, lobes 5, ca. 3 mm long and 1.7 mm broad, margin broad- ly rounded and subentire. Fruits unknown.
Trees of lowland deciduous forest formations near the Bay of Nicoya. Immature inflorescences were collected in June (Brenes 15694 the type), and inflorescences with falling corollas were col- lected in July. Fruiting in July-August and Oc- tober-November. The species is known only from along the Pacific coast of central Costa Rica.
Guettarda brenesii is recognized by the bracteate inflorescences, the long narrow corollas, the broad- ly ovate leaves, and the rocky, seasonally very dry
seaside habitat. This species is probably related to G. foliacea, which shares characters of the inflo- rescence. The type appears to have immature leaves and inflorescences; hence, Standley's description represents minimal measurements. A highly re- stricted range and short flowering season may ex- plain the paucity of collections.
Guettarda combsii Urban, Symb. Ant. 6: 48. 1 909.
Small to large trees (shrubs), 5-30 m tall, trunks to 60 cm dbh, leafy branchlets 2-6 mm thick, with soft whitish erect or ascending hairs 0.5-1 mm long but soon gla- brescent, terete; stipules 6-14 mm long, ovate to lan- ceolate, acuminate, pubescent along the midrib. Leaves often clustered at the ends of branchlets, petioles 2-9 cm long, 1.3-2 mm thick, minutely pubescent; leaf blades 7-20(-28) cm long, 5-13 (-20) cm broad, very variable in shape, from broadly ovate to ovate-oblong or subor- bicular, apex abruptly narrowed and obtuse or short acu- minate, base broadly obtuse to rounded and truncate to subcordate, drying thin-chartaceous, subglabrous or sparsely pubescent above with thin hairs ca. 1 mm long, more densely pubescent beneath with thin whitish hairs 0.5-1 mm long, 2° veins 8-1 I/side, 3° veins parallel and prominent. Inflorescences 12-18 cm long, with long (5- 1 5 cm) peduncles and 2 or 4 distal cincinnoid branches 2-5 cm long, minutely (0.1-0.4 mm) pubescent, bracts 2-4 mm long and ca. 1 mm broad, flowers sessile and closely spaced. Flowers with hypanthium ca. 1 mm long and 1.3 mm diam., with a dense greenish white or pale grayish white tomentum, calyx tube 1.5-2.5 mm long, entire or 2-lobed; corolla greenish white to cream white, tube 15-18 mm long, 1-1.5 mm diam., with dense re- trorse sericeous hairs, corolla lobes 5-7, ca. 3-4 mm long; ovary 4- or 5-locular. Fruits 6-8 mm long, subglobose, covered with a dense minute (0.05 mm) tomentum, gray- ish green, the surface becoming wrinkled.
Trees of evergreen Caribbean rain forest for- mations, from near sea level to 900 m elevation. Flowering in May; fruiting in September in Belize. The species ranges from Belize to southeastern Nicaragua.
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Guettarda combsii is recognized by the broad pubescent leaf blades usually rounded and trun- cate to subcordate at the base, the long-pendun- culate inflorescences with four short cincinnoid branches, and the subglobose fruit. The fact that these plants become very tall trees may account for the paucity of collections in southern Central America. A sterile collection from a tall tree at Minefields. Nicaragua (Proctor et al. 27130 F), sug- gests that this species is also likely to occur in northern Costa Rica.
Guettarda conferta Benth., Hot. voy. Sulph. 106. 1845.
Trees to 10 m tall, with dense, ferruginous hairs on the branchlets, petioles, peduncles, and nerves of the leaves; stipules broadly obovate, 12 mm long, hirsute externally at base, otherwise glabrous, about equaling the petioles, deciduous. Leaves with blades 10-20 cm long, 6-9 cm broad, ovate, apex acuminate, base acute, hirsute on both sides. Inflorescences 2.5-3 cm long, cy- mose, subsessile, the branches recurved and 2.5 cm long or less. Flowers ca. 8 mm long, calyx tube 1-2 mm long, shallowly 3- or 4-dentate; corolla white, sericeous-hir- tellous, tube ca. 10 mm long, corolla lobes 4, ca. 4 mm long, obtuse crispate. Fruits ovoid-tetragonous, 4 mm long, 3 mm diam.. hirsute, 4-locular.
Plants of Cocos Island and the Pacific slope of southern Costa Rica and the Osa Peninsula, 0- 300 m elevation. Flowering in August and Decem- ber; fruiting in January.
Guettarda conferta is distinguished by its short sessile inflorescences. In general aspect this species resembles G. crispiflora.
Guettarda crispiflora Vahl, Ecolog. Amer. 36: pi. 6. 1797. G. chiriquiensisSlandl., Ann. Missouri Hot. Gard. 25: 838. 1938. Figure 32.
Small to medium-sized trees 4-20 m tall, leafy branch- lets 1.5-6 mm thick, at first quadrangular but soon be- coming terete, with short (0.3 mm) thin appressed-as- cending hairs but glabrescent; stipules 8-18(-22) mm long, to 1 cm broad, ovate-elliptic to slightly obovate, apex acute to acuminate, with thin ascending sericeous hairs along the midrib and at the base. Leaves clustered distally, petioles 2-7 cm long, 1.3-2 mm thick, minutely puberulent and with longer hairs along the adaxial side; leaf blades (6-)8-22 cm long, (3-)5-l 1 cm broad, ovate- elliptic or ovate-oblong to ovate-lanceolate or broadly ovate, apex tapering gradually and acuminate or acute, base obtuse to slightly rounded or attenuate and acute, drying stiffly chartaceous, glabrous or minutely (0.2 mm) puberulent above, pubescent beneath with larger (0.4-1 mm) hairs on the major veins and smaller (0.3 mm) thin
whitish hairs on the 3° veins, tufts of hairs (domatia) sometimes present in the vein axils beneath, 2° veins 6- 10/side, distal 2° veins strongly ascending, 3° veins often parallel. Inflorescences 2-6(-8) cm long, equally wide, with a short (4-25 mm) peduncle and 2 diverging cin- cinnoid branches 1 5-30(-60) mm long and enlarging in fruit, the rachis minutely grayish white tomentulose, the flowers sessile and closely (0-4 mm) spaced. Flowers sweet-scented, hypanthium 1.5-2.5 mm long, 1.5-2 mm diam., densely grayish white tomentulous, calyx tube ca. 0.5 mm long, calyx lobes 4, ca. 0.5 mm high; corolla white or pinkish, tube 13-18 mm long, 1-1 .5 mm diam.. narrowly tubular, densely short-sericeous with retrorse or spreading hairs, lobes 5-6 mm long, the lobes with smaller undulate marginal lobes. Fruits ca. 8 mm long and 6 mm diam., oblong and with 4 prominent longi- tudinal ribs, becoming purple and with a white pulp.
Trees of the Caribbean slope cloud forest for- mations and southern Pacific wet forest forma- tions, from 300 to 1700 m elevation. Flowering in January, April, and June-September, with Jan- uary and August collections being most frequent; fruiting in March and June—January. In our area the species is known from the Caribbean slope of the Central Highlands (from near Monteverde eastward to Tapanti, Cartago, and San Joaquin de Dota, San Jose), on the Osa Peninsula, near San Vito, and in the Chiriqui Highlands. The species also occurs in the Lesser Antilles and Trinidad.
Guettarda crispiflora is recognized by the short- pedunculate inflorescences with bifurcate cincin- noid branching, the white corollas with crisped and undulate corolla lobes, the four-angled fruit, longer petioles, and the subparallel (almost lineo- late) minor venation. Guettarda poasana is closely related to G. crispiflora, and material of the two species should be compared when making iden- tifications. Guettarda chiriquiensis was distin- guished by its more densely pubescent vegetative parts, but there are a few intermediate collections in Costa Rica. Nevertheless, the distinctive pop- ulations of the Chiriqui Highlands and adjacent Costa Rica may be worthy of subspecific rank.
Guettarda foliacea Standley, Contr. U.S. Natl. Herb. 18: 139. 1916. Figure 33.
Shrubs or small trees, 3-6(-10) m tall, often branching from the base and with clambering branches, leafy stems 0.9-4 mm thick, at first strigose with thin ascending hairs 0.3-1 mm long, glabrescent, becoming brown and terete, spines often present; stipules 3-5(-12) mm long, trian- gular to lanceolate, strigulose. caducous. Leaves oppo- site, petioles 3-25 mm long, 0.4-1.2 mm thick, ap- pressed strigose to sericeous; leaf blades 3-16 cm long, 2-7 cm broad, elliptic to ovate or obovate, apex acute to short-acuminate, base cuneate to slightly rounded or
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
153
subtruncate, drying thin-chartaceous and greenish, with thin whitish hairs 0.2-0.9 mm long on both surfaces (dense only on the major veins), 2° veins 4-8/side, 4° veins often parallel, domatia of dense hairs present in the vein axils beneath. Inflorescences 2-7 cm long, to 5 cm broad, peduncles 5-30 mm long, 0.5-1.1 mm thick, with thin ascending hairs, usually with a single pair of dichotomous distal branches (each with 3-7 flowers), subtended by lanceolate ciliate bracts 3-8 mm long, flow- ers sessile and crowded in cymes. Flowers densely mi- nutely sericeous externally, hypanthium ca. 1 mm long, calyx tube 1-1.5 mm long, ca. 1.3 mm diam., entire; corolla white, tube 12-20 mm long, lobes 4(-5), 2-5 mm long, rounded distally. Fruits 1-3 cm diam., globose, with a dense minutely velutinous surface, becoming red.
Plants of evergreen or partly deciduous forests, 0-300(-1000) m elevation. In central Panama flowering occurs primarily in late June-early July (Croat, 1978) and fruiting in September-Novem- ber. This species ranges from the Cordillera de Tilaran to Colombia.
Guettarda foliacea is recognized by its small in- florescences, lack of calyx lobes, thin variable leaves on slender petioles, and minor venation, often with a small group of parallel veins (sublineolate). The spines are rarely seen on herbarium collections. The westernmost collection (84°53'W, Haber et al. 4775 CR, MO) came from the edge of cloud forest at 1 000 m elevation with immature flowers in May.
tomentulose, calyx tube 0.5-1 mm long, entire; corolla white or yellowish, tube 8-13 mm long, 0.5-1.3 mm diam., narrowly tubular, lobes 3-4 mm long, rounded distally. Fruits 10-18 mm long, globose to oblong, yel- lowish brown or grayish with a dense covering of minute (0.1-0.2 mm) velvet-like or matted hairs.
Trees and shrubs of both evergreen and decid- uous forest formations, from near sea level to 1 000 m elevation on the Pacific slope and from near sea level to ca. 500 m on the Caribbean slope. Flow- ering in March-November, with the majority col- lected in May; fruiting in July-January. The spe- cies ranges from southern Mexico to Panama.
Guettarda macrosperma is recognized by its small cymose inflorescences, rounded fleshy fruit usually over 1 cm diam., and often smaller leaves with the subparallel 3° veins usually meeting at angles between the 2° veins (> -shaped). Trees in the Caribbean lowlands may reach 30 m in height; the same trunk and bark characteristics are found in trees of both seasonally dry forest and evergreen rain forests (N. Zamora, pers. comm.). Some spec- imens may resemble Chomelia panamensis. It ap- pears that Guettarda divaricata (Roem. & Schult.) Standl. of Mexico is closely related, and the two may be part of a more broadly defined taxon. Ma- lacahuite is a common name.
Guettarda macrosperma J. D. Smith, Bot. Gaz. 1 8: 204. 1893. Figure 33.
Shrubs or more often trees, 4-12(-30) m tall, trunks often fluted or with deep depressions, with dark bark exfoliating in patches, leafy branchlets 1.2-4 mm thick, sparsely puberulent with thin ascending hairs 0.5-1.5 mm long, soon glabrescent and becoming dark brown or blackish with short (0.3-1.2 mm) grayish lenticels; stipules 3-8(-12) mm long, triangular-lanceolate, dense- ly sericeous on the back with longer hairs, caducous. Leaves often crowded at the ends of branchlets, petioles (4-)10-45 mm long, 0.8-1.7 mm thick, with straight ascending hairs; leaf blades (4-)6-18 cm long, (2-)3-l 1 cm broad, broadly elliptic, broadly ovate-elliptic, obo- vate-elliptic, or elliptic-oblong, apex abruptly narrowed and short-acuminate to acute or obtuse, base obtuse to rounded and subtruncate, drying chartaceous to stiffly chartaceous, sparsely pubescent with thin short (0.3-0.7 mm) hairs and glabrescent, with thin short (ca. 0.3 mm) ascending hairs along the 3° veins beneath and with lon- ger and denser hairs along the major veins, 2° veins 4- 8/side, 3° veins rarely subparallel, often with tufts of hairs (domatia) in the vein axils. Inflorescences 2-6 cm long, primary peduncles 4-38 mm long, usually with 2 short distal branches, densely ascending sericeous, bracts 2-3 mm long, flowers sessile and closely crowded (not clearly cincinnoid in arrangement). Flowers with hypanthium 1-1.5 mm long and ca. 1 mm diam., densely whitish
Guettarda poasana Standl., J. Wash. Acad. Sci. 1 8: 182. 1928. Figure 32.
Small trees, 3-10(-15) m tall, trunk to 40 cm thick, bark shredding off in oblong patches, leafy branchlets 2- 5 mm diam., somewhat flattened in early stages and glabrous, drying dark but becoming grayish in age, len- ticels difficult to see; stipules 12-20 mm long, 8-10 mm broad, ovate and long-acuminate at apex, the stipules overlapping on the sides, glabrous. Leaves clustered near the ends of branches, petioles (2-)3-7(-10) cm long, 0.8- 1.6 mm thick, glabrous and drying dark; leaf blades 7- 14(-16) cm long, 2.5-6(-9) cm broad, broadly elliptic- ovate to broadly elliptic or elliptic, apex gradually ta- pering and short-acuminate, base obtuse to acute and slightly decurrent on petiole, drying stiffly chartaceous, glabrous and lustrous above, minor venation with thin whitish ascending hairs ca. 0.3 mm long beneath, with longer (0.5-1 mm) straight hairs on the mid vein and secondaries beneath, 2° veins 5-8/side, the distal strongly ascending, 3° veins often subparallel but not prominent beneath, small tufts of hairs (domatia) often present in the vein axils beneath. Inflorescences axillary, 3-6 cm long and equally wide, peduncles 1-3 cm long, glabrous or sparsely puberulent, with 2 cincinnoid branches 2- 3.5 cm long, the flowers 5-11 on each branch and sessile, ca. 1-4 mm distant. Flowers sweet-scented, hypanthium 1.5-2 mm long, 1.3 mm diam., glabrous or sparsely puberulent, calyx tube 0.5-1 mm long, lobes ca. 0.3 mm long; corolla white, reddish, or lavender, tube 1 6-20 mm
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long 1-1.4 mm diam., densely tomentulose externally, lobes 4-5, 4-6 mm long, usually white with fringed- undulate smaller lobes; tips of the anthers exserted 1-2 mm from the mouth of the tube. Fruits becoming 8 mm long and 6 mm diam., oblong and with 4 prominent longitudinal ridges, reddish purple.
Trees of evergreen cloud forest formations, 1300-2200(-2700?) m elevation (down to 1 100 m on the northern vocanoes). Flowering in March and May-November (mostly in June); fruiting probably throughout the year. This species is en- demic to Costa Rica and ranges from the Cordille- ra de Guanacaste in the west to the eastern slopes of Volcan Barva.
Guettarda poasana is recognized by its glabrous stipules and stems, long-petiolate leaves, narrowly tubular flowers with whitish fringed lobes, four- angled fruit, and restricted cloud forest range. The petioles dry dark and are sometimes pink in life. This species is closely related to G. crispiflora and might be considered a subspecific element of that species. However, though their ranges overlap slightly, G. crispiflora and G. poasana do not grow in the same locality; G. poasana is generally found at higher altitudes.
Guettarda sanblasensis Dwyer, Ann. Missouri Bot. Gard. 67: 204. 1980. Figure 33.
Trees, 12-20 m tall, leafy branchlets 1.5-4 mm thick, with short (0.3 mm) appressed-ascending hairs, quickly glabrescent, terete; stipules 3-6 mm long, densely seri- ceous with pale yellowish ascending hairs 0.5-1 mm long, early caducous. Leaves 1-5 cm distant at the ends of branchlets, petioles 2-7 cm long, 0.7-1 .3 mm thick, with slender appressed-ascending hairs; leaf blades 1 1-22 cm long, (4-)6-l 2 cm broad, broadly elliptic to broadly ovate- elliptic, apex abruptly narrowed and short-acuminate (or acute), base obtuse to slightly rounded, drying charta- ceous and dark brown above, lustrous above and with scattered thin appressed hairs 0.2-0.3 mm long, the hairs more numerous and longer (0.3-0.5 mm) beneath, 2° veins 5-9/side, the distal arcuate-ascending, 3° veins prominent above and below and paler in color beneath, subparallel or > -shaped between the secondaries, with minute tufts of hair (domatia) in the vein axils beneath. Inflorescences 6-10 cm long, primary peduncles to 6 cm long, ca. 1.2 mm thick and minutely appressed-puber- ulent, bifid but the 2 branches with additional dichot- omous branches (not scorpioid/cincinnoid), pedicels 0- 2 mm long, bracts minute (0.5 mm) and caducous. Flow- ers with hypanthium ca. 1.5 mm long and 1.2 mm diam., with longitudinal ribs, calyx tube 2-3 mm long and ca. 2 mm diam., minutely velutinous, subentire distally; corolla white or pink, tube 13-18 mm long, 1-1.3 mm diam., minutely (0.2 mm) ascending-sericeous, lobes 5- 6, 4-5 mm long and 1-1.5 mm broad, oblong and entire, stigma 0.5 mm long, subglobose. Fruits 2-2.2 cm long
and 8-12 mm thick, oblong-obovoid, abruptly rounded (truncated) at apex, persisting calyx 1-3 mm long, 1.5- 2 mm diam., surface minutely velutinous and yellowish or grayish brown.
Trees of evergreen forest formations of the Pa- cific lowlands, from near sea level to 600 m ele- vation and usually on limestone. Flowering in July- September; fruiting in September-November. This species is known only from a few collections in the Golfo Dulce region of Costa Rica; it ranges to eastern Panama.
Guettarda sanblasensis is recognized by the broadly elliptic leaves on long slender petioles, the long-pedunculate inflorescences with dichoto- mous or cymose distal branching, long-tubular co- rollas with five or six entire oblong perianth lobes, and oblong fruit with truncated apex and usually persisting calyx tube.
Guettarda tournefortiopsis Standl., Publ. Field Columb. Mus., Bot. Ser. 7; 293. 1931. Tour- nefortiopsis reticulata Rusby, Bull. New York Bot. Gard. 4: 369. 1907, non G. reticulata Gri- seb., 1863. Figure 32.
Small trees, 4-10 m tall, leafy branchlets 1.5-6 mm thick, at first densely tomentose with soft wooly hairs to 2 mm long, 4-angular, soon .glabrescent and dark with lenticels 0.3-0.7 long, becoming terete and pale grayish; stipules 1 2-20 mm long, narrowly ovate-triangular and acuminate, tomentose, usually early deciduous. Leaves with petioles 13-35 mm long, 1.5-2.2 mm thick, gla- brescent and drying dark; leaf blades 9-16 cm long, 2- 5 cm broad, lanceolate to narrowly oblong-lanceolate or narrowly elliptic-oblong, apex acute to short acuminate, base obtuse to acute and usually with the margin revo- lute, drying subcoriaceous, dark brown above, glabrous and usually lustrous above, with the major and minor veins slightly impressed above and the surface slightly rugose, densely tomentulous between the veins beneath and whitish to pale brown in color, 2° veins 9-1 I/side, domatia present. Inflorescences 2-5 cm long and equally wide, becoming 8 cm long in fruit, peduncles 1 2-20 mm long (to 30 mm in fruit), 1 .5 mm thick and tomentulous, with 2 cincinnoid branches 2-6 cm long and each branch bearing up to 20 flowers in 2 rows along 1 side, flowers sessile and closely crowded, bracts absent. Flowers with hypanthium 1 .5-2 mm long, covered by a dense tomen- tum, calyx lobes ca. 1 mm broad, broadly obtuse and difficult to see; corolla reddish to coral pink, white with- in, tube 4-7 mm long, ca. 1.5 mm diam., densely pu- berulent with retrorse hairs externally, lobes 5, 1-2 mm long. Fruits 5-8 mm long, 4-6 mm diam., globose to rounded-oblong, 5-angled during development, sessile, becoming purple or black.
Trees of wet montane cloud forest formations, from 800 to 1900 m elevation. Collections with
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
155
flowers (or flower buds) and fruits have been made in January-March and August-October. Known only from the Cordillera de Tilaran and the west- ern parts of the Cordillera de Talamanca, prov- inces of Cartage and San Jose, in Costa Rica. The species is also known from the Chiriqui Highlands and was originally described from Bolivia.
Guettarda tournefortiopsis is easily recognized because of its stiff narrow leaves densely tomen- tulose beneath (when young), the two-branched scorpioid inflorescences (resembling those found in the Boraginaceae), small tomentulose red flow- ers, and small sessile fruits.
and long corolla tubes. The tall size of this dis- tinctive species may explain why it had not been collected before 1986. The collections from the Osa Peninsula differ in a number of ways from the type; more material is needed to assess the pattern of variation.
Hamelia Jacquin
REFERENCE— T. S. Elias, A monograph of the genus Hamelia (Rubiaceae). Mem. New York Bot. Gard. 26: 81-144. 1976.
Guettarda turrialbana Zamora & Poveda, Ann. Missouri Bot. Card. 75: 1 157. 1988. Figure 30.
Trees to 30 m tall and with trunks 40 cm dbh, bark exfoliating, leafy branchlets 1.5-6 mm thick, glabrous and dark in early stages, terete, becoming pale grayish and with few broadly ellipsoid lenticels 0.8-1 .5 mm long; stipules ca. 1 2-25 mm long, triangular and acuminate, overlapping, glabrous, deciduous. Leaves clustered at the ends of branchlets, petioles 2-3.5 cm long, 1-2 mm thick, glabrous and drying dark; leaf blades 9-26 cm long, 4- 1 2 cm broad, oblong to broadly elliptic-oblong or ovate- oblong, apex abruptly narrowed and short-acuminate, base abruptly narrowed or rounded and obtuse, drying stiffly chartaceous, dark brown above, glabrous and lus- trous above, subglabrous beneath with tufts of hairs (domatia) in the vein axils, 2° veins 8-10/side, 3° veins slightly raised above but not clearly subparallel. Inflo- rescences 5-11 cm long, primary peduncles 3-5 cm long, 1-1.3 mm thick, glabrous and drying dark, cymose with 2 primary branches, the branches 2-3 cm long and with 3-5 flowers 2-5 mm distant on the rachis (or the branch- es very short and all the flowers close together in Hammel et al. 16848 CR, MO), pedicels short (1 mm) or absent, flower subtended by short (0.5 mm) broad bracts. Flow- ers glabrous externally, with the sweet odor of Coffea flowers, hypanthium ca. 2 mm long and 1.2 mm diam.. calyx cup ca. 2 mm long, ca. 3 mm diam. distally and entire; corolla white, tube 20—40 mm long, 2-2.8 mm diam., puberulent within, lobes 5, ca. 5 mm long and 2 mm broad, oblong and entire, distally rounded. Fruits 24-28 mm long, 6-12 mm diam., oblong, glabrous.
Trees of lowland rain forest formations; col- lected at 600 m elevation, below the CATIE site near Turrialba on the Caribbean slope, and on the Osa Peninsula at 200-300 m elevation. Flowering in May (Hammel et al. 16848 CR, F, MO) and June (H err era 4246 MO, Zamora et al. 1263 CR holo- type, F); fruiting in August and September. En- demic.
Guettarda turrialbana is recognized by its tall stature, mostly glabrous parts, oblong leaves, few- flowered bifurcate inflorescences, entire calyx cup,
Shrubs or small trees, branchlets usually slender, terete or 4-angled in cross-section, glabrous to densely puber- ulent; stipules interpetiolar, usually small, entire or with an awn (or trilobate), caducous or less often persistent. Leaves opposite and decussate or in whorls of 3-5/node, usually regularly spaced by well-developed internodes, often long-petiolate, usually thin in texture, glabrous or puberulent, pinnately veined, domatia present in some species, conspicuous raphides visible on the dried leaf surfaces in a few species. Inflorescences terminal or less often axillary, 3-many-flowered, often thyrse-like with cymose branches or with helicoid branches, pedunculate, the flowers often all along 1 side of the distal branches, sessile or short-pedicellate. Flowers bisexual and radially symmetrical, monomorphic, hypanthium urceolate to tubular, calyx tube minute or absent, calyx lobes 5, small, rounded to elongate, usually persisting; corolla narrowly tubular to funnelform or campanulate-urceolate (with a short narrow base), bright yellow to orange, orange-red, or deep red, with 5 longitudinal ribs alternating with the lobes, corolla lobes 5, erect to recurved, slightly imbri- cate in bud; stamens 5, filaments inserted at the base of the corolla tube, flattened, anthers long-linear, sagittate at the base and usually with the connective prolonged distally, included or partly exserted; ovary (4-)5-locular, with axile placentation and many ovules in each locule, style narrowly cylindrical, stigmas 1-5. Fruit a fleshy berry, oblong to ovoid or subglobose, with 5 longitudinal ribs, terminated by the circular calyx scar and a conical disc; seeds numerous, plano-convex or angled, foveolate.
Hamelia ranges from southern Florida, U.S.A., through Mexico, Central America, and the West Indies into tropical South America. Elias recog- nized 16 species in his fine monograph. The nar- rowly tubular or funnelform yellow to orange or red flowers are usually all aligned on the upper- most sides of the inflorescence branches. In some, the inflorescences have longer cincinnus-like branches. Many of our species have three to four leaves at distal nodes and a number have long- petiolate leaves; most have small tufts of hairs (domatia) in the vein axils beneath. Individual plants may vary greatly within many species, and this makes identification difficult. In addition, there
156
FIELDIANA: BOTANY
may be intermediates or hybrids between some of our species.
The genus is divided into two subgenera. Sub- genus Hamelia has narrowly tubular flowers that do not enlarge distally at anthesis and are red, orange, or yellow in color. Subgenus Amphituba has the yellow corolla tube slightly to conspicu- ously expanded distally at anthesis. However, the
flowers of subgenus Amphituba remain narrowly tubular until just before anthesis and may be dif- ficult to distinguish from those of subgenus Ha- melia on this account. Species of Hoffmannia may be similar, but they have consistently axillary in- florescences; compare Deppea, with papery cap- sules.
Key to the Species of Hamelia
la. Young stems and undersides of leaves conspicuously pubescent with hairs 0.2-0.5 mm long .. 2 Ib. Young stems and undersides of leaves glabrous or with minute (0.1-0.2 mm) inconspicuous hairs
4
2a. Stipules 7-17 mm long; leaf blades with 8-18 major 2° veins on each side, with hairs to 1 mm long; corolla expanded distally at anthesis, corolla lobes 2-6 mm long [calyx lobes 0.5-2 mm
long] H. xerocarpa
2b. Stipules 2-5(-8) mm long; leaf blades with 4-9 major 2° veins on each side, with hairs to 0.5
mm long; corolla narrowly tubular, corolla lobes 1-2 mm long 3
3a. Calyx lobes 0-0.7 mm long; seeds 0.6-0.9 mm long; petioles 10-80 mm long; common wide- spread plants, 0-1 500 m elevation H. patens
3b. Calyx lobes 2—4 mm long; seeds 1-1.2 mm long; petioles 3-20 mm long; uncommon plants of
the Caribbean lowlands, 0-300 m elevation H. rovirosae
4a. Corolla tube 8-1 3 mm long at anthesis; fruits 5-10 mm long 5
4b. Corolla tube 1 3-35 mm long at anthesis; fruits 7-16 mm long 6
5a. Inflorescences 3-9 cm long, corolla tube becoming expanded distally at anthesis; leaf blades with 5-9 major 2° veins on each side, 5-1 7(-23) cm long, usually drying greenish to pale grayish;
fruits 4-7 mm long; Caribbean and evergreen Pacific lowlands H. axillaris
5b. Inflorescences 8-18 cm long, corolla tube narrowly tubular at anthesis; leaf blades with 7-12 major 2° veins on each side, 10-27(-32) cm long, usually drying reddish brown to pinkish gray;
fruits 6-10 mm long; evergreen Pacific slope H. magnifolia
6a. Leaf blades with 8-13 major 2° veins on each side; corolla tube distally widened (4-7 mm) at
anthesis; corolla lobes 2-4 mm long 7
6b. Leaf blades with 3-9 major 2° veins on each side; corolla tube narrowly (2-4 mm) tubular at anthesis
(in H. patens, often broader in H. calycosa)
7a. Leaf blades 3-6 cm broad; corolla tube 1 3-22 mm long; 0-800 m elevation ...//. xerocarpa
7b. Leaf blades 4-12 cm broad; corolla tube 25-35 mm long; 700-1600 m elevation
H. macrantha
8a. Leaf blades with 6-9 pairs of 2° veins, petioles 10-80 mm long; sepal lobes 0-0.8 mm long, corolla
lobes 1-2 mm long; common, 0-1 500 m elevation H. patens
8b. Leaf blades with 3-7 pairs of 2° veins, petioles 8-20 mm long; sepal lobes 1-2.5 mm long, corolla lobes 3-6 mm long; not reported from Costa Rica H. calycosa
Hamelia axillaris Sw., Prodr. 46. 1788. H. lutea Rohr ex Smith in Rees, Cyclop. 17.1811. Figure
42.
Shrubs, 1-5 m tall or small treelets to 5 m tall, leafy branchlets 1 .2-4 mm thick, glabrous, with 4 longitudinal ribs and 4-angular in cross-section but becoming terete; stipules 2-6(-8) mm long, ca. 1 mm broad, triangular to narrow with folded margins. Leaves opposite (rarely 4/node), petioles l-4(-7) cm long, ca. 1 mm wide, gla-
brous to sparsely and minutely papillate-puberulent; leaf blades 5-17(-23) cm long, 2-8 cm broad, narrowly el- liptic, elliptic-oblong or obovate to narrowly obovate- oblong, apex acuminate with tip ca. 1 cm long, base attenuate and decurrent on petiole, drying chartaceous or membranaceous, glabrous above, glabrous or mi- nutely (0. 1 mm) papillate-puberulent beneath, with tufts of hairs (domatia) in the vein axils, 2° veins 5-9/side and loop-connected near the margin, short (0. 1-0.3 mm) linear cystoliths visible on the lower surface when dry. Inflorescences axillary or terminal, 3-8 cm long, 3-8 cm
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
157
broad, compound dichasia with lateral branches 1-4.5 cm long, often scorpioid with 3-15 secund flowers on the uppermost side, peduncles 5-15 mm long, glabrous or minutely and sparsely puberulent, bracts 0.5-1 mm long, narrow, flowers sessile or with pedicels. Flowers with hypanthium 1.5-3 mm long, ca. 1.2 mm diam., glabrous, calyx lobes 0.5-1.5 mm long, 1 mm wide at the base, glabrous to puberulent; corollas yellow, nar- rowly urceolate to funnelform, tube 8-13 mm long, ca. 1 mm diam. near the base and 3-5 mm wide distally, glabrous, lobes 1-2 mm long, broadly triangular; sta- mens with filaments 4-5 mm long, anthers 5-8 mm long with apical connective 0.5 mm long, style 8-10 mm long, stigmas 3-4 mm long. Fruits 4.5-7 mm long, 3-4 mm diam., ovoid-oblong to subglobose, disc 0.5 mm high and 0.7 mm broad; seeds ca. 1 mm long.
Plants of the evergreen Caribbean slope and the Osa Peninsula, from near sea level to 600(-1000) m elevation. Flowering primarily in late June-Oc- tober; fruiting in February and June-December. The species is known from southern Mexico, Be- lize, Guatemala, the larger West Indian islands, and from Nicaragua southward to Venezuela and Bolivia.
Hamelia axillaris is recognized by its short yel- low corollas funnelform at anthesis, short rounded fruit, general lack of pubescence, and usually com- pact inflorescences with flowers along one side of the distal branches. There may be intermediates between this species and H. magnifolia on the Osa Peninsula; compare the extreme upper-right figure in Figure 42 (based on Utley & Utley 1208 F).
Hamelia calycosa J. D. Smith, Bot. Gaz. 12: 132. 1887.
Shrubs or small trees to 12 m tall, leafy branchlets 1- 3 mm thick, glabrous or glabrescent; stipules l-2.5(-4) mm long, ca. 1 mm broad at the base, subulate or with a narrow awn, minutely puberulent. Leaves opposite or 3-4/node on distal branches, petioles 6-20 mm long, 0.4-0.8 mm broad, glabrate; leaf blades 3-1 1(-15) cm long, l-3.5(-5) cm broad, lanceolate to narrowly elliptic- oblong or elliptic-oblanceolate, apex tapering gradually and acuminate, tip ca. 7 mm long, base acute to cuneate and decurrent on petiole, drying membranaceous to chartaceous, glabrous above, glabrous or minutely (0. 1- 0.2 mm) pubescent beneath or rarely with longer (0.7 mm) thin hairs, with tufts of hairs (domatia) in the vein axils beneath, 2° veins 3-7/side. Inflorescences terminal or rarely axillary, 3-10 cm long, with 4-24 flowers, pe- duncles to 2 cm long and often with 2 dichotomous branches distally, usually minutely puberulent, bracts 1- 2 mm long, subulate or triangular, flowers secund and with pedicels l-5(-8) mm long. Flowers with hypanthi- um 2-4 mm long, oblong, glabrous or sparsely pubescent, calyx lobes 1-3 mm long, ca. 0.5 mm wide, narrowly oblong, caducous; corolla funnelform, yellow or pale or- ange (striped with maroon), tube 1 5-24(-32) mm long,
constricted (1.5 mm) near the base and 8-10 mm diam. distally, pubescent externally, lobes 3-6 mm long, ovate; stamens 5, filaments 4—6 mm long, anthers 15-18 mm long, 1 mm wide, distal rounded connective ca. 1 mm long, style to 14 mm long, stigmas 5 and connate, ca. 1.2 mm long. Fruits 7-16 mm long, 4-8 mm thick, cy- lindrical, disc conical and 1-3 mm long; seeds 1-1.2 mm long.
Trees and shrubs of Caribbean rain forest for- mations, from near sea level to 1500 m elevation. Flowering is in April-September in northern Cen- tral America. The species ranges from southern Mexico, Belize, Guatemala, Honduras, Panama, and Colombia to Peru.
Hamelia calycosa is recognized by the larger sepal lobes, longer corollas expanded distally, ped- icellate fruit, and smaller leaves with short peti- oles. Although not yet collected in Costa Rica, this species is likely to be present.
Hamelia macrantha Little, Carib. Forester 9: 274. 1948.
Shrubs or small trees to 8(-l 2) m tall and 1 3 cm dbh, leafy branchlets 2-6 mm thick, 4-angular in cross-sec- tion, glabrous or minutely puberulent; stipules 3-7 mm long, 1-2.5 mm broad at the base, triangular to awl- shaped, glabrous or rarely puberulent. Leaves opposite but 3-4 at distal nodes, petioles (1 .5-)6-l 1 cm long, 1 .3- 2.7mm broad, glabrous or puberulent; leaf blades (7-) 1 2- 23(-27) cm long, (3-)4-12(-15) cm broad, broadly ob- long or elliptic-oblong to ovate or obovate, apex obtuse to abruptly short-acuminate, base rounded and subtrun- cate to obtuse (cuneate), drying membranaceous to char- taceous, glabrous above, glabrous to minutely (0. 1 mm) papillate puberulent beneath in Costa Rica, domatia sometimes present, 2° veins 8-12/side and loop-con- nected near the margin. Inflorescences terminal or ax- illary, 6-17 cm long and wide, to 15 cm broad, panic- ulate, 20-many-flowered, peduncles 2-8 cm long, 1-3 mm thick, lateral branches to 6 cm long, bracts 0.4-0.7 mm long, ovate and acute, glabrate or puberulent, flow- ers sessile to short (2 mm) pedicellate. Flowers with hy- panthium 3-5 mm long, 1.5-2 mm diam., glabrous, calyx lobes 0.7-1 .5(-2) mm long, 1-1.5 mm broad at the base, triangular-subulate; corolla yellow, tubular-funnelform, tube 23-35 mm long, constricted near the base and 5-
7 mm diam. distally, glabrous, lobes 3-5 mm long, ovate, glabrous; stamens with filaments 6-8 mm long, anthers 16-20 mm long, included, connective prolonged ca. 1 mm beyond the thecae; style to 20 mm long, stigmas 5-
8 mm long. Fruits 11-15 mm long, 3.5-5 mm diam., ellipsoid, red becoming black and lustrous, ovarian disc conical; seeds 0.8-1 mm long.
Trees and shrubs of evergreen cloud forest and lower montane forest formations on both the Ca- ribbean and Pacific slopes, from 700 to 1500 m elevation in Costa Rica. Flowering in June-Sep-
158
FIELDIANA: BOTANY
tember in Costa Rica and Panama; fruiting in Au- gust-September. The species ranges from Costa Rica and Panama to Colombia and Ecuador.
Hamelia macrantha is recognized by the large yellow corolla tube slightly widened distally at an- thesis, larger leaves with many secondary veins and often with long narrow petioles, and the mon- tane habitat (in Costa Rica). Note that the widened corolla tube is only seen at anthesis. The foliage of this species resembles that of some specimens of H. xerocarpa variety costaricensis, and the two species may be difficult to separate.
Hamelia magnifolia Wernham, J. Bot. 49: 210. 1911. Figure 42.
Shrubs or small trees to 5(-6) m tall, trunks to 1 2 cm dbh, leafy branchlets 2-7 mm thick, with 4 longitudinal ridges and quadrangular in cross-section, glabrous or minutely (0. 1 mm) papillate-puberulent; stipules 4-9 mm long, to 2 mm wide at the base, narrowly triangular (rarely bifid), glabrate. Leaves opposite (rarely 4/node), petioles l-5(-7) cm long, 1.7-2.3 mm broad; leaf blades 10-27(-32) cm long, 4-ll(-15) cm broad, broadly ob- long to ovate-oblong, elliptic-oblong or ovate-elliptic, apex short-to long-acuminate with tip to 2 cm long, base rounded and truncate to obtuse, drying stiffly charta- ceous, glabrous above, glabrous beneath, 2° veins 7- 1 2(-l 5)/side, some 3° veins subparallel, domatia absent. Inflorescences terminal, 8-12(-18) cm long, to 15 cm wide, paniculate with opposite branching, with more than 60 flowers, peduncles 1-3 cm long, distal branches to 7 cm long and with secund flowers, minutely papillate- puberulent or glabrous, bracts 0.5-1 mm long, narrow pedicels 0-1 mm long. Flowers with hypanthium 2-3 mm long, 0.8-1 .5 mm diam., minutely puberulent, calyx lobes 0.3-0.6 mm long, 1 mm broad at the base, deltoid or broadly rounded, puberulent along the margin; corolla narrowly tubular at anthesis (also somewhat narrowed above the base and below the middle), yellow, tube 10- 13 mm long, 2-3 mm diam., lobes 1.2-3 mm long; sta- mens with filaments 2-3 mm long, anthers 6-8 mm long, apical connective ovate and ca. 1 mm long. Fruits 6-10 mm long, 3-4 mm diam., oblong to broadly ellipsoid, glabrous, ovarian disc ca. 1 mm long, red; seeds 0.4-0.5 mm diam.
Plants of evergreen forest formations on the Pa- cific slope, from 1 00 to 900 m elevation in Costa Rica. Flowering in January-August; fruiting in June-September and December-January. The species is restricted to the evergreen formations of the Pacific slopes of southern Costa Rica and ad- jacent areas in Chiriqui Province, Panama.
Hamelia magnifolia is recognized by the small narrowly tubular flowers in large inflorescences, the large leaves often rounded at the base and with many secondary veins, the smaller fruits, and the
restricted geographic range. Standley (1938) listed zorillo Colorado as a common name.
Hamelia patens Jacq., Enum. PI. Carib. 16. 1760. H. patens var. glabra Oersted, Vidensk. Meddel. Naturhist. Foren. Kjobenhavn 1852: 42. 1853. H. viridifolia Wernham, J. Bot. 49: 213. 1911. Figure 42.
Shrubs or small trees, 2-7 m tall, leafy branchlets 1- 7 mm thick, glabrous or pubescent with crooked trans- lucent or yellowish hairs to 0.6 mm long (in variety patens), 4-angled in early stages but becoming terete; stipules 2-6 mm long, narrowly oblong to linear, pu- bescent, as many as the leaves at each node. Leaves usually 3(-4) at distal nodes (less often opposite or rarely 5/node), often unequal at the same node, separated by well-developed internodes, petioles (8-)20-55(-80) mm long, 0.6-1.6 mm wide, glabrous to densely pubescent; leaf blades 5-17(-23) cm long, l-7(-10) cm broad, el- liptic-oblong, ovate-elliptic, ovate-oblong, or elliptic ob- ovate-elliptic, apex usually short-acuminate, base acute to attenuate (obtuse in larger leaves) and decurrent on petiole, drying membranaceous to thin-chartaceous, gla- brous (or sparsely pubescent in variety patens) above, glabrous to densely villosulous beneath with thin straight or curved whitish hairs 0.2-0.5 mm long (rarely with scurfy yellowish hairs), tufts of hairs (domatia) often present in vein axils beneath, 2° veins 6-9/side, small (0.2 mm) linear raphides (cystoliths) resembling ap- pressed hairs often visible on ,the dark upper surface of dried leaves. Inflorescences terminal, solitary or 2-4, 4- 9(-15) cm long, to 12(-20) cm broad, an open panicle with 2-3 nodes on the rachis and 2-4 branches at each node, peduncles 1-3 cm long (to 5 cm in fruit), minutely papillate-puberulent (less often with conspicuous hairs), often orange to coral red, primary branches often ending in a flower and 2 long distal secondary branches bearing a row of sessile or subsessile flowers along the upper side, bracts 0.3-1 mm long, triangular, distal flowers with pedicels 1-5 mm long. Flowers with hypanthium 1.5-3 mm long, 1-2 mm diam., longitudinally ribbed, deep red, glabrate to densely short-villous. calyx lobes 0.5-1 mm high, rounded or broadly triangular, persisting; co- rolla narrowly tubular at anthesis, orange to reddish or- ange, tube (12-)14-18(-23) mm long, 1.5-3 mm diam., lobes 1-2.5 mm long, 1-2 mm broad at the base, tri- angular; stamens with filaments 5-7 mm long, anthers 8-12 mm long, included or slightly (3 mm) exserted. the apiculatc connective 0.5 mm long; stigmas 3-5 mm long. Fruits 6-1 3 mm long, 4-10 mm diam., oblong to ovoid- oblong (subglobose), red becoming black or bluish, gla- brous or with hairs ca. 0.2 mm long, raphides often visible on the surface, with a ring around the top formed by the calyx scar ca. 3 mm diam., disc to 1 mm high and inconspicuous; seeds 0.5-1 mm long.
Common shrubs and treelets of open early sec- ondary growth in evergreen and partly deciduous forest formations, from near sea level to 1 600 m elevation. Flowering and fruiting in all months of
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
159
the year in Costa Rica. The species ranges from southern Florida, Mexico, the West Indies, and Central America southward to northern Argentina and Paraguay (the species is not found in the Guianas, northeastern Brazil, the central Amazon basin, or central Brazil).
Hamelia patens is recognized by its bright or- ange or red-orange flowers with narrowly tubular corollas with small lobes and included anthers and the usually three- or four-leaved distal nodes. The open inflorescences with dichotomous or dichasial branching and with secund flowers often in a row along the upper side of distal inflorescence branch- es are also distinctive. These shrubs and little tree- lets of open evergreen lowland secondary sites are among the most common and conspicuous of Cos- ta Rica's woody flora. They appear to germinate only in open sunny sites. The breeding biology of this species was studied by Bawa and Beach ( 1 983). Standley ( 1 938) listed the common names anileto, azulillo, coralillo, palo camaron, pissi, zorillo, and zorillo real, and he cited Pittier for the Indian names pili-tso (Guatuso) and tsus-krd (Brunka).
Hamelia patens variety patens has the leaves sparsely to densely villous (especially on the lower surface), and the flowers are sparsely to densely villous externally. Variety glabra Oersted has the leaves glabrous above and sparsely villous or pu- berulent on the veins beneath, and the flowers are usually glabrous externally. There seem to be no ecological or geographic distinctions between the varieties, and they may be no more than glabrous and puberulent forms found within the same pop- ulations.
Hamelia rovirosae Wernham, J. Bot. 49: 211. 1911. Figure 34.
Shrubs or slender treelets to 5(-10) m tall, leafy branchlets 0.8-3.5 mm diam., with curved or crooked multiccllular hairs 0.3-1 mm long and often in longi- tudinal rows, with 4 longitudinal ribs and quadrangular in early stages but becoming terete and glabrescent; stip- ules 2-6(-8) mm long, with a very short (1 mm) broad base and long narrow linear awn, puberulent, caducous or persisting with the leaves. Leaves usually 3/node (rare- ly opposite), petioles 3-14(-20) mm long, 0.5-1 mm thick, pubescent with short crooked hairs; leaf blades (2.5-)5-15 cm long, (1.5-)2-6 cm wide, elliptic-oblong to elliptic-ovate, elliptic or elliptic-obovate, apex acute to short-acuminate, base acute to attenuate and slightly decurrent on petiole, drying thin-chartaceous, glabrous to sparsely pubescent above, sparsely to densely villous beneath with thin straight or crooked hairs 0.2-0.5 mm long, with denser tufts of hairs (domatia) in the vein axils, 2° veins 3-7 /side and weakly loop-connected near
margin. Inflorescences 4-12 cm long, equally broad, pe- duncles to 3 cm long, villous, floral rachis with dichot- omous distal branches bearing 2-8 flowers, bracts to 0.5 mm long, caducous, flowers sessile or short (1-2 mm) pedicellate. Flowers with hypanthium 2-4.5 mm long, villous with curly hairs ca. 0.5 mm long, calyx lobes 2- 4 mm long and enlarging in fruit, 1 mm wide and oblong, villous; corolla narrowly tubular at anthesis, reddish or- ange to bright red or dark red, tube 16-22 mm long, 2- 3 mm diam., minutely villous with hairs ca. 0.5 mm long (often in longitudinal rows), lobes 1-2 mm long, 1- 1.5 mm broad at the base, ovate; stamens with filaments 7-9 mm long, anthers 10-12 mm long, slightly exserted, connective appendage ca. 0.5 m long; stigmas connate, 2-3 mm long. Fruit 8-14 mm long, 4-8 mm diam., cylindrical to oblong-ellipsoid, red becoming black, cov- ered with small (ca 0.5 mm) crooked hairs, persisting sepals to 6 mm long and 2 mm broad; seeds ca. 1 mm long.
Shrubs of Caribbean lowland evergreen forest formations, from near sea level to 200 m elevation. Flowering in February, April, and July-October in Central America. The species ranges along the Caribbean lowlands, from Tabasco, Mexico, to Bocas del Toro, Panama.
Hamelia rovirosae is recognized by the unusual crooked multicellular hairs, corollas narrowly tu- bular at anthesis, puberulent fruit with prominent calyx lobes, and restriction to the Caribbean low- lands. This species is frequently confused with H. patens.
Hamelia xerocarpa Kuntze, Rev. gen. pi. 1: 284. 1 89 1 . H. costaricensis Standl., Contr. U.S. Natl. Mus. 20: 207. 1919. H. panamensis Standl., loc. cit. 208. 1919. H. rowlei Standl., J. Wash. Acad. Sci. 15: 7. 1925. H. storkii Standl., loc. cit. 7. 1925. Figure 42.
Shrubs or small trees to 5 m tall, leafy branchlets 1.5- 5 mm thick, with 4 longitudinal ridges and quadrangular in cross-section, glabrous to sparsely pilose with stiff erect hairs to 0.9 mm long; stipules 6-1 3(-l 7) mm long, cuspidate and often with 2 small lateral teeth, glabrate or pubescent along the edge, drying black. Leaves usually 3 or 4/node, petioles (10-15)-85 mm long, about 1 mm broad, glabrous to densely pubescent; leaf blades 8.5- 1 7(-37?) cm long, 3.5-9(-l 4?) cm broad, ovate to broad- ly elliptic-oblong or ovate-rotund, apex acute to acu- minate, base obtuse to cuneate and decurrent on petiole, leaves drying thin-chartaceous, glabrate above, usually minutely papillate-puberulent or with straight or curved hairs 0.2-0.9 mm long beneath, 2° veins (7-)9-13(-18)/ side and loop-connected near the margin, 3° veins often subparallel, domatia present or absent. Inflorescences terminal on short lateral branches (and apparently ax- illary), 5-15 cm long and equally wide, peduncles 1- 2.5(-5) cm long, the dichasia to 6(-l 5) cm long and with 3-9(-26) flowers along 1 side, glabrous or with yellowish
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hairs to 1 mm long, bracts 0.6-0.8 mm long, the secund flowers sessile or subsessile. Flowers with hypanthium 2-3 mm long, glabrous to pilose, calyx lobes 0.5-2 mm long, ovate, glabrous to pilose; corolla funnelform (but narrowly tubular until anthesis), yellow, tube 1 8-36 mm long, 4-7 mm diam., glabrous to densely puberulent with hairs 0.2-0.4 mm long, lobes 2-5(-8) mm long, 2-5 mm broad at the base, ovate and acute; stamens with fila- ments 8-16 mm long, anthers 10-16 mm long, connec- tive little ( 1 mm) extended distally; style ca. 20 mm long, stigmas 5, to 5 mm long. Fruits 11-14 mm long and 3- 5 mm diam., oblong to ovoid-oblong; seeds 0.5-1 mm long.
Plants of evergreen lowland formations and gal- lery forests in deciduous areas, from near sea level to 800 m elevation. Flowering in May-December (throughout the year in Panama). The species ranges from Nicaragua to northern Colombia.
Hamelia xerocarpa is recognized by the broader corolla tube (at anthesis) with longer corolla lobes, the unusual yellowish puberulence (when present), the lowland evergreen habitat, and the leaves usu- ally with many secondary veins and often with long petioles. Unfortunately, few herbarium col- lections exhibit the broader funnelform corollas in anthesis, and it is easy to misidentify this spe- cies. This species may be difficult to separate from H. macrantha. The breeding biology was studied by Bawa and Beach (1983).
Hamelia xerocarpa variety xerocarpa is distin- guished by the conspicuous hairs on many parts of the plant and the leaves with 11-18 pairs of secondary veins. Variety costaricensis (Standl.) Elias is recognized by the lack of pubescence and leaves with 9-12 pairs of secondary veins.
1 1 illia Jacquin
REFERENCE— C. M. Taylor, Revision of Hillia subg. Ravnia (Rubiaceae: Cinchonoideae). Sel- byana 11: 26-34. 1989.
Shrubs, small trees, or lianas, epiphytic or less often terrestrial, branchlets thick and terete, glabrous; stipules interpetiolar and intrapetiolar but splitting apart along the edges. Ungulate and blunt at apex, caducous. Leaves opposite, equal or unequal at each node, subsessile to short-petiolate; leaf blades elliptic to obovate, entire, decurrent on the petiole, semisucculent and drying co- riaceous, without domatia. Inflorescences of solitary ter- minal flowers (or 3-flowered dichasia in H. thflora), bracts reduced or absent (the flowers at first enclosed within the large untied stipules), pedicels short or absent. Flow- ers bisexual, monomorphic, often large, glabrous exter- nally, hypanthium continuous with the pedicel, calyx tube often absent, calyx lobes 2-5 and distant (or none), sometimes with a secondary smaller set of calyx lobes alternate with the larger lobes; corolla salverform to fun- nelform or tubular, white to yellowish, pinkish, orange, or red, semisucculent, corolla lobes (3-)5-7(-9), con- volute in bud, becoming reflexed; stamens (4-)5-7, fil- aments very short and inserted below the throat, anthers basifixed, elongate-linear, obtuse at each end, included (except in H. longifilamentosa); ovary 2-locular, ovules many and ascending in each locule on scptal placentas, style as long as the corolla tubes, stigmas subcapitate or 2. Fruits woody capsules, narrowly cylindrical to very narrowly oblong, truncated distally, dark brown and smooth, dehiscing septicidally and basipetally into 2 flat- tened valves; seeds many and imbricated, rhombic and flattened, with a minute circumferential wing, appen- daged at the base and with a tuft of hairs at the distal apex.
Hillia is a genus of about 20 species, ranging from southern Mexico to Brazil and Peru. The genus is recognized by the larger flowers with long tubes, many ascending imbricated ovules, long tu- bular ("cigar-shaped") capsules, and flattened winged seeds with a tuft of hairs at one end. The more colorful flowers of subgenus Ravnia (see fol- lowing key) appear to be an adaptation to bird pollination. Specimens lacking flowers or fruit can be very difficult to identify to species. In addition, one group of our species may be part of a poly- morphic complex; see the discussion under H. maxonii. Some of our species of Hillia are very similar to Cosmibuena, but that genus has seeds lacking the tufted hairs at one end.
Key to the Species of Hillia
la. Corolla pinkish to red or orange, funnelform or tubular with the central part of the tube inflated
and narrowed at both ends, flowers not scented; subgenus Ravnia .
Ib. Corolla greenish or white, long-tubular with rotate lobes or salverform, corolla tube expanded only at the corolla lobes, flowers often sweet-scented; subgenus Hillia . 4
2a. Corolla tube rose red and inflated in the middle, 3-5 cm long, flowers often in groups of 3; a
commonly collected species "• triflora
2b. Corolla tube yellow-orange to peach or rose, funnelform and widest at the mouth, 4-6.5 cm
long, flowers solitary; rarely collected species .... 3a. Corolla lobes 6-12 mm long; free portion of the filaments ca. 1 mm long H. allenii
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
161
3b. Corolla lobes 13-17 mm long; free portion of the filaments 9-10 mm long
H. longifilamentosa
4a. Plants of lower elevation forest, not known from above 300 m in Costa Rica; larger leaf blades
more than 10 cm long and 5 cm broad 5
4b. Plants of higher elevations, rarely collected below 600 m; larger leaf blades rarely more than 1 0 cm
long or 5 cm broad 6
5a. Leaf blades with thin texture, the 5-6 pairs of 2° veins arising at ca. 60° angles from the midvein; corolla tubes 8-10 cm long H. macrophylla
5b. Leaf blades thick-textured, the 4-5 pairs of 2° veins arising at ca. 30° angles from the midvein;
corolla tubes 4-5 cm long H. grayumii
6a. Leaf blades 4-10 cm long, usually tapering gradually to a bluntly acute apex, broadest at the middle
or below; corolla tubes 4-5 cm long; seed hairs ca. 1 7 mm long H. loranthoides
6b. Leaf blades 0.7-3.7 cm long, usually bluntly obtuse to rounded at the apex; corolla tubes rarely
exceeding 4 cm in length; seed hairs ca. 10 mm long 7
7a. Leaf blades 12-37 mm long, stipules 6-16 mm long; corolla lobes suborbicular, corolla tubes 24-
42 mm long; seeds ca. 3 mm long; 1400-2400 m elevation H. maxonii
7b. Leaf blades 7-14 mm long, stipules 3-7 mm long; corolla lobes broadly to narrowly ovate, corolla
tubes 15-35 mm long; seeds ca. 2 mm long; 600-1600 m elevation H. panamensis
Hillia allenii C. M. Taylor, Selbyana 1 1: 32. 1989. Ravnia panamensis Steyerm., Ceiba 3: 22. 1 952, not Hillia panamensis Standl.
Epiphytic shrubs to 1.5 m tall, leafy stems 2.5-7 mm thick, glabrous, terete; stipules 10-12 mm long, Ungulate, quickly caducous. Leaves with short (1—4 mm), thick (2- 3 mm) petioles, glabrous and drying dark; leaf blades 4-
1 1 cm long, 1.5-5 cm broad, elliptic to narrowly elliptic- oblong, apex slender acuminate, base cuneate or slightly rounded, drying subcoriaceous, dark brown above, gla- brous above and below, 2° veins 5-9/side, usually ob- scure. Inflorescences of solitary terminal flowers borne on short (3—4 mm) thick (2 mm) glabrous pedicels drying black. Flowers glabrous externally, hypanthium 4-7 mm long and 3.5 mm thick, calyx tube not developed, calyx lobes 6, 6-14 mm long, 2-3 mm broad, narrowly spat- ulate-oblong; corolla funnelform, pale red and pale yel- low to salmon-pink, tube 25-40 mm long, lobes 6, 6-
1 2 mm long and 1 0 mm broad at the base, bluntly acute; anthers 5-7 mm long on filaments ca. 1 mm long. Fruits not seen.
Plants of montane cloud forest formations at ca. 1000 m elevation. Flowering in June and Septem- ber. This species known only from the Cordillera de Tilaran and in western Panama.
Hillia allenii is recognized by the brightly col- ored funnelform corolla and the short filaments. Fruiting material may be very difficult to distin- guish from that of H. longifilamentosa.
Hillia grayumii C. M. Taylor, Selbyana 12: 137. 1991. Figure 28.
Epiphytic shrubs, ca. 1 m tall, leafy stems 4-7 mm thick, slightly quadrangular, glabrous, brownish, mi-
nutely grooved; stipules ca. 40 mm long, 6-8 mm broad, lanceolate, caducous. Leaves isophyllous, usually decus- sate, petioles 6-20 mm long, 2-3 mm thick, articulated at the stem; leaf blades 9-19 cm long, 3-6.5 cm broad, elliptic to elliptic-oblong, apex acuminate with tip 1-2 cm long, base obtuse or acute, drying stiffly chartaceous or subcoriaceous and dark brown above, glabrous above and below, 2° veins 4-6/side and strongly ascending (ca. 30°), obscure beneath. Inflorescences of solitary terminal flowers, subtending stipules caducous, peduncles 2-3 mm long, bracts 1-3 mm long, triangular, acute. Flowers gla- brous, hypanthium 7-8 mm long, cylindrical, calyx limb to 0.5 mm long, truncate or slightly lobed; corolla tu- bular-funnelform, bright pale green to yellow, tube 43- 50 mm long, lobes 6, 8-9 mm long, triangular, obtuse to rounded; filaments ca. 10 mm long, anthers 6, ca. 9 mm long. Fruits 10-12 cm long and 8-14 mm broad, drying dark brown, stipe ca. 3 mm long; seeds 3 mm long, 0.5 mm broad.
Plants of lowland rain forest and swamp forest formations, collected from near sea level to 600 m elevation. Flowering in May-June; fruiting in March and May-June. The species is known only from the Caribbean lowlands of northern and cen- tral Costa Rica.
Hillia grayumii is distinguished from its con- geners by the lowland habitat, yellowish funnel- form flowers, and large fruit. Among Costa Rican species, it is similar to H. macrophylla, but that species grows in cloud forests and has thin-tex- tured leaves and tubular flowers.
Hillia longifilamentosa (Steyerm.) C. M. Taylor, Selbyana 1 1: 32. 1989. Ravnia longifilamentosa Steyerm., Ceiba 3: 21. 1952.
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FIELDIANA: BOTANY
Epiphytic or clambering shrubs to 6 m tall, leafy stems 2-7 mm thick, glabrous, smooth, brown to gray; stipules 22-37 mm long, elliptic to oblanceolate, caducous. Leaves with petioles 2-8 mm long, thick, glabrous; leaf blades 6-16 cm long, 2-7 cm broad, elliptic or elliptic-oblong, apex acute or slightly acuminate, base obtuse to cuneate, drying subcoriaceous, grayish green, glabrous above and below, 2° veins 4-6/side, strongly ascending. Inflores- cences of solitary terminal flowers, pedicles ca. 1 mm long, bracts 2-3 mm long, triangular, acute. Flowers with hypanthium 4-10 mm long, obconic to ellipsoid, calyx lobes 6, 6-14 mm long, narrowly triangular to Ungulate or oblanceolate; corolla tubular-rotate, orange-red to salmon-pink, or white marked with pink, tube 32-43 mm long, ca. 3 mm diam., lobes 6, 13-17 mm long, triangular to Ungulate; stamens 4-6, anthers ca. 5 mm long, well exserted, dark green. Fruits 10 cm long.
Plants of montane cloud forest formations, 1 1 00- 1700 m elevation. Flowering in April, July-Au- gust, and November. This species is known only from a few collections, ranging from Zarcero, Ala- juela, to Chiriqui, Panama.
Hillia longifilamentosa is distinguished by its solitary terminal flowers with anthers extended 8- 10 mm beyond the throat of the tube. Compare H. allenii.
Hillia loranthoides Standl., J. Wash. Acad. Sci. 18: 165. 1928. Figure 27.
Epiphytic shrubs, 0.4-1.5 m tall but with vining branches, leafy stems 3-8 mm thick, glabrous, grayish, terete or quadrangular; stipules 18-22 mm long (to 35 mm beneath the flowers), 6-10 mm broad, oblong-ob- ovate, obtuse or rounded distally. Leaves closely clus- tered or distant, decussate, petioles 4-8(-15) mm long, 1.5-2.8 mm thick; leaf blades (3-)4-10 cm long, (1 .3-)2- 4.5 cm broad, elliptic to elliptic-oblong, ovate or ovate- lanceolate, apex bluntly acute or obtuse, base obtuse to cuneate, drying coriaceous, dark grayish, 2° veins 4-57 side, strongly ascending or obscure. Inflorescences of sol- itary terminal sessile flowers, subtended by a pair of enlarged (8-25 mm) oblong-obovate bract-like reddish stipules, 1-2 smaller (1-4 mm) pairs of triangular or Ungulate bracteoles often present at the base of the short pedicel. Flowers 6-7 cm long, hypanthium ca. 4 mm long, calyx tube minute, calyx lobes 4, 8-10 mm long, 1-2 mm broad, linear-Ungulate; corolla tubular with ro- tate lobes, white or cream, carnose, tube 40-60 mm long, 2-5 mm diam., lobes 4, 10-25 mm long, 8-16 mm broad, rounded distally. Fruits 3-8 cm long, 7-10 mm diam.; seeds ca. 2.5 mm long and 0.7 mm thick, distal hairs 6- 17 mm long.
Plants of evergreen montane forests, from (300-)700 to 1400 m elevation. Flowering in Feb- ruary-May; fruiting in January and May. In Costa Rica, this species is known only from the Cordil- lera de Tilaran, near San Ramon, and El Retiro
(Cartago) and from a single lower elevation col- lection on the Osa Peninsula. The species is also known from southern Mexico.
Hillia loranthoides is distinguished by its thick leaves, usually tapering gradually to the apex, larg- er flowers with slender tubes, and seeds with longer bristles. Its distribution is unusual and may be an artifact of the difficulty of collecting epiphytes.
Hillia macrophylla Standl., Publ. Field Columb. Mus., Hot. Ser. 7: 201. 1931. Figure 28.
Epiphytic shrubs or vines, 3-7 m tall, leafy branchlets 3-7 mm thick, glabrous, pale brown and lenticelate; stip- ules 10-35 mm long, 12-25 mm broad, narrowly lan- ceolate (in Costa Rica) to oval-oblong, the basal sheath 1-2 mm long, caducous. Leaves decussate, petioles 6- 20 mm long, 2-3.5 mm thick, terete, clearly differenti- ated from the lamina base; leaf blades 9-2 1 cm long, 6- 10 cm broad, ovate-elliptic to elliptic-oblong or elliptic, apex acuminate (acute), base obtuse, drying membra- naceous to chartaceous, dark brown above, glabrous above and below, major veins with a rugose texture. 2° veins 6-9/side arising at ca. 60° angles and loop-con- nected near the margin. Inflorescences of large solitary sessile terminal flowers, subtended by elongated (20-50 x 5-24 mm) caducous stipules, bracteoles usually ab- sent. Flowers with hypanthium 6-7 mm long, 3-3.5 mm diam., cylindrical, calyx tube 0-0.5 mm long, subentire; corolla tubular-salverform, white, tube 5.5-1 1 cm long, 2-4.5 mm diam., lobes 5 or 6, 2.5-5 cm long. ca. 8 mm broad at the base and narrow (4 mm) distally, linear- lanceolate to narrowly triangular. Fruits 7-12 cm long, the opened valves becoming 14-20 mm broad, rounded at the base, acute at apex, sessile; seeds 1.5-4 mm long with distal hairs 14-18 mm long.
Plants of moist cloud forests from 800 to 1 800 m elevation. Flowering in March-June; a single fruiting collection was made in December. This species is known only from near Monteverde, Ca- taratas de San Ramon, and above the Rio Gato, Cartago, in Costa Rica. It is also found in Colom- bia, Ecuador, and Peru.
Hillia macrophylla is unique among our species of Hillia because of the larger thin-textured leaves, more numerous ascending secondary veins, and long-tubular flowers with corolla lobes that have long (3-4 cm), narrow tips. It is a rarely collected species. South American collections appear to have larger (3-4 mm) seeds and larger stipules sub- tending the flowers.
Hillia maxonii Standl., J. Wash. Acad. Sci. 18: 163. 1 928. H.palmana Standl. Joe. cit. 18: 164. 1928. H. hathewayi Fosberg, Sida 2: 387. 1966. Figure 27.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
163
Epiphytic shrubs, 0.7-2. 5(-5) m tall, often pendant to 5 mm long, leafy stems 1-5 mm thick, dark or pale grayish, glabrous, older nodes articulate with transverse ridges; stipules 6-16(-32) mm long, 2-6 mm broad, ob- long-obovate to obovate, rounded, larger and bract-like beneath the flowers. Leaves decussate and often crowded on short (3-15 mm) internodes, petioles 2-6(-15) mm long; leaf blades (12-)18-37(-60) mm long, 6-20(-30) mm broad, obovate to oblong-obovate or ovate-elliptic, apex bluntly obtuse to rounded, base obtuse to cuneate, decurrent on the petiole, drying subcoriaceous and dark grayish or brownish, glabrous above and below, 2° veins 3-4/side, strongly ascending or obscure. Inflorescences of solitary, terminal, sessile flowers subtended by a pair of enlarged bract-like stipules rounded at apex. Flowers glabrous externally, fragrant, hypanthium ca. 2.5 mm long and 1 .8 mm thick, calyx tube very short, calyx lobes absent or 4-6 mm long, 0.5-1.8 mm broad; corolla tu- bular with rotate lobes, slightly fleshy, tube 24-42 mm long, 1.4-4 mm diam., greenish or white, lobes 4, 10- 14(-22) mm long and usually equally broad or broader than long, suborbicular and rounded distally, bright white or yellowish white. Fruits (22-)30-60 mm long, 5-9 mm thick, the valves to 8 mm broad when opened and ex- panded; seeds 3-4 mm long, 0.5-1 mm diam., distal hairs 6-13 mm long.
Plants of evergreen montane forest formations from 500 to 2400 m elevation. Flowering in April- September and December; fruiting in September and November-March. This species has been col- lected near Managua, Nicaragua, in the Cordillera de Tilaran, Cordillera Central, in the western part of the Cordillera de Talamanca, and in Chiriqui and Veraguas, Panama.
Hillia maxonii is recognized by its usually epi- phytic habit, smaller stiff leaves, bract-like stipules subtending the flowers, and tubular corollas with broadly rounded lobes. Smaller-leaved specimens of this species may represent intermediates with H. panamensis. Larger-leaved specimens resem- ble H. tetrandra Sw. of the West Indies and north- ern Central America. However, H. tetrandra has larger (5-1 1 x 2.5-6 cm) leaves that are more often obovate, quite unlike those of southern Cen- tral America. All three taxa exhibit a wide range of variation, and it is possible that they are ele- ments of a single polymorphic species. (A similar problem is found in the epiphytic species of Psy- chotria; see the discussion under P. guadalupen- sis.)
Hillia panamensis Standl., N. Amer. Fl. 32: 117. 1921. H. chiapensis Standl., J. Wash. Acad. Sci. 16: 16. 1926. Figure 27.
Epiphytic shrubs or small treelets to 4 m tall, leafy stems 1-2.7 mm thick, glabrous and grayish, often de-
veloping opposite longitudinal sulci and expanded nodes; stipules 3-7 mm long, 1-2 mm broad, oblong to narrowly obovate and rounded distally, translucent. Leaves de- cussate and usually closely crowded on short (0.5-5 mm) internodes, petioles 1-4 m long; leaf blades 7-12(-14) mm long, 4-8(-10) mm broad, elliptic to elliptic-oblong or obovate, apex acute to bluntly obtuse or rounded, base cuneate to obtuse, drying stiffly chartaceous to sub- coriaceous, grayish to very dark, margins often becoming slightly revolute, glabrous above and below, 2° veins 2- 3/side, strongly ascending or obscure. Inflorescences of solitary terminal flowers, the distal stipule pair slightly expanded (6 x 2.5 mm) and bract-like smaller (1-2 mm) rounded bracts sometimes present at the base of the sessile flowers. Flowers glabrous externally, hypanthium ca. 1.5 mm long, 1 mm thick, tubular, calyx lobes to 1 1 mm long, 0.3-0.6 mm broad, linear, caducous; corolla tubular with rotate lobes, white, slightly succulent, tube 15-30(-38) mm long, 0.9-1.5 mm diam., lobes 4, 5-7 mm long, 3.5-6 mm broad, ovate to lanceolate, bluntly obtuse. Fruits 20-42 mm long, opened expanded valves 3-4 mm wide; seeds ca. 2 mm long and 0.6 mm thick, distal hairs 10 mm long.
Plants of evergreen montane forest formations from 600 to 1 600 m elevation. Flowering in May- August; fruiting in December. This species is found in southern Mexico-Guatemala, in the Cordilleras de Guanacaste and Tilaran in Costa Rica, and in western and central Panama.
Hillia panamensis is distinguished by its epi- phytic habit, very small stiffclosely crowded leaves, fragrant flowers with long slender corolla tubes, and rotate lobes usually narrower than long. It has been called jasmin del volcdn. This species may not be specifically distinct from the very similar, and partly sympatric, H. maxonii. But the differ- ences used in the keys do seem to separate a great majority of specimens. See the discussion under H. maxonii.
Hillia triflora (Oersted) C. M. Taylor, Selbyana 1 1: 30. 1989. Ravnia triflora Oersted, Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1852: 49. 1853. Ravnia pittien 'Standl., N. Amer. Fl. 32: 1 14. \92l.LagenanthusparviflorusE\van, Mutisia 4: 5. 1952. H. triflora var. pittieri (Standl.) C M. Taylor, Selbyana 11:31. 1989. Figure 27.
Epiphytic (rarely terrestrial) shrubs, 0.5-1 .5(-3) m tall (branches to 2 m long and pendulous), leafy stems 1.8- 5 m thick, glabrous; stipules 15^43 mm long, 4-10 mm broad, largest beneath the inflorescences, elliptic and acute, glabrous and caducous. Leaves usually decussate, isophyllous or anisophyllous at a node, petioles 3-7(-l 2) mm long, 1.5-2 mm thick, poorly differentiated from the base; leaf blades 5-1 3(-l 9) cm long, 1 .5-4.5 cm broad, narrowly elliptic-oblong to oblanceolate, elliptic-oblong
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FIELDIANA: BOTANY
or oblong (linear-lanceolate in variety pittieri), apex ta- pering gradually and narrowly acuminate, tapering grad- ually to the acute or cuneate base, drying coriaceous, dark green or grayish green above, 2° veins 3-5/side. Inflorescences of 1 or 3 terminal flowers borne on a very short (0.5-2 mm) thick peduncle, bracts ca. 3 mm long or absent, pedicels 1-6 mm long but difficult to distin- guish from the hypanthium. Flowers glabrous externally, hypanthium ca. 5 mm long, 1.2-1.8 mm diam. calyx lobes 6 (5, 7), 4-14 mm long, 0.5-1 mm broad, linear- oblong; corolla red, tubular with relatively small lobes, tube 40-65 mm long, 1.5-10 mm broad, lobes 6 (5, 7), 3-4 mm long, to 5 mm broad at the base; stamens 5-6, anthers 3-5 mm long, partially exserted. Fruits 5-10 cm long, ca. 8 mm diam., valves becoming up to 12 mm broad; seeds 0.8-2.5 mm long, distal hairs 15-30 mm long.
Plants of montane and premontane evergreen forest formations from (100-)800 to 2400 m ele-
vation. The lowest elevation collections are from La Selva; this species has not been collected from below 1000 m on the Pacific slope. Flowering in all months, but most commonly February-Au- gust; fruiting in October-February. The species ranges from southern Mexico to northwestern Co- lombia.
Hillia triflora is distinguish by its epiphytic hab- it, the tubular reddish flowers usually in terminal triads, long narrow capsules, and seeds with a dis- tal tuft of hairs. There are few fruiting collections. There may be some local differentiation between the two varieties; at Monteverde, variety pittieri is found at slightly lower elevations than is variety triflora. The following key distinguishes the two varieties.
la. Leaves at each node approximately equal in size at the same node on flowering shoots; mature corolla tube not or only slightly inflated, up to 1.6 times broader in the broadest portion as at the apex H. triflora var. triflora
Ib. Leaves at each node strongly differing in size on flowering shoots, larger leaves ca. 1.5 times longer than the other leaf of the same node; mature corolla tube strongly inflated, ca. 1 .8 times broader in center than at the apex H. triflora var. pittieri
Hippotis Ruiz Lopez & Pavon
Trees or shrubs, puberulent (in ours) or glabrous; stip- ules interpetiolar, triangular to obovate, large and ca- ducous. Leaves opposite, decussate, petiolate; leaf blades usually thin, pinnately veined, without domatia, with the minor venation parallel (lineolate) within the areolae formed by the 3° and 4° veins. Inflorescences axillary to leaves or undeveloped leaves and axillary in fruit, short cymose and 2-3-flowered, capitate or of solitary flowers. Flowers bisexual, monomorphic, radially symmetrical or bilaterally symmetrical by curvature of the corolla, usually large, sericeous externally, calyx bilobate or spa- thaceous; corolla tubular or funnelform, white, rose, red, or pale orange, corolla tube straight or curved, corolla lobes 5, short and truncated or emarginate, plicate-val- vate in bud; stamens 5, filaments usually unequal, borne at the middle or lower half of the tube, anthers included; ovary 2-locular, placentas peltate from the center of the septum, with many horizontal ovules in each locule, stigmas 2, subcapitate. Fruits baccate, ovoid to ellipsoid or globose; seeds many, small, angular.
A genus of around 12 species in northern South America, Ecuador, and Peru; 1 species reaches northernmost Costa Rica. The larger thin hairy leaves, unusual lineolate minor venation, few larg- er pseudoterminal flowers, spathaceous calyx, and hairy fleshy fruit with many small seeds help to distinguish our species.
Hippotis albiflora Karst., Fl. Colomb. 1: 33, pi. 1 7. 1 858. Duroia panamensis Dwyer, Ann. Mis- souri Bot. Gard. 55: 38. 1968. Figure 25.
Small trees, 3-17 m tall, leafy stems 2.5-8 mm thick, densely pilose with yellowish hairs 1-3 mm long, terete or slightly quadrangular, older stems glabrescent and pale grayish; stipules 15-30 mm long, 5-12 mm broad, nar- rowly ovate or lanceolate, with long yellowish hairs ex- ternally, leaves with petioles 10-28(-35) mm long, 1.5- 2.5 mm thick, densely pilose with erect or ascending hairs 1-2 mm long; leaf blades 14-30(-35) cm long, 6- 14(-18) cm broad, obovate to oblong-obovate, apex ta- pering gradually or abruptly and acuminate or caudate- acuminate, tip 1-2 cm long, base obtuse to rounded and subtruncate, drying thin-chartaceous to chartaceous, densely pilose on both surfaces with thin straight or curved hairs 0.7-2 mm long, 2° veins 7-10/side, 3° veins often subparallel, the smallest veins distinctly parallel (lineo- late) in small groups. Inflorescences of 1-3 flowers in the axils of the distal leaves but often appearing terminal, peduncles absent or very short, bracts 3-5 mm long, ovate, pedicels 3-8 mm long, 1-2 mm thick and densely yellowish velutinous. Flowers 5-6 cm long, hypanthium and calyx 3.3-3.8 cm long, ca. 8 mm broad, densely pilose, spathe-like and split ca. 10 mm down 1 side; corolla ca. 5 cm long and 2 cm broad distally, funnel- form, white to cream, pilose externally, tube 35-45 mm long, greenish white, straight or slightly curved, lobes 5- 8 mm long and 10 mm broad, obtuse. Fruits 3-4 cm long (not including the persisting calyx), 1-2 cm diam.,
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
165
ellipsoid and hirsute, becoming purple; seeds 2 mm long, foveolate.
Plants of rain forest formations in the Caribbean lowlands, from 10 to 400 m elevation (to 1600 m in Venezuela). Flowering in February, April, and July-October; fruiting in May-September. This species has been collected near Rio Colorado in northern Costa Rica, from La Selva and the Hitoy Cerere reserve. It is also known from the province of Bocas del Toro, in Panama, and from Colombia and Venezuela.
Hippotis albiflora is recognized by the conspic- uous pubescence, larger thin leaves with lineolate minor venation, few large pseudoterminal flowers, spathaceous calyx, and larger, hirsute, many-seed- ed fruit. Our material may differ in some details from that described from Venezuela; it could be that Costa Rican populations are worthy of sub- specific recognition. Compare this species with Duroia costaricensis.
Hoffmannia Swart/
Herbs, herbaceous subshrubs, or slender shrubs, main vertical stems usually with few or no lateral branches, glabrous or pubescent, terete or quadrangular, internodes often hollow when dried; stipules interpetiolar, usually small and triangular, often succulent and divergent, ca- ducous. Leaves opposite and decussate or rarely verti- cillate, equal at the node or sometimes slightly unequal, petiolate or occasionally subsessile, petioles with inflated chambers (ant vesicles) in a few species; leaf blades usu- ally drying membranaceous to chartaceous (rarely sub- coriaceous), entire, often decurrent on the petiole, pin- nately veined, often with conspicuous raphides on the dried leaf surfaces, domatia rarely present. Inflores- cences axillary, sessile or pedunculate, with few to many flowers, usually cymose or dichasial and often with heli- coid branches, fasciculate to capitate in a few species, flowers usually pedicellate, bracts and bracteoles rarely
present. Flowers bisexual and monomorphic, glabrous or pubescent externally, hypanthium often with longi- tudinal ribs, calyx tube usually very short, calyx lobes 4(-5), usually short and often triangular, persisting; co- rolla salverform to funnelform or rotate, white, yellow, orange, or rose to deep red or purple, corolla tube short and usually narrow, glabrous on the interior, corolla lobes 4(-5), imbricate along the edges or apparently valvate, apex usually acute; stamens 4 (3-5), borne within the tube of the corolla, filaments very short, anthers linear to narrowly oblong, usually white; ovary 2- (3, 4) locular, each locule with many ovules borne longitudinally on bilamellate axile placentas, style slender, stigmas bilo- bate or clavate. Fruits baccate, usually small, 2- (3, 4) locular, often becoming enlarged and spongy at maturity, with many minute multiseriate horizontal seeds; seeds angulate, their surfaces often reticulate to foveolate.
A genus of about 1 00 species, ranging from Mex- ico and the West Indies to South America. A ma- jority of the species are found in Mexico and north- ern Central America. The usually single-stemmed habit, presence of raphides in many parts, small triangular stipules (often caducous), axillary inflo- rescences, four-parted flowers, and many-seeded fleshy fruits help to distinguish the species of Hoff- mannia from other Rubiaceae in Costa Rica. Most of the Costa Rican species are restricted to areas with very high rainfall (none are found in decid- uous vegetation), and relatively few species grow below 800 m elevation.
Hoffmannia is probably Costa Rica's taxonom- ically most difficult genus of Rubiaceae. These semisucculent subshrubs do not dry well, and the flowers, often in small dense clusters, are also not well preserved. In addition, there appears to be great variation from plant to plant within many species. The genus is related to Hamelia and Dep- pea; it is currently being studied by Dr. John Dwyer (MO), and the review presented here should be con- sidered no more than tentative.
Key to the Species of Hoffmannia
la. Petioles with lateral longitudinal inflated vesicles; plants to 1.2 m tall, surfaces with longer (1.5 mm) multicellular hairs H. vesiculifera
Ib. Petioles without lateral inflated structures; plants 0.5-3 m tall, glabrous to villous with multicellular hairs 2
2a. Inflorescences sessile and capitate in the leaf axils in early stages (rarely becoming pedunculate or branched in later stages), often verticillate at the nodes 3
2b. Inflorescences sessile and fasciculate to pedunculate and paniculate or cymose, neither densely
capitate nor verticillate 4
3a. Leaf blades neither bullate nor areolate, to 30(— 40) cm long, glabrous or pubescent; corolla ca. 7 mm long; commonly collected plants H. congesta
166
FIELDIANA: BOTANY
3b. Leaf blades bullate or areolate, to 20(-25) cm long, pubescent; corolla ca. 4.5 mm long; rare
H. areolata
4a. Leaves usually 3/node, leaf blades cuneate at the base with long-decurrent lateral margins on the petiole and these slightly expanded near the stem, lateral margins of the petiole 2-4 mm broad [leaf blades with 1 1-18 pairs of 2° veins]; Vara Blanca to the upper Rio Grande de Orosi, 1400-
1600 m elevation H. amplexifolia
4b. Leaves opposite (rarely with 3 leaves per node), leaf blades obtuse to cuneate at the base, if long- decurrent on the petiole never forming a lateral margin 2-4 mm wide along the petiole and not
expanded near the base; species widely distributed, 10-2300 m elevation 5
5a. Plants of Cocos Island 6
5b. Plants of mainland Central America 7
6a. Petioles 2-5 cm long; corolla glabrous on the exterior H. piratarum
6b. Petioles 6-8 cm long; corolla puberulent on the exterior H. nesiota
7a. Young stems glabrous or subglabrous, or with very small (0.2 mm) hairs in longitudinal lines along
the young stern 8
7b. Young stems densely pubescent with crooked multicellular hairs, the hairs varying 0.3-2 mm long,
older stems pubescent or glabrescent 19
8a. Inflorescences short, the 1° peduncle and rachis usually less than 3 cm long (if plants grow below 600 m elevation and have petioles 2-6 cm long and leaves 4-15 cm broad, go to
dichotomy 12) 9
8b. Inflorescences small to large, peduncle and rachis of some inflorescences usually exceeding
4 cm in length 12
9a. Corolla tubes 3-4 mm diam. and 3-4 mm long; leaf blades 15-35 cm long with 9-15
pairs of 2° veins, cuneate basally and long-decurrent on the petiole H. dolae
9b. Corolla tubes 1-3 mm diam. (dried) and 1-6 mm long; leaf blades 5-15 cm long with 5-9 pairs of 2° veins, abruptly narrowed at the base (but leaves to 23 cm long, with 7-
12 pairs of 2° veins, and cuneate long-decurrent bases in H. hamelioides) 10
lOa. Corolla 9-14 mm long, corolla lobes 3-6 mm long; calyx lobes ca. 1.5 mm long, ovary 2-3 mm long; flowers fasciculate (inflorescences rarely pedunculate), usually more than
1 5/node; fruit elongate when dried H. psychotriifolia
lOb. Corolla less than 9 mm long, corolla lobes 1.5-4 mm long; calyx lobes 0.5-1 mm long, ovary 1-2 mm long; flowers fasciculate or borne on slender pedunculate; fruit rounded
when dried 11
1 la. Inflorescences compact, fasciculate or with short stiff peduncles, often less than 2 cm long; leaf blades to 1 5 cm long, rarely tapering gradually at the base and decurrent on
the petiole; a commonly collected species H. longipetiolata
lib. Inflorescences open with thin peduncles and pedicels, usually more than 2 cm long; leaf blades to 23 cm long, usually tapering gradually at the base and decurrent on the
petiole; uncommon plants H. hamelioides
12a. Sepal lobes 2-4 mm long; corolla tube 1-3 mm long, corolla lobes 3-7 mm long; plants not collected from about 700 m elevation [leaf blades usually with an arcuate submarginal vein
and abruptly narrowed at the base] H. liesneriana
12b. Sepal lobes rarely more than 1 .8 mm long; corolla tube usually more than 2 mm long (except in H. laxa), corolla lobes 2-6 mm long; plants rarely found below 800 m elevation (except
H. pallidiflord) 13
1 3a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein; in- florescences with a prominent erect peduncle 4-10 cm long and usually terminated by 3
branches; 1200-1500 m elevation H. davidsoniae
13b. Secondary veins loop-connected only in the distal part of the lamina, not forming an arcuate submarginal vein; inflorescences often pendant, with shorter slender peduncles (400-)800-
2300 m elevation -I4
14a. Corolla tubes 4-8 mm long; inflorescences short to long and with many lateral branches; leaf blades drying stiffly chartaceous and often large, 12-37 cm long with 9-15 pairs of 2° veins; 1000-2200 m elevation • ! 5
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 167
14b. Corolla tubes 1 .5-4 mm long; inflorescences short (to 6 cm) with few lateral branches [corolla tube 1.5—4 mm long]; leaf blades drying membranaceous to thin-chartaceous, 7-20(-25) cm
long with 7-10 pairs of 2° veins; (400-)800-1 500 m elevation 17
1 5a. Leaves usually drying grayish and stiffly chartaceous, petioles 2-10 cm long; leaf blades cuneate and slightly decurrent on the petiole; (corolla tubes 4-8 mm long); fruit rose- red to white; central Costa Rica H. leucocarpa
1 5b. Leaves usually drying dark brown and chartaceous, petioles 2-5 cm long; leaf blades usually cuneate and long-decurrent at the base; fruit red; central Costa Rica to the
Chiriqui Highlands (note: the following 2 species may intergrade) 16
16a. Corolla tubes 6-10 mm long; calyx lobes 1.2-3 mm long; leaf blades often narrowly obovate and long attenuate at the base; central Costa Rica to Panama, 1000-2500 m
elevation H. arborescens
16b. Corolla tubes 3-7 mm long; calyx lobes 0.5-1.5 mm long; leaf blades not usually narrowly obovate and long attenuate at the base; westernmost Costa Rica and the
Chiriqui Highlands, 1 100-1 700 m elevation H. pittieri
1 7a. Inflorescences usually unbranched (sometimes bifurcate) [drying yellowish and not very slen- der, probably erect; pedicels 0-3 mm long and ca. 0.5 mm thick (dried); corolla tube 2.5—4 mm long; 400-1300 m elevation on the Caribbean slope] H. pallidiflora
1 7b. Inflorescences usually branched 18
18a. Inflorescences with very thin (0.2-0.3 mm) branches that usually dry black, pendulous, flowering portion 2—4 cm long, pedicels 4-9 mm long and 0.2-0.3 mm thick (dried); 1000-
1 500 m elevation H. laxa
18b. Inflorescences with thicker branches that do not dry black, flowering portion 1-2 cm long,
pedicels 2-6 mm long, not thin and black; 0-200 m elevation, Osa Peninsula
H. hammelii
1 9a. (from 7b) Leaf blades rounded to bluntly obtuse at the apex, often bullate, many 4° veins parallel (sublineolate but difficult to see); inflorescences with few (3-7) subsessile flowers on long (3-7 cm)
peduncles; plants to 40 cm tall [200-1200 m elevation] H. bullata
1 9b. Leaf blades acute to acuminate (rarely obtuse) at the apex, never bullate, 4° veins partly parallel only in H. aeruginosa; inflorescences with short (0-2 cm) peduncles or with more than 7 flowers
if peduncles are long, pedicels 0-2 cm long; plants usually more than 40 cm tall 20
20a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein [inflores- cences subsessile and usually less than 2 cm long] 21
20b. Secondary veins loop-connected only in the distal part of the blade, a submarginal vein absent
(except in the distal third) 22
2 la. Leaf blades cuneate and slightly decurrent at the base, with 9-1 5 pairs of 2° veins; 400-1000
m elevation H. inamoena
21b. Leaf blades acute to obtuse at the base, not usually decurrent on the petiole, with 7-12 pairs
of 2° veins; 1000-1500 m elevation in north-central Costa Rica H. aeruginosa
22a. Plants not found above 900 m elevation; leaf blades usually less than 15 cm long, rarely cuneate
and long-decurrent at the base, with 6-10 pairs of 2° veins H. valerii
22b. Plants not found below 1 000 m elevation; leaf blades often exceeding 1 5 cm in length, usually
gradually cuneate and long-decurrent on the petioles, with 7-12 pairs of 2° veins 23
23a. Inflorescences with thick densely villous peduncles and pedicels; calyx lobes 1.5-3 mm long; corolla
tube 6-8 mm long (lobes 3-4.5 mm long) H. asclepiadea
23b. Inflorescences with slender glabrescent or sparsely puberulent peduncles and pedicels; calyx lobes 0.3-1.5 mm long; corolla tubes 1.5-4 mm long (note: the following 2 species may be synonymous)
24
24a. Inflorescences short but open and few-flowered; corolla lobes mostly 2—4 mm long
H. hamelioides
24b. Inflorescences short but crowded and the flowers clustered around the node; corolla lobes 3-7 mm long H. decurrens
168 FIELDIANA: BOTANY
Hoffmannia aeruginosa Stand!. , Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1313. 1938. Figure 9.
Herbaceous subshrubs or slender shrubs, 0.5-2.5(-4) m tall, with few branches, leafy stems 2-6 mm thick, quadrangular or rounded, densely pilose with reddish brown crooked hairs 0.2-0.7 mm long; stipules 1-2 mm long, 2-3 mm broad at the base, broadly triangular, vil- lous, persisting. Leaves with petioles (10-)20-70 mm long, 0.8-2 mm thick, densely villous, often with petioles of the same node differing in length; leaf blades (6-)9- 20 cm long, (3-)4-10 cm broad, elliptic-oblong to ovate- elliptic or broadly elliptic, apex acuminate or short-acu- minate to acute, base obtuse to acute (not or only slightly decurrent on the petiole); drying thin-chartaceous, gray- ish or grayish brown above, glabrous above, puberulent beneath with short (0.1-0.2 mm) hairs on 2° veins and longer (0.5-2 mm) hairs on the midvein, 2° veins 7-12/ side and evenly loop-connected 2-4 mm from the mar- gin, many 4° veins parallel (forming rectangular areolae). Inflorescences solitary and axillary (2/node), 1-2 cm long with 2-10 flowers, glomerulate, sessile or on short (3 mm) peduncles, unbranched, flowers sessile or subses- sile. Flowers densely covered with curly-crooked hairs 0.3-0.5 mm long, dark reddish brown when dried, hy- panthium 2.5-3 mm long, 2 mm diam. distally, obconic, calyx lobes 1 .3-2.5 mm long, ca. 0.6 mm wide, narrowly oblong; corolla rotate, reddish, tube 1-2 mm long, lobes 4-6 mm long, 2-3 mm broad; anthers ca. 2.5 mm long; style and stigma white. Fruits not seen.
Plants of wet cloud forest formations on the Caribbean slope from (?100-)800 to 1600 m ele- vation. Flowering in April-August. The species is known from north-central Costa Rica, near Ciu- dad Quesada and Zarcero, and from western Pan- ama.
Hoffmannia aeruginosa is recognized by the dense reddish (in life) multicellular hairs on all parts, long-petiolate leaves (with blades scarcely decurrent), small dense inflorescences, and re- stricted range. The petioles often unequal at a node, the subparallel minor venation and arcuate sub- marginal vein are also distinctive features. Com- pare this species with material placed under H. inamoena with less dense pubescence, especially on the petioles.
Hoffmannia attinis Hemsley, Diagn. PI. Nov. 31. 1879.
TYPE— Endres 150, without locality.
Branches terete, puberulent when young. Leaves with petioles ca. 6 mm long; leaf blades 10-12.5 cm long, ovate-oblong, obtusely acuminate, atten- uate at the base, minutely puberulent beneath. In-
florescences umbellate-cymose, fasciculate, with ca. 6 flowers, peduncles 8-16 mm long, slender, pedicels 2-4 mm long. Flowers 6-8 mm long, pu- berulent. Based on Standley (1938), who had only seen the description.
Hoffmannia amplexifolia Standl., J. Wash. Acad. Sci. 15: 8. 1925. Figure 8.
Herbs or subshrubs, l-2(-3) m tall, leafy stems 3-10 mm thick, glabrous or minutely puberulent on the ridges in early stages, with 3 longitudinal ridges; stipules 1-2 mm long and 2 mm broad at the base, triangular, pu- berulent along the edge. Leaves 3 (4)/node, petioles ab- sent or short (3 mm) when the decurrent leaf base is reduced to a lateral ridge, minutely puberulent beneath; leaf blades (11-) 16-37 cm long, 3-13 cm broad, nar- rowly elliptic to oblanceolate or very narrowly elliptic, gradually narrowed to the long-acuminate apex, gradu- ally narrowed to the cuneate and long-decurrent base with a margin 2-4 mm wide along the petiole, often expanded (to 6 mm) and auriculate at the base, leaves drying chartaceous and dark brown to grayish above, glabrescent above, minutely (0.1-0.2 mm) puberulent below, 2° veins 11-1 8(-22)/side, usually loop-connected 1-4 mm from the leaf edge. Inflorescences axillary to leaves, 2-12/node, to 10 cm long, with short (3-15 mm) or long (to 60 mm), thick (1.5-2 mm) peduncles and long (to 7 cm) slender lateral branches from the apex of the peduncle, minutely puberulent, pedicels 0-5 mm long with hairs 0. 1-0.4 mm long. Flowers sparsely to densely puberulent with minute crooked hairs that dry reddish, hypanthium 2-3 mm long, 1.5-2 m diam. distally, tur- binate to obconic, calyx reddish, calyx cup ca. 0.7 mm deep, calyx lobes 1-2.5 mm long and 0.5-1 mm broad, triangular to oblong; corolla rotate, greenish white to pale yellow, tube 1-4 mm long, 1-2 mm diam., lobes 4-9 mm long, 1.5-3 mm broad; anthers 3-5 mm long and 0.7-1.5 mm broad, connivent, yellow. Fruits becoming pink or red, to 8 mm diam. in life (4-5 mm long dried), ellipsoid to subglobose.
Plants of the very wet forests of the Caribbean slope, 800-2000 m elevation. Flowering in March- July; fruiting in October-December and March. The species is known only from central Costa Rica (Vara Blanca de Sarapiqui to Tapanti).
Hoffmannia amplexifolia is recognized by the nodes with three leaves, long-decurrent leaf base forming a leafy lateral margin along the petiole and often slightly expanded (auriculate) at the base (appearing to be amplexicaul), distinctive puber- ulence (but not on the stems), much-branched in- florescences, and broad connivent anthers. Al- meda el al. 6665 (CAS, F) from near Las Alturas, Puntarenas, is tentatively placed here; it has very pubescent inflorescences with corolla tubes 5-6
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
169
mm long. Pressed specimens can be similar to H. subauriculata with two leaves per node.
Hoffmannia arborescens J. D. Smith, Bot. Gaz. 37: 417.1 904. H.josefina Standl., J. Wash. Acad. Sci. 15: 8. 1925. (?//. steinworthii Fosberg, Sida 2: 388. 1966.) Figure 10.
Herbaceous subshrubs, 1.5-2 m tall, leafy stems 2-6 mm thick when dried, glabrescent; stipules ca. 2 mm long. Leaves with petioles 3-8(-25) mm long; leaf blades 7-18 cm long, 2.5-7 cm broad, oblanceolate to narrowly obovate, base acute to attenuate and long-decurrent on petiole, glabrous above and below, drying stiffly char- taceous, often dark reddish brown above, 2° veins 6- 127 side. Inflorescences 1-3 in each leaf axil (2-6/node), 3- 5 cm long, to 4 cm broad, open cymose with many branches, peduncles 3-20 mm long, puberulent with stiff hairs, flowers crowded distally, pedicels 2-6 mm long. Flowers with hypanthium 2-3 mm long, ca. 2 mm diam., calyx lobes 1-3 mm long, 1 mm broad at the base, blunt at apex; corolla funnelform, white to yellow or flushed with red, glabrous or puberulent, tube 6-8(-l 0) mm long, 1-2 mm diam., lobes 3-6 mm long. Fruits ca. 8 x 4.5 mm, red.
In evergreen montane forests from 1 600 to 2500 m elevation. Flowering in February, April-Au- gust, and November-December; fruits were col- lected in February (immature fruits in August). This species ranges from Costa Rica to western Panama.
Hoffmannia arborescens is recognized by the usually narrowly obovate leaves gradually long- attenuate at the base, flowers on short inflores- cences in leaf axils (or below), larger flowers with prominent calyx lobes, long corollas, and the high- er altitude range. Occasional individual plants may be puberulent (Q. Jimenez 231 CR). This species may resemble H. hamelioides, while large-leaved specimens may be confused with the more com- mon and closely related H. leucocarpa. The dif- ferences between H. arborescens and H. pittieri are minor, and the two species may intergrade in the highlands of western Panama, where H. pittier is common and distinctive.
This species was described from material orig- inating from Sta. Rosa del Copey at ca. 1 800 m elevation (Tonduz 12230 = J. D. Smith n. 8121 us holotype). An isotype labeled as H. arborescens at CR lacks any indication of having the leaf-like floral bracts 1 cm long mentioned in the original description. This may have been an error on the part of J. D. Smith, as no such bracts have been seen in any Costa Rican material of Hoffmannia. The type bears the same Herb. Nat. C.R. number
(12230) as does the type of H. decurrens Standl. (q.v.), and they probably represent a mixed col- lection of two different species.
Hoffmannia areolata Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 52. 1940. Figure 9.
Herbaceous subshrubs or slender stemmed shrublets, 0.6-1 .5(-2) m tall, leafy stems 2-9 mm thick, glabrescent or sparsely pilose with crooked hairs 0.3-0.7 mm long; stipules 3-6 mm long and 2-4 mm broad at the base, glabrous, ligulate, caducous. Leaves with petioles 3- 40(-90) mm long, difficult to distinguish from the de- current lamina base; leaf blades (6-)8-20(-30) cm long, 3-8(-12) cm broad, elliptic to ovate-elliptic or oblan- ceolate, apex tapering gradually and acute or short-acu- minate, more abruptly narrowed to an obtuse or cuneate base and long-decurrent on petiole, drying membrana- ceous to stiffly chartaceous, dark greenish above, the surface distinctly bullate and with short (0.2-0.5 mm) scabrous hairs above, with thinner hairs ca. 0.3 mm long beneath, 2° veins 1 1-16/side. Inflorescences verticillate in leaf axils, glomerulate, to 2 cm long and 8-15 mm broad, sessile, often obscuring the node, with pink hairs, the flowers sessile or subsessile. Flowers closely con- gested within the inflorescence, reddish tomentulous, ca- lyx lobes 3-5 mm long and 0.5 mm broad, greenish to magenta; corolla funnelform, white, rose or translucent, ca. 4.5 mm long; tube ca. 2.5 mm long with lobes ca. 2 mm long; anthers ca. 1.5 mm long. Fruits 4-8 mm long, 3-6 mm diam. (to 10 mm in life), ellipsoid to subglobose, rose to deep red, spongy.
A little-collected species of montane cloud forest formation, 900-2100 m elevation. Flowering in March-December; fruiting in April, July-Septem- ber, and January. The species ranges from the Cor- dillera de Tilaran eastward to the Chiriqui High- lands of Panama.
Hoffmannia areolata is recognized by its rugose- bullate leaves (areolate beneath), densely flowered axillary sessile glomerulate (usually verticillate) in- florescences, very small flowers, and hairs with thickened bases. The leaves are very dark green above in life. In general aspect, these plants re- semble H. congesta and some Psychotria spp. with axillary inflorescences.
Hoffmannia asclepiadea Standl., J. Wash. Acad. Sci. 15: 7. 1925. Figure 10.
Slender herbaceous shrubs, 1.5-2(-3) m tall, usually unbranched, leafy stems 2.5-7 mm thick, sparsely to densely pilose with crooked hairs 0.3-0.5 mm long (?rarely glabrous as in Williams el al. 28096 CR, F); stipules 2-4 mm long and 3 mm broad at the base, triangular, pu- berulent, caducous. Leaves with petioles 1.5-5 cm long,
170
FIELDIANA: BOTANY
1-3 mm thick, densely to sparsely puberulent; leaf blades 6-21 cm long, 4-9(-ll) cm broad, elliptic to elliptic- oblong or ovate-elliptic, apex acuminate with tip 1-2 cm long, base rounded to obtuse to cuneate-decurrent, dry- ing chartaceous, dark brown above (much paler be- neath), glabrous or sparsely pubescent above with hairs to 0.7 mm long, more densely pubescent beneath with crooked reddish hairs, 2° veins 9-12/side, often loop- connected near the margin. Inflorescences 2/node, open cymose, 2-4 cm long, with many closely spaced flowers on pubescent peduncles 3-10 mm long and ca. 0.7 mm thick, usually with 3 major branches and small (0.6 mm) bracteoles, pedicels 1-3 mm long, short villous. Flowers pubescent, with 5 distal longitudinal lines of hairs on the unopened buds, hypanthium 1-1.5 mm long, calyx lobes 0.5-1 mm long, narrowly triangular; corolla funnelform, pale yellow, tube 6-8 mm long, ca. 1.5 mm diam., lobes 2-3(-4-5?) mm long, to 2 mm broad. Fruits said to be subglobose and red.
Plants of wet cloud forest formations from 1 900 to 2300 m elevation. Flowering in April-July. The species is known only from along the Cordillera Central and western portion of the Cordillera de Talamanaca in central Costa Rica.
Hoffmannia asclepiadea is recognized by the dense puberulence on leaves and inflorescences, compact many-flowered inflorescences on very short peduncles, longer corolla tubes, and restric- tion to higher-elevation cloud forests. This species may be no more than a very puberulent form of another species, such as H. arborescens.
6 mm long and 2 mm broad; anthers ca. 3.2 mm long. Fruits becoming red or orange-red, 6-10 mm long and 4-9 mm diam., subglobose but with a truncated apex and persisting calyx.
Plants of wet evergreen forest formations from 50 to 1200 m elevation in Costa Rica. Sometimes forming a ground cover under the deep shade of the forest. Flowering in April-August; fruiting Jan- uary-July in Costa Rica. This species ranges from Veracruz, Mexico, to central Panama.
Hoffmannia bullata is recognized by its smail stature with roots at lower nodes, prominently bul- late leaves that are obtuse or rounded apically, few-flowered inflorescences with long peduncles, and puberulence on many parts. The sublineolate 4° venation is distinctive but difficult to see in some collections. The midvein is often white on the dark green upper surface in living material. This species displays much variation within its range and within Costa Rica. Collections from the Pacific slope and near the Panama border tend to have smaller stature and narrowly elliptic-oblong leaves with more deeply impressed venation and acute apices (fig. 9, uppermost left). These differ- ences are worthy of more detailed study. In ad- dition, H. discolor (Lemaire) Hemsl. of Veracruz, Mexico, is closely related, but the leaves dry darker, the flowers have longer (2-4 mm) linear calyx lobes, and the fruits are larger.
Hoffmannia bullata L. O. Williams, Fieldiana Bot. 36: 52. 1973. Figure 9.
Herbs or herbaceous subshrubs, 10-60 cm tall, erect or decumbent, stems unbranched, lower nodes with roots, leafy stems l-4(-7) mm thick, densely pilosulous with curved hairs 0.1-0.3 mm long drying reddish brown, terete; stipules 2-3 mm long, triangular, deciduous. Leaves opposite, petioles 4-45(-55) mm long, 1-2.5 mm thick, pilosulous with curved or crooked multicellular hairs; leaf blades 4-22 cm long, 2-9(-12) cm broad, obovate to oblong-obovate, apex rounded to bluntly obtuse, base obtuse to cuneate (not or only slightly decurrent on pet- iole), drying chartaceous, grayish or grayish brown above, glabrous above, minutely puberulent on the veins be- neath with curved reddish hairs ca. 0.2 mm long, 2° veins 5-12/side, many 4° veins subparallel (sublineolate) and demarking narrow rectangular areas on the upper dried surface (but often difficult to see). Inflorescences usually solitary in leaf axils ( 1 -2/node), 3-8 cm long, glomerulate or subumbellate, peduncles 2-7 cm long, 0.5-1 mm thick, glabrous or sparsely pilose, with 3-9 flowers along the same side of bifurcate branches, pedicels 1-5 mm long. Flowers sparsely puberulent or subglabrous, hypanthium 2-3 mm long, 1.5-2 mm broad distally, urceolate or turbinate, red, calyx lobes 1-2.5 mm long, 0.4 mm broad and acute; corolla rotate, pale red or rose, 7-10 mm long in bud, tube 3-4 mm long and 0.8 mm diam., lobes 3-
Hoffmannia congesta (Oerst.) Dwyer, comb. nov. Xerococcus congestus Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1852: 52. 1853. Figure 9.
Herbs or subshrubs, 0.5-2(-3) m tall, stem erect and usually unbranched, terete, leafy stems 3-15 mm thick, glabrous or with a few large (1 mm) crooked hairs; stip- ules 6-14(-20) mm long, 4-6 mm broad at the base, triangular-ovate, with or without an awn, glabrous, co- riaceous, deciduous. Leaves with petioles 2-8(-15) cm long, 1.2-3 mm broad, glabrous or with crooked mul- ticellular hairs to 2 mm long; leaf blades 15-30MO) cm long, 8-15(-25) cm broad, broadly elliptic to broadly ovate-elliptic or elliptic-obovate, apex short-acuminate, somewhat rounded to abruptly narrowed at the obtuse to acute base and slightly decurrent on petiole (rarely long-decurrent or sometimes rounded to a truncated base in large leaves), drying stiffly chartaceous, dark brown or dark grayish above, glabrous or with scattered crooked hairs, 2° veins 7-14(-17)/side. Inflorescences solitary in leaf axils (2/node), sessile and usually glomerulate, many- flowered, 1-2 cm long and to 6 cm broad (across the node), subglobose to verticillate, reddish, pedicels 0-2.5 mm long. Flowers tightly congested, sparsely and mi- nutely puberulent, hypanthium 3 mm long and 1-2 mm
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diam.. calyx red, calyx lobes 3-6 mm long, 1-1.5 mm broad, obtuse at the tip, with parallel venation; corolla salverform, white, tube 2-3.5 mm long, lobes 2-3.5 mm long and 1 mm broad. Fruit becoming white and spongy, ca. 5 x 3 mm when dried, oblong.
Understory plants of the wet Caribbean slopes from 700 to 1800(-2400?) m elevation. Flowering in January-October; fruiting in October-March. The species ranges from the Cordillera de Tilaran eastward to Veraguas, Panama.
Hoffmannia congesta is characterized by its ses- sile (often verticillate) glomerulate reddish inflo- rescences, long calyx lobes, white fruit, un- branched stems, large leaves, and prominent stipules. This is one of our most commonly col- lected and most distinctive species of Hoffmannia. In the past it was assigned to its own genus (Xero- coccus); specimens are often found in the uniden- tified section of Rubiaceae collections.
Hoffmannia davidsoniae Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 53. 1946. (?= H. lan- cistigma Dwyer, Ann. Missouri Bot. Gard. 56: 279. 1969). Figure 7.
Herbs or slender subshrubs, 0.5-1 .7 m tall, stems usu- ally unbranched, leafy stems 1.5-5 mm thick, glabrescent (rarely densely puberulent, cf. Davidse et al. 29175 CR); stipules 1.5-3 mm long, ca. 1 mm broad at the base, narrowly triangular, caducous. Leaves with petioles 1-7 cm long, 0.8-2 mm thick, glabrous; leaf blades 8-17 cm long, 3-9(-12) cm broad, broadly oblong to elliptic-ob- long or ovate-oblong (rarely lanceolate), apex narrowed abruptly and short-acuminate, base obtuse to acute and slightly decurrent on petiole, drying grayish to dark brown or dark grayish green above, glabrous above and below or minutely puberulent, 2° veins 10-14/side, sometimes arising from the midvein at nearly 90°, loop-connected distally and forming an arcuate submarginal vein 1-4 mm from the edge. Inflorescences borne on lower leafless nodes, 1-2/node, 5-15 cm long, paniculate and with a prominent peduncle 4-6(-12) cm long and 1-2.3 mm thick, glabrous, with 3(-4) distal branches and each branch with 7-17 flowers, pedicels 2-4(-6) mm long. Flowers glabrous, hypanthium 2-3 mm long, ca. 1 mm diam. distally, turbinate, calyx lobes 0.3-1 mm long, triangular; corolla rotate to fimnelform, yellow or flushed with pur- ple, tube 1-4 mm long, lobes 2-4(-6) mm long; anthers to 3.8 mm long. Fruit ca. 8 mm long and 4 mm diam., ellipsoid to subglobose, becoming red.
Plants of wet cloud forest formations at 700- 2300 m elevation. Flowering in January, April, July, and September; fruiting in September and December-January. The species ranges from cen- tral Costa Rica eastward to the Chiriqui High- lands.
Hoffmannia davidsoniae is a little-collected spe- cies characterized by the long stiff peduncles usu- ally terminated by three branches and the long- petiolate leaves with an arcuate submarginal vein. This species is poorly known in Costa Rica, and there appears to be great variation from collection to collection. The secondary veins often arise at nearly 90° from the midvein, but there are collec- tions in which this is not the case. This species is closely related to H. liesneriana.
Hoffmannia decurrens Standl., Contr. U.S. Natl. Herb. 20:205. 1919.
Slender shrubs or subshrubs, 1.5-3 m tall, leafy stems 1.5-4 mm thick, densely puberulent with crooked yel- lowish brown or reddish brown hairs 0.1-0.3 mm long; stipules 1.5-3 mm long, ca. 2 mm broad at the base, triangular, hirstulous, deciduous. Leaves with petioles 5-40 mm long, ca. 1 mm thick, difficult to distinguish from the lamina base, densely hirsutulous with hairs 0.3- 0.5 mm long; leaf blades 7-21 cm long, 2-7 cm broad, elliptic-obovate to oblanceolate or elliptic, apex acu- minate, gradually narrowed to a narrowly cuneate base and long-decurrent on petiole, drying membranaceous or thin-chartaceous and dark greenish brown or grayish green above, glabrous on the upper surface, pubescent on the veins beneath with hairs 0.2-0.7 mm long, 2° veins 7-10/side. Inflorescences 1-3/axil, l-2(-3)cm long, cymose or fasciculate with few (1-5) flowers or rarely 2-branched and scorpioid with 5-10 flowers, peduncles 5-10(-24) mm long, 0.2-0.3 mm thick, puberulent, ped- icels 2-6 mm long, slender. Flowers sparsely to con- spicuously pubescent, hypanthium 0.7-1 .3(-2) mm long, 0.6-1(-1.7) mm diam. distally, turbinate, calyx lobes 0.7-1.5 mm long, triangular; corolla yellow or whitish, tube 2-4 mm long, lobes 3-7 mm long, ca. 2 mm wide at the base. Fruits becoming 8-9 mm long, 5-6 mm diam. (dried), ovoid to globose, red, borne on slender peduncles and pedicels.
Plants of montane evergreen forest formations, 1 100-2000 m elevation. Flowering in April-June and December; fruiting in December-February. This poorly characterized species is endemic to Costa Rica and presently known to range from Monteverde to Sta. Maria de Dota.
Hoffmannia decurrens is recognized by the pu- berulent young stems, narrowly obovate leaves with cuneate long-decurrent base, and very short inflo- rescences with very slender peduncle and pedicels. This species is poorly characterized at present, with a diverse array of specimens placed here. This species may include material presently assigned to H. hamelioides. The type, Tonduz 12230 us, bears the same collection number as the type of H. ar- borescens; these types (not seen) probably repre- sent a mixed collection.
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Hoffmannia dotae Standl., J. Wash. Acad. Sci. 18: 181. 1928. H. ramonensis Standl., loc. cit. 180. 1928. Figure 10.
Single-stemmed shrubs or herbaceous subshrubs, (0.5-)l-3(-6?) m tall, leafy stems 2-10 mm thick, gla- brous to pilosulous, older stems terete and hollow, woody; stipules 3-4 mm long and 4 mm broad at the base, glabrous, deciduous. Leaves 2 (rarely 3) at a node, pet- ioles 4-20 mm long (to 5 cm if including the decurrent lamina base), ca. 1.5 mm broad, glabrous; leaf blades 1 5-30(-35) cm long, (4-) 7-1 2 cm broad, elliptic-obovate to obovate-oblong or oblanceolate, apex short-acumi- nate, tapering gradually or abruptly to the cuneate and long-decurrent base, the narrowed basal part of the lam- ina to 6 cm long, drying membranaceous to thin-char- taceous, usually dark brown or gray above, glabrous above, pubescent in very early stages beneath and be- coming glabrous or with reddish brown hairs 0.1-0.4 mm long along the major veins, 2° veins 9-15/side and often loop-connected near the edge. Inflorescences ax- illary or at older leafless nodes, l-3/axil(2-6/node), 1.5- 3(-5) cm long, ca. 1 5 mm broad, usually paniculate with
3 primary branches and 4-9 flowers, peduncles 0-5 mm long (to 35 mm when the primary peduncle is reduced and the 1° branches function as peduncles), minutely puberulent, pedicels 3-6 mm long. Flowers conspicu- ously (0. 1-0.3 mm) puberulent or glabrous, hypanthium 3-4 mm long, 2-3 mm diam. distally, obconic and red- dish, calyx tube ca. 0.5 mm long and rotate, lobes (l-)2-
4 mm long, 1-2 mm broad at the base, bluntly triangular to oblong, often held horizontally; corolla rotate, usually red or orange at the base and orange or yellowish distally, fleshy and glabrous, buds ca. 10 mm long, tube 2-5 mm long, 2.5-4 mm diam., lobes 4-9 mm long, 2-3 mm broad, yellowish within. Fruits 6-8 mm long and 4-6 mm diam. (dried), ca. 8 mm diam. in life, orange or reddish.
Plants of wet montane cloud forest formations, from 850 to 2100 m elevation. Flowering in Feb- ruary-September; fruiting in September-January. It is uncommon in Costa Rica, except for the San Vito area. This species ranges from the Cordillera de Tilaran to the Chiriqui Highlands of Panama.
Hoffmannia dotae is recognized by its larger leaf blades with long-decurrent base forming narrowly winged margins along the sides of the petiole (but not auriculate at the base), the often cauliflorus few-flowered inflorescences, and the short thick corolla tube with larger stiff yellow petals. The flower buds with their short conical corolla tubes and broad (3-4 mm) base distinguish this species from all other Costa Rican species, none of which have such thick corolla tubes. Compare H. leu- cocarpa with more numerous slender flowers and leaves acute to cuneate at the base.
HofTmannia gesnerioides (Oerst.) Kuntze, Rev. gen. pi. 285. 1891. Ophryococcus gesnerioides Oerst.,
Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 53. 1852.
This species ranges from Guatemala to central Nicaragua at elevations of 1000-1 500 m. The small stature (to 60 cm), long reddish hairs, and few- flowered cymes on peduncles that elongate to 3 cm in fruit help to distinguish this species, which is not known from Costa Rica.
HofTmannia hamelioides Standl., J. Wash. Acad. Sci. 15: 8. 1925. Figure 11.
Herbs or shrubs, (0.5-)1.5-3 m tall, stems often branched, leafy stems 1.2-3 mm thick, usually densely covered with crooked reddish brown multicellular hairs 0.3-1 mm long (sometimes glabrescent); stipules 2-3 mm long, triangular, drying reddish brown. Leaves with petioles 6-25 mm long, 0.7-1 .5 mm thick, usually dense- ly puberulent; leaf blades 8-23 cm long, 3-8 cm broad, oblanceolate-elliptic to narrowly obovate or elliptic-ob- long, apex acuminate, base gradually narrowed and acute or long-cuneate, decurrent on petiole, drying membra- naceous to thin-chartaceous, dark brown to blackish brown above, glabrous or pubescent beneath with crook- ed hairs 0.2-0.7(-1) mm long, 2° veins 7-12/side. Inflo- rescences axillary or at older nodes, solitary (1-2/node), 1-3 cm long, usually on short (1-10 mm) slender pe- duncles, with 3-7 flowers, pedicels 0.7-1.5 mm long, 0.2-0.4 mm thick, minutely (0.1-0.2 mm) puberulent. Flowers with short (0.2-0.7 mm) crooked hairs, buds ca. 8 mm long, hypanthium ca. 2 mm long and 1.5-2 mm diam., calyx lobes 0.7-1.5 mm long, 0.3-0.6 mm broad, narrowly oblong to linear; corolla yellowish white or yellow, tube 1.5-2 mm long, 1.5-2 mm diam., lobes 2- 4 mm long, 1.5-2 mm broad, apex acute, becoming re- flexed. Fruits 4-8 mm long, 3-6 mm diam. (dried) red- dish at maturity, with persisting calyx lobes.
Plants of evergreen forests in the central high- lands, from 450 to 2200 m elevation. Flowering in February and May-July; fruiting in July and December. This species is known only from the Central Volcanic highlands and the Cordillera de Talamanca in Costa Rica.
Hoffmannia hamelioides is recognized by the usually puberulent young stems, narrowly oblan- ceolate leaves gradually tapering to the long-at- tenuate base on slender petioles, small inflores- cences with filamentous peduncles and pedicels, and short corolla tubes. Compare this species with H. decurrens (perhaps synonymous) and H. valerii with cuneate to truncated leaf bases and lower elevation habitat. Glabrous individuals resemble a number of species, especially H. longepetiolata.
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Hoffmannia hammelii C. M. Taylor, sp. nov.
Species Hoffmanniae laxae similis, sed ab ea stipulis brevioribus (0.7-1.5 mm longis) ac cymis secundis dif- fert; etiam H. pallidiflorae similis, sed foliis basi acutis ac lobulis corollinis brevioribus (3-4 mm longis) differt.
TYPUS— Liesner2907 (holotypus CR, isotypus MO), from near Sirena, Corocovado National Park, 0-200 m alt., 5 July 1977, Puntarenas, Costa Rica.
Herbaceous subshrubs, 0.3-0.5(-1) m tall, stems branched only near the ground, leafy stems 1.3-3 mm thick, glabrous and terete, often drying dark; stipules 0.7- 1.5 mm long, ca. 1 mm broad at the base, glabrous, deciduous. Leaves with petioles 6-27 mm long, 0.7-1.4 mm broad, glabrous; leaf blades (4-) 7- 18 cm long, (1.5->4-7 cm broad, elliptic to elliptic-obovate or ellip- tic-oblanceolate, apex acuminate, base acute (not de- current), drying stiffly chartaceous, grayish above, usu- ally paler or yellowish beneath, glabrous above and below, 2° veins 7-13/side. Inflorescences solitary and axillary to leaves (1-2/node), 2-7 cm long but the flowering por- tion only 1-2 cm long, with a single rachis or cymose with a terminal flower and 2 lateral branches with 5-12 flowers each, peduncles 1 5-50 mm long, rachis with 2- 7(-l 1) flowers along 1 side, glabrous, pedicels 2-6 mm long. Flowers with ovary/hypanthium 2 mm long and 1 .5 mm diam., calyx tube 0.5 mm long, calyx lobes 0.5- 0.8 mm long, ca. 1 mm broad at the base, triangular; corolla rotate, white to yellow-green, tube 2-3 mm long, lobes 3-4 mm long. Fruits ca. 5 mm diam., ellipsoid to subglobose, red, glabrous.
Plants of the evergreen Pacific lowlands, from near sea level to 200 m elevation. Flowering in July-September and November; fruiting in Jan- uary. This species is endemic to the Golfo Dulce area and the Osa Peninsula.
Hoffmannia hammelii is recognized by its small stature, leaves tapering equally to apex and base, small flower groups on long slender peduncles, leaves often drying grayish or reddish brown, and restricted geographic range. This species appears to be part of a species group including H. laxa, H. pallidijlora, H. bullata, and H. discolor (Lemaire) Hemsl. of Mexico. The species is named in honor of Barry Hammel, who has made many important contributions to our knowledge of Costa Rica's Rubiaceae. Other collections seen were M. Cha- varria et al. 254 CR, Kernan 813 & 1249 CR, MO, and Knapp 2184 CR, MO.
Hoffmannia inamoena Standl., J. Wash. Acad. Sci. 18: 179. 1928. H. fimbrianthera Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969. Figure 11.
Herbaceous subshrubs. 1-2 m tall, with few branches, leafy stems 2.5-5 mm thick, sparsely to densely minutely (0.1-0.2 mm) puberulent in early stages but soon gla-
brous and drying grayish, quadrangular; stipules 2-3 mm long. Leaves with petioles 10-60 mm long, 0.8-2 mm thick, minutely and sparsely puberulent or glabrous; leaf blades 8-22 cm long, 3.5-10 cm broad, elliptic-oblong to oblong or obovate, apex acuminate with tip 3-1 5 mm long, base obtuse to acute and usually slightly decurrent on petiole, drying membranaceous or thin-chartaceous and grayish green above, glabrous above, very minutely (0.05 mm) puberulent on the veins beneath or glabrous, 2° veins 9-14/side and loop-connected near the margin. Inflorescences axillary or on older leafless nodes (1-2/ node), ca. 1 cm long and 1-5 flowers, glomerulate, sessile or with peduncles 1-3 mm long, pedicels 1-3 mm long, pubescent with hairs 0. 1-0.4 mm long or glabrous. Flow- ers with crooked reddish hairs to 0.3 mm long (rarely glabrous), hypanthium ca. 2 mm long and 1.7 mm diam. distally, urceolate, calyx lobes 1.3-2.3 mm long, nar- rowly oblong to narrowly triangular, apex obtuse; corolla rotate, deep red, puberulent, tube ca. 1.5 mm long, lobes ca. 4 mm long. Fruits 5-7 mm long, subglobose, becom- ing white, puberulent.
Plants of evergreen forest formations from 400 to 1000 m elevation. Flowering in January and May-July; fruiting in June and November-De- cember. The species ranges from Bijagua, Alajue- la, eastward along the Caribbean slope to Rio Re- ventazon, and in the General Valley of the Pacific slope.
Hoffmannia inamoena is recognized by the minute puberulence on many parts, secondary veins clearly loop-connected near the margin of the leaf, very small sessile inflorescences, and rel- atively large narrow calyx lobes. The material placed here is rather poor and may not represent a single species. Holm and Iltis (58 F) noted that both flowers and fruit fall easily from the stems.
Hoffmannia laxa Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 1 16. 1940. Figure 11.
Herbaceous subshrubs or slender shrublets 1-2 m tall, branched only near the base, leafy stems 1-6 mm thick, glabrous; stipules 1-3 mm long, triangular, glabrous, de- ciduous. Leaves with petioles 1 .5-6(-9) cm long, 0.8-2.2 mm broad, glabrous; leaf blades 7-18(-21) cm long, 4- 9.5(-ll) cm broad, elliptic to elliptic-ovate or ovate, apex acuminate, tapering gradually or abruptly to the cuneate or acute base and slightly decurrent on petiole, drying thin-chartaceous, grayish green to dark brownish green above, glabrous above and below, 2° veins 7-1 1/ side. Inflorescences 1-3 in each leaf axil (2-6/node), 3- 8 cm long, ca. 2 cm broad, pendant, open paniculate or racemose with few (8-15) distant flowers, reddish in life, peduncles 10-45 mm long, 0.2-0.8 mm thick, glabrous, rachis often elongate with alternate flowers and minute (0.5 mm) bracteoles, pedicels 4-9 mm long (to 16 mm in fruit), 0.2-0.3 mm thick (dried). Flowers glabrous, hypanthium ca. 2 mm long and 1.8 mm diam., subglo- bose, calyx lobes 0.2-0.7 mm long, triangular; corolla
174
FIELDIANA: BOTANY
funnelform, white to pale green, tube l-2(-3) mm long, lobes 3-5 mm long and 1 mm broad at the base; anthers ca. 2 mm long. Fruits to 8 mm long, 4-5 mm diam., globose to ellipsoid, greenish white when immature, whitish purple or red to bluish black or purple-black at maturity, with a small persisting calyx tube; seeds black.
Plants of evergreen lower montane forest for- mation on both the Caribbean and Pacific slopes, 200-1600 m elevation. Flowering in February- June and August-October; fruiting in January and March-October. This species is known only from Monteverde eastward to the slopes of Volcan Poas and in western Panama.
Hoffmannia laxa is recognized by the often long- petiolate somewhat larger leaves, the lack of pu- bescence, the open few-flowered inflorescences with long thin filamentous (when dried) peduncles and pedicels, and the small flowers with very short corolla tubes. The name H. capillacea Dwyer has been applied incorrectly to material of this species.
Hoffmannia leucocarpa Stand!.. J. Wash. Acad. Sci. 1 5: 9. 1925. //. carpinteraeStendl, N. Amer. Fl. 32: 199. 1934, nom. nov. for//, macrophylla Standl., J. Wash. Acad. Sci. 15: 9. 1925, non //. macrophylla Hemsl. (?= H. trichocalyx Standl., J. Wash. Acad. Sci. 18: 181. 1928.) Figure 10.
Slender shrubs or small treelets, l-2.5(-4) m tall, with few or no lateral branches, leafy stems 2.5-8 mm thick, glabrous (rarely ferruginous-villous), usually hollow; stipules 1.5-4 mm long, 2-4 mm broad at the base, triangular, glabrous, deciduous. Leaves with petioles 1.5- 7(-10)cm long, 1.3-3.8 mm broad, glabrous: leaf blades 14-24(-35) cm long, 5-12(-15) cm broad, elliptic to broadly elliptic, apex tapering abruptly and acuminate, tapering more gradually to the cuneate base and slightly decurrent on petiole, drying chartaceous to stiffly char- taceous, pale greenish brown to dark grayish green above, glabrous above and below, 2° veins 11-14/side. Inflo- rescences 1-3 /axil below the leaves (2-6/node), 4-8(-l 5) cm long, with 1 or 2 orders of branching (rarely with a single rachis), often pendant, peduncles 2-5(-8) cm long, 0.5-2 mm thick, usually terminated by 3 subequal 1° branches, glabrous or less often minutely puberulent, pedicels 2-8(-10) mm long, sparsely to densely puber- ulent or villose. Flowers with hypanthium ca. 2 mm long and 1.5 mm diam., obconic, puberulent with thin hairs ca. 0.2 mm long, calyx green to pink, orange or red, calyx tube 1-2 mm long, lobes 0.5-2 mm long, bluntly tri- angular; corolla rotate to funnelform, pale pink to yellow marked with orange or red, glabrous to villosulous, tube 4-8 mm long, 1.5-2 mm diam., lobes (2-)3-7 mm long, 2 mm broad at the base; anthers ca. 4 mm long. Fruits 8-10 mm long and 7-8 mm diam., globose to obovoid (ca. 1 cm diam. in life), often bisulcate in development, white to pink or red.
Plants of the wet Caribbean cloud forests and nearby areas, from 1000 to 2300 m elevation. Flowering in February-July; fruiting in February, April, June-July, and October-December. The species ranges from the eastern part of the Cor- dillera de Tilaran (Monteverde) to the western slopes of Volcan Irazu in central Costa Rica (but see below).
Hoffmannia leucocarpa is recognized by the larger firm-textured leaves on long petioles, pe- dunculate and branched (usually pendant) inflo- rescences, and the larger flowers with corolla tubes and lobes usually equal in length. The fruit can be white or pink to red. This species differs from larger-leaved specimens of H. longepetiolata by the more rigid leaf blades with short-acuminate apices, larger inflorescences, and larger corollas. This species may intergrade with H. arborescens, forming a complex that includes H. pittieri. Dried specimens can be similar to H. dotae, but that species has smaller inflorescences and much broader and more succulent corolla tubes and the leaves have winged petioles. Most material placed here has branched inflorescences, but a few have monopodial circinnate inflorescences; this is an- other example of the great variability within Hoff- mannia species and why determining species in the genus is so difficult.
The preceding circumscription excludes a num- ber of specimens from the San Vito area formerly placed here. These are treated as unusual elements of//, pittieri or H. arborescens. Hoffmannia tricho- calyx probably represents an unusually pubescent representative of this species; it is based on Stan- dley & Torres 47690 (holotype), 47480, 47819. & 47924, all us.
Hoffmannia liesneriana L. O. Williams, Fieldiana Bot. 36: 54. 1973. Figure 8.
Herbs or slender subshrubs, 0.3-1(-1.5) m tall, stems usually unbranched, quadrangular with opposing flat and sulcate surfaces, leafy stems 4-12 mm thick, glabrous (strigillose): stipules 1-3 mm long, 3 mm broad at the base, triangular, glabrous. Leaves with petioles (2-)6-l 3 cm long, 1.5-3 mm thick, glabrous; leaf blades (1 1-)13- 24 cm long, (6-) 7- 13 cm broad, broadly ovate-elliptic to broadly elliptic-oblong, apex abruptly narrowed and short acuminate, abruptly narrowed to an obtuse or rounded base, drying chartaceous, dark grayish or gray- ish brown above, glabrous above, sparsely to densely minutely (0. 1-0.2 mm) puberulent on the veins beneath, 2° veins (7-)10-14(-l 7)/side, usually loop-connected near the margin. Inflorescences borne at older leafless nodes or near the ground, 1-4/node, 2-11 cm long, with many flowers in 1 -several dense clusters or on helicoid branch-
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es but rarely expanded, reddish to reddish purple, pe- duncles 4-50 mm long, glabrous or sparsely puberulent, pedicels 0-4 mm long, slender. Flowers subglabrous or puberulent with hairs ca. 0.2 mm long, reddish, hypan- thium 3-4 mm long, 2-2.6 mm diam., calyx lobes 2-5 mm long, ca. 0.4 mm broad; corolla salverform, rose or pinkish red to red-brown, tube 1-3 mm long, lobes 3-7 mm long, ca. 1.3 mm broad; anthers ca. 4 mm long, white. Fruits 4-6 mm long, 2-3 mm diam. (dried), red; seeds ca. 0.2 mm long.
Plants of lowland rain forest formations on the Caribbean slope, 50-600(-1 100) m elevation. Flowering in March and May-November; fruiting in the same months and in January. This species apparently ranges from Rio San Lorenzo, Alajuela, eastward to Guapiles, Limon, in Costa Rica.
Hoffmannia liesneriana is recognized by the large leaves with long petioles, the short unbranched stems, flowers in close clusters on cauliflorous in- florescences, long calyx lobes, and lower-elevation habitat. Unlike so many of our other Hoffmannia species, this species has leaves that are often abruptly narrowed at the base (but some collec- tions do have long-decurrent leaf blades). This spe- cies resembles Psychotria capacifolia with white flowers. Compare H. davidsoniae and H. eliasii Dwyer of Panama.
Hoffmannia longipetiolata Polak., Linnaea 4 1 : 567. 1877. H. tonduzii Standl., Contr. U.S. Natl. Herb. 20: 205. 1919. H. woodsonii Standl., Ann. Missouri Dot. Card. 28: 471. 1941. Figure 11.
Small shrubs or herbaceous subshrubs, 1-2 m tall, usually few-branched and slender-stemmed, leafy stems 0.8-5 mm thick, glabrous or minutely puberulent along narrow longitudinal ridges or with crooked hairs 0.5 mm long in early stages; stipules 0.7-3 mm long, 2-3 mm broad at the base, triangular, glabrous, deciduous. Leaves with petioles 3-15(-40) mm long, 0.5-1 mm broad, gla- brous or puberulent; leaf blades 4- 1 2(- 1 5) cm long, 1 . 5- 4(-6) cm broad, elliptic-obovate to elliptic or ovate-el- liptic, apex short- to long-acuminate, tip to 1 5 mm long, base acute to cuneate and somewhat decurrent, drying membranaceous to thin-chartaceous, brownish green to grayish green above, glabrous above, glabrous or mi- nutely puberulent on the veins beneath, 2° veins 4-7 (-9)7 side. Inflorescences 1-3/axil (2-6/node), 1-2 cm long, cymes or fascicles with (l-)3-6(-12) flowers, peduncles 2-5 mm long, pedicels 2-8(-10) mm long, ca. 0.3 mm thick, glabrous or rarely sparsely and minutely (0. 1 mm) puberulent. Flowers greenish yellow, glabrous, hypan- thium 1.5-2 mm long, 1.3 mm diam. distally, obconic, calyx lobes 0.5-1 (-2) mm long, ca. 1 mm broad at the base, triangular and acute; corolla rostrate, yellow, pink or cream white, tube 1.5-3 mm long, 1.3-1.5 mm diam., lobes 2-4 mm long, 1 .5-2 mm broad at the base, reflexed; anthers 3-3.5 mm long. Fruits 5-6 mm diam., subglo- bose, red to pink.
Plants of wet evergreen forest formations on both the Caribbean and Pacific slopes, 600-1 800(-2000) m elevation. Flowering in January-October, with the majority in May-July; fruiting in April and September-February. The species ranges from the Cordillera de Guanacaste eastward to central Pan- ama.
Hoffmannia longipetiolata is recognized by the smaller or medium-sized leaves that taper grad- ually to apex and base, little or no pubescence, terete stems, very small few-flowered inflores- cences, and small glabrous yellow flowers. The thin peduncles and pedicels, short corolla tubes, and larger corolla lobes are also helpful in recognizing this species. This is one of the most commonly collected species of Hoffmannia in Costa Rica. Originally it was thought that the smaller-leaved specimens determined as H. tonduzii were dis- tinct, but there are too many intermediate collec- tions to be able to segregate the larger-leaved spec- imens (some of which have long petioles). Collections from lower (600-900 m) elevations and from western Panama often have larger leaves than collections from central highland Costa Rica. Two species that are very similar to H. longipe- tiolata are H. decurrens (with densely puberulent young stems) and H. psychotriifolia (with larger fasciculate flowers and narrowly ellipsoid fruits).
Hoffmannia nesiota J. D. Smith, Bot. Gaz. 61: 374. 1916.
Shrublets, branchlets subterete, glabrous; stipules not seen. Leaves with petioles 7-8 cm long, 2-3 mm thick, pubescent; mature leaf blades 18-26 cm long, 10-13 cm broad, elliptic to broadly elliptic-oblong, acuminate, base obtuse and slightly decurrent, stiffly chartaceous and pu- bescent or glabrescent, 2° veins 12-16/side. Inflores- cences cymose with peduncles 1-5 cm long, pubescent or glabrescent, lacking bracts and bracteoles, pedicles 6- 10 mm, slender. Flowers 15-16 mm long, with calyx lobes 1.5-2 mm long, triangular; corolla 11 mm long, rotate, puberulent, tube 5-6 mm long, lobes 5-6 mm long, linear; ovary trilocular, style 12 mm long. Fruits ca. 1 cm diam.; seeds ellipsoid ca. 0.5 mm long.
A poorly known species endemic to Cocos Is- land. The species is only known from the type collection: Pittier 12387 us, collected at Wafer Bay, June 1898, and Barclay 2178 us, April 1838. The Barclay collection is much more pubescent than the type but appears to represent the same species. Fruits of the Barclay collection are two-locular, not three-locular as described for the type.
Hoffmannia nicotianifolia (Mart. & Gal.) L. O.
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Williams is a species of Mexico and northern Cen- tral America; the name has been incorrectly ap- plied to Costa Rican collections.
Hoffmannia pallidiflora Standl., J. Wash. Acad. Sci. 15:9. 1 925. H. rexmontis Dwyer, Ann. Mis- souri Hot. Gard. 67: 241. 1980. Figure 11.
Herbs or subshrubs, 0.5-1.5 m tall, erect or leaning on others, usually unbranched, leafy stems 2-6 mm thick, glabrous, apparently succulent in life and distorted on drying; stipules 2-3 mm long, 2 mm broad at the base, glabrous, caducous. Leaves with petioles (1.2-)2-6 cm long, 0.8-1 .8 mm broad, petioles of the same node often unequal in length, glabrous; leaf blades 10-25 cm long, 4-10 cm broad, broadly elliptic to elliptic-oblong or el- liptic-obovate, apex acuminate with a narrowed tip 5- 15 mm long, tapering gradually to a cuneate base (or abruptly obtuse) and decurrent on petiole, drying mem- branaceous, dark yellowish green or grayish green, gla- brous above and below, 2° veins 8- 11 /side. Inflores- cences solitary and axillary to basal leaves or cauliflorus (2/node), 3-8 cm long, with a single unbranched rachis or with 2 lateral helicoid branches 1-3 cm long, pedun- cles 15-35 mm long, 0.3-1 mm thick, flowers 3-30, pedicels 0-3 (-6) mm long, ca. 0.5 mm thick. Flowers glabrous, hypanthium 2-3 mm long and 1 mm diam. at apex, narrowly obconic, calyx lobes 1-2 m long, trian- gular to ligulate, acute; corolla funnelform, white or pale yellowish green, tube 2-4 mm long, ca. 1.3 mm diam., lobes 4.5-8 mm long, 0.8-1.7 broad. Fruits white be- coming red or dark reddish purple, 4-6 mm diam. (to 1 cm in life), globose or ellipsoid-oblong during develop- ment.
Plants of the wet cloud forests along the Carib- bean escarpment, (400-)700-1400 m elevation. Flowering in January-June; fruit in February-June, August, and November-December. Collections have been made from Bijagua, Alajuela, south- eastward to the upper Rio Grande de Orosi, Car- tago, and in western Panama.
Hoffmannia pallidiflora is recognized by the gla- brous, slender, usually simple unbranched or two- branched pedunculate inflorescences, very thin leaves usually decurrent at the base and often long- petiolate, narrowly obconic hypanthium, and re- striction to the middle elevations of the Caribbean slope. The thin foliage dries yellowish to very dark green. The slender unbranched inflorescences have flowers along only one side and usually dry yel- lowish.
Hoffmannia piratarum Standl., J. Wash. Acad. Sci. 18: 180. 1928.
Shrublets, branchlets obtusely tetragonal, glabrous;
stipules caducous. Leaves with petioles 25-45 mm long; leaf blades 10-20 cm long, 5-6 cm broad, elliptic-lan- ceolate, long-acuminate, the tip often falcate, drying membranaceous, dark green and glabrous above, sparse- ly short villous and glabrescent beneath, with 8-9 2° veins on each side. Inflorescences in leaf axils, 1-3/nodc, to 3 cm long, peduncles 1-2.5 cm long, short villous, pedicels 2-5 mm long. Flowers puberulcnt proximally, hypanthium 3 mm long, calyx lobes 2-3 mm long, tri- angular or rounded; corolla glabrous, tube ca. 4 mm long. 2 mm diam. at base and 5 mm at apex, corolla lobes ca. 6 mm long. Fruits ca. 6 mm long.
Plants of Wafer Bay, Cocos Island. Flowering in January. Standley knew it only from the type collection Pittier 16259 us, and the preceding de- scription is based on his description.
Hoffmannia pirtieri Standl., Contr. U.S. Natl. Herb. 18: 140. 1916. H. panamensis Standl., Contr. U.S. Natl. Herb. 20: 204. 1919.
Shrubs or subshrubs, 1.5-3(-4) m tall, usually with unbranched main stems, leafy stems 2-9 mm thick, be- coming woody and 12 mm thick, glabrous or densely pubescent on the young stems; stipules ca. 1 .5 mm long, ca. 2.5 mm broad at the base, broadly triangular, gla- brous. Leaves with petioles 1.5-7 cm long, 1.5-5 mm broad, glabrous or rarely puberulent in early stages, with narrow lateral margins continuous with the lamina mar- gins; leaf blades 1 2-26(-35) cm long, 4-1 2(-l 6) cm broad, elliptic-obovate to elliptic-oblong or oblanceolate, apex acuminate with tip 8-18 mm long, gradually narrowed to the cuneate base and long-decurrent on the petiole, drying chartaccous and dark brown or dark grayish green above, glabrous above and below (rarely puberulent in early stages), 2° veins 9-14/side and weakly loop-con- nected near the margin. Inflorescences 1-5/axil from thickened reduced peduncles, 2-5(-9) cm long, with (5-)10-15 flowers, peduncles 4-40 mm long, with 1° branches 3-30 mm long (often 2-branched with alternate flowers along 1 side), glabrous to densely reddish pu- berulent, pedicels to 9 mm long. Flowers glabrous to reddish puberulent, hypanthium 2-3 mm long, 1.5-2 mm diam., oblong-obconic, calyx orange-red, calyx lobes 0.5-1 mm long; corolla yellowish tipped with red or orange, tube 3-6 mm long, 0.7-1.6 mm thick, lobes 3- 5 mm long, 1.7 mm broad at the base. Fruits ca. 5 mm diam., deep green becoming deep red.
Plants of evergreen montane forest formations, 1 100-1 700(-2200) m elevation. Flowering in Feb- ruary-August; fruiting in January-February and August. The species (as here interpreted, see be- low) ranges from the San Vito de Coto Brus area into the adjacent Chiriqui Highlands of Panama.
Hoffmannia pittieri is recognized by the taller, more woody stems, larger leaves with cuneate long- decurrent leaf bases, inflorescences with slender- pedicellate flowers, short corolla lobes, and re-
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stricted latitudinal range. The inflorescences vary from short to moderately long with few to many branches. The interpretation presented here as- sumes that the very similar H. leucocarpa does not range eastward of Volcan Irazu. There is probably intergradation with H. arborescens (which usually has larger more branched inflorescences), and it may be necessary to place all this material under a more broadly denned H. arborescens.
Hoffmannia psychotriifolia (Benth. in Oerst.) Gri- seb., Fl. Brit. W. Ind. 321. 1861. H igginsia psy- chothaefolia Benth. in Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 50. 1853. Figure 11.
Slender shrubs or herbaceous subshrubs, l-2.5(-4) m tall, with few lateral branches, leafy stems 0.8-4.5 mm thick, glabrous, terete; stipules 1-1.5 mm long or reduced to a ridge, 1-2 mm broad at the base, triangular, glabrous. Leaves with petioles 8-35(-50) mm long, 0.7-1.5 mm thick, glabrous; leaf blades 5-14(-20) cm long, 1 .5^.5(-6) cm broad, narrowly elliptic to oblanceolate or narrowly elliptic-obovate, apex acuminate with tip 1-2 cm long, gradually narrowed to the acute or cuneate and decurrent base, drying membranaceous to thin-chartaceous, dark green or dark grayish brown above, distinctly paler be- neath, glabrous above and below, 2° veins 5-9/side. In- florescences usually fasciculate from a thickened solitary short-shoot (brachyblasts) in the leaf axils, with 5-30 flowers per node, peduncles rarely present or 0-3(-8) mm long, pedicels 2-8 mm long, ca. 0.3 mm thick, glabrous or minutely puberulent. Flowers glabrous, to 18 mm long, hypanthium 2-3 mm long, 0.7-1.7 mm diam. dis- tally, oblong-obconic, with longitudinal ridges, calyx lobes 0.6-2(-3) mm long, triangular to ligulate; corolla fun- nelform, pale yellow to white or pink, tube 4-9 mm long, 1-3 mm diam., lobes 3-6 mm long, 1.4-2 mm broad at the base. Fruits 6-12 mm long, 5-8 mm diam., oblong- ellipsoid, becoming red and translucent.
Plants of evergreen montane forest formations, (900-)1 100-2100(-2400) m elevation. Flowering in January-July and September-November; fruit- ing in February-September and November. The species ranges from southern Mexico and Guate- mala to Panama and occurs in the West Indies.
Hoffmannia psychotriifolia is recognized by the thin-glabrous leaves tapering gradually to both apex and base, stems rounded and slender, small, usu- ally fasciculate inflorescences, slender pedicels, larger flowers with well-developed calyx lobes, longer ridged ovary, and oblong fruit. This species may be difficult to separate from H. longipetiolata (including H. tonduzii), but that species has small- er flowers, few-flowered, often pedunculate inflo- rescences, and young stems that may have puber- ulent longitudinal ridges.
Hoffmannia refulgens (Hooker) Hemsley was based on cultivated material thought to have been collected in South America but most probably from northern Central America or southern Mexico (D. Lorence, pers. comm.). This name has been used for Costa Rican collections now placed under H. bullata.
Hoffmannia subauriculata Stand! .. J. Wash. Acad. Sci. 18: 179. 1928. H. haydenii Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969. Figure 8.
Herbaceous subshrubs, 1-2 m tall few-branched, leafy stems 1 .5-5(-7) mm thick, subglabrous; stipules 2-3 mm long. Leaves subsessile with poorly denned petioles be- cause of the long-attenuate lamina base and winged lat- eral margins 2-4 mm broad (short petioles 1-3 mm long sometimes present below the auricles); leaf blades 8- 2 5 (-40?) cm long, 3-10(-13) cm broad, obovate to ob- lanceolate or broadly elliptic, apex short-acuminate, ta- pering very gradually to the cuneate or long-attenuate base (5-7 x l cm) and usually slightly auriculate near the stem, drying dark, glabrous above, subglabrous or minutely puberulent on the veins below, 2° veins 9- 137 side. Inflorescences 2-4 on lower leafless nodes, 3-9 cm long, usually with few or no lateral branches (cymose to cincinnoid), with ca. 6-12 flowers, peduncles 2-4 cm long, glabrous or with reddish curved hairs to 0.2 mm long, pedicels 2-6(-12) mm long. Flowers with minute curved hairs or glabrous, with calyx lobes ca. 1.5 mm long, triangular; corolla not seen at anthesis. Fruits ca. 8x6 mm, ellipsoid, bright red.
Plants of wet evergreen forests in the central mountains and on the Caribbean slope, from 100 to 1 800 m elevation. Flowering in June-July and November; fruiting in December-January. The species ranges from near Monteverde to central Panama.
Hoffmannia subauriculata is distinguished by the long-decurrent leaves with broad (ca. 5 mm) lateral margins (along what would otherwise be a petiole) and their slight auriculate expansion near the base. The inflorescences appear to develop only at leafless nodes. This species is similar to H. am- plexifolia, which has three leaves per node.
Hoffmannia valerii Standl., J. Wash. Acad. Sci. 18: 178. 1928. Figure 9.
Weak-stemmed shrubs or subshrubs, 0.5-1. 5(-3?) m tall, usually without lateral branches, leafy stems 1-4 mm thick, with thin crooked multicellular hairs 0.3-1.5 mm long; stipules 1-4 mm long, rounded distally or with an awn, caducous. Leaves with petioles 8-60 mm long, 0.8-1.6 mm thick, villous with hairs 0.3-1 mm long; leaf blades 4-14(-19) cm long, 1.8-7(-9) cm broad,
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broadly elliptic to narrowly elliptic-oblong or ovate-el- liptic, apex acuminate, base obtuse to acute and some- what decurrent on petiole, drying membranaceous to thin-chartaceous, dark brown or grayish green above, glabrous or with few scattered hairs above, with more numerous crooked hairs 0.3-1.5 mm long beneath, 2° veins 6-10/side. Inflorescences axillary or at leafless nodes, 2-4/node, l-2(-3) cm long, fasciculate or con- tracted cymes with 3-9 flowers, peduncles 1-4 mm long, pedicels 2-8(-12) mm long, filiform, villous or subgla- brous. Flowers puberulent with crooked hairs that dry reddish, hypanthium 2-3 mm long, 1.5-2 mm diam. distally, calyx lobes l-3(-4) mm long, narrowly trian- gular; corolla rotate, pale yellow, tube 1-3 mm long, lobes 3-5(-7) mm long; anthers 2-3 mm long. Fruits 4- 6 mm long, 4-5 mm diam., ellipsoid, becoming orange or bright red.
Plants of the wet evergreen Caribbean slope and lowlands, often along streams and in wet areas, 50-950 m elevation. Flowering in February-Au- gust and November; fruiting in January, July, and November. The species ranges from near Arenal, Alajuela, to western Panama.
Hoffmannia valerii is recognized by the long hairs on many younger parts, slender unbranched stems, small short-pedunculate or fasciculate inflores- cences, short corolla tubes, and lower-elevation habitats. The lype (Valeria 57 us) was collected at 600 m elevation near Arenal, Guanacaste Prov- ince. This species resembles H. gesnerioides (Oer- sted) Kuntze of Nicaragua at 800-1500 m eleva- tion, but that species has longer (2-4 cm) peduncles, attenuate leaf bases, and dense spreading villose pubescence on all parts. Costa Rican material identified as H. boraginoides Dwyer ined. appear to be plants of//, valerii that have unusually long (2.5-3 mm) calyx lobes. The excellent DUKE col- lections from La Selva display considerable vari- ation in sepal lobe length, and it seems best to consider these plants all part of H. valerii.
Hoffmannia vesiculifera Standl., Publ. Field Co- lumb. Mus., Bot. Ser. 4: 285. 1929. H. kirkbridei Dwyer, Ann. Missouri Bot. Gard. 56: 278. 1969. //. morn Dwyer, loc. cit. 67: 236. 1980. Figure 8.
Herbs or subshrubs, 0.3-1.5 m tall, main stems usually unbranched, leafy stems 2.5-1 0 mm thick, usually densely villous with multicellular crooked reddish brown hairs to 1.5(-3) mm long; stipules to 2-5 mm long, little de- veloped and difficult to see among the pubescence, ca- ducous. Leaves with petioles 14-45 mm long, with a lateral longitudinal inflated chamber (vessicle or pouch) 12-20 mm long and 4-9 mm broad on each side (2/ petiole), with conspicuous reddish brown hairs; leaf blades 12-27(-40) cm long, 7-15(-20) cm broad, elliptic to el- liptic-obovate (less often obovate-oblong or oblanceo-
late), apex acuminate to short-acuminate, base obtuse to cuneate and decurrent (to the inflated vessicles), drying thin-chartaceous, dark brown or grayish above, paler grayish or reddish brown beneath, with few scattered crooked multicellular hairs above and below, more densely hirsutulous on the midveins above and below, the crooked reddish hairs 0.4-1 .3(-3) mm long, 2° veins 8-17/side. Inflorescences axillary or terminal on short lateral axillary shoots and subtended by leaf-like bracts ca. 2 cm long, solitary (2/node), sessile or with peduncles (= lateral branches?) to 3 cm long, flowers 10-many. in dense subsessile capitulae or on condensed helicoid cymes (with the inflorescences occasionally expanding to 7 cm and becoming branched in fruit, as in the type: Cooper 231 F), pedicels to 4 mm long. Flowers yellowish but drying reddish brown, with conspicuous hairs 1-2 mm long, hypanthium ca. 2.5 mm long, and 1.5 mm diam., calyx lobes 2.5-7 mm long, linear, with few to many reddish hairs; corolla rotate, glabrous, cream yellow to reddish, tube 2-4(-8) mm long, corolla lobes 4-17 mm long, to 3 mm broad at the base, lanceolate; anthers 3.5- 4 mm long. Fruits 4-6 mm long, 3-5 mm diam., be- coming bright red or white, thin-walled with longitudinal ribs, often with long ( 1 mm) crooked hairs.
Plants of wet evergreen forests of the Caribbean slope and lowlands, (20-)300-1500 m elevation. (Note: All Costa Rican collections came from above 300 m elevation.) Flowering in February-June and September; fruiting in April and October-Febru- ary. The species ranges from central Costa Rica to central Panama.
Hoffmannia vesiculifera is our most distinctive species of Hoffmannia. The unusual inflated struc- tures of the petiole are found in no other Costa Rican species of Rubiaceae and resemble those found on some Melastomaceae. The long crooked multicellular hairs, condensed inflorescences, and long-linear calyx lobes are additional distinctions. The variation in leaf size and form and inflores- cences (dense heads to helicoid cymes or, rarely, branching infructescences) is considerable but does not warrant segregation of species or subspecies. The inflorescences may all be borne on lateral ax- illary branches that are variously reduced, as ev- idenced by the paired leaf-like bracts subtending the inflorescences. Only one of our collections, Lent 91 1 F, cited small ants in the petiole chambers.
Holtonia Standley
Holtonia Standley is now considered part of Elaeagia.
Isertia Schreber
REFERENCE— B. Boom, A revision of the genus
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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Isertia (Isertieae: Rubiaceae). Brittonia 36: 425- 454. 1984.
Shrubs or trees, branchlets slender and subterete or thick and quadrangular, glabrous or puberulent; stipules interpetiolar or intrapetiolar, sometimes deeply divided and apparently 2/node, persisting. Leaves opposite, pet- iolate, often large; leaf blades drying thin-chartaceous to coriaceous, venation pinnate, domatia absent. Inflores- cences terminal and solitary, paniculate-thrysiform or racemose-thyrsiform, often large, the secondary branch- es terminating in dichasia or scorpioid cymes, flowers sessile or pedicellate. Flowers bisexual, radially sym- metrical, hypanthium ovoid to subglobose, calyx lobes 4-6, small, equal or unequal, persisting; corolla tubular- funnelform to salverform, corolla tube short or elongate, villous in the throat, corolla lobes 4-6(7), short and spreading, valvate or imbricate in bud; stamens 4-7, filaments inserted near the mouth of the tube, anthers dorsifixed, loculate with the interior of the thecae divided into small chambers, included or exserted; ovary 2-6(-7)-
locular, ovules numerous on axile placentas, style linear, stigma with 2-6(7) oblong lobes. Fruits berry-like with fleshy exocarp and 2-6 pyrenes, each pyrene (nutlet) with a bony endocarp and 2-many seeds, globose; seeds min- ute, angular, brownish, the testa deeply foveolate.
A genus of 14 species, ranging from Central America through northern South America and oc- curring naturally only in western Cuba and Gua- deloupe in the Caribbean. Boom has divided the genus into two sections: section Cassupa with 2- 3(— 4) locules and stigmas and fleshy fruit and sec- tion Isertia with (4-)5-7 locules and stigmas and hard fruit. Our two species represent both sections of the genus. The large leaves, large many-branched solitary terminal inflorescences, flowers with long narrow corolla tubes, and loculate anthers distin- guish this genus.
Key to the Species of Isertia
la. Leaf blades usually dull greenish or grayish beneath, usually acute at the base and decurrent on the petiole; corolla yellow to orange or red, corolla tubes to 28 mm long; fruit oblate, ca. 7 mm diam. I. haenkeana
1 b. Leaf blades usually whitish gray beneath, subtruncate to obtuse at the base and not conspicuously decurrent on the petiole; corolla white, corolla tubes to 55 mm long; fruit ellipsoid, ca. 10 mm diam. . /. laevis
Isertia haenkeana DC., Prodr. 4: 437. 1830. 7. deamii Bartlett, Proc. Amer. Acad. Sci. 33: 59. 1907. 7. deamii var. stenophylla J. D. Smith, Bot. Gaz. 61: 374. 1916. Figure 50.
Shrubs or small trees, 2-6(-20) m tall, branches thick and often slightly narrower at the node, leafy branchlets 3-10 mm thick, quadrangular with rounded edges, be- coming terete, densely puberulent with short (0.2-0.6 mm) grayish hairs; stipules 4/node (or interpreted as 2 with 2 almost separate lobes), 7-14(-45) mm long, 2-8 mm broad at base, narrowly triangular with long-acute apex, glabrous abaxially or sericeous along the midvein (strigulose), drying dark. Leaves often smaller beneath the inflorescence, petioles 5-50 mm long, 1.8-4 mm thick, with adaxial margins continuous with the lamina mar- gins, densely puberulent; leaf blades (7-)14-45(-64) cm long, (4-)7-16(-28) cm broad, elliptic to elliptic-oblong or elliptic-obovate, apex short- or long-acuminate, base gradually narrowed and acute to attenuate, decurrent on petiole, drying chartaceous, dark brown above, glabrous and often lustrous above, with thin erect or appressed whitish hairs 0.2-0.5 mm long on the veins beneath, 2° veins 14-22/side and loop-connected near the margin. Inflorescences 8-22 cm long, 6-12 cm broad, peduncle a thick extension of the stem, to 5 cm long, with a thick central rachis and many lateral branches, strigulose, the lateral branches (dichasia) with a short (1-2 cm) reddish orange peduncle and a terminal flower at the dichotomy
of 2 longer scorpioid branches with 4-9 flowers, the sec- ondary branches subtended by narrow bracts 3-1 1 mm long, pedicles 0-2 mm long, bracteoles 1-2 mm long. Flowers with hypanthium and calyx tube 2-3 mm long and equally broad, cupulate, subentire with 4 minute lobes, sparsely strigulose externally; corolla bright yellow turning orange or reddish, tube 17-25(-28) mm long, ca. 1.5 mm diam. at base and 2-3 mm distally, lobes 5-6, 5-7 mm long, with prominent dense yellowish hairs ca. 2 mm long on the inner face basally; stamens 5 or 6, filaments and anthers 3.5-6 mm long; ovary 4-6(-7) loc- ular. Fruits berry-like, 4-5 mm long and 6-8 mm diam., oblate, smooth and strigulose, usually with 5 or 6 car- tilaginous lobes and 5-6 multiseeded pyrenes; seeds 0.6- 0.9 mm long.
Plants of evergreen lowland wet forest forma- tions, from near sea level to 600 m elevation in southern Central America. Probably flowering throughout the year; in central Panama flowering is primarily in the early rainy season (May-July), with fruit maturing in the late wet season and early dry season (Croat, 1978). This species ranges from Mexico to Panama, Columbia, and Venezuela and is found in westernmost Cuba.
Isertia haenkeana is recognized by its large op- posite leaves with many secondary veins and de-
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FIELDIANA: BOTANY
current lamina base, four persisting stipules at each node, cupular calyx with minute lobes, narrow yel- low to red corolla tubes, short corolla lobes with bearded hairs within, and rounded berry-like fruit with hardened pyrenes within. This species resem- bles Palicourea guianensis.
Isertia laevis (Triana) B. M. Boom, Brittonia 36: 433. 1984. Cassupa laevis Triana, Ann. Sci. Nat. Paris, Ser. IV 9: 44. 1858. C. panamensis Standl., Contr. U.S. Natl. Herb. 18: 135. 1916. /. pana- mensis (Standl.) Standl., Publ. Field Columb. Mus., Bot. Ser. 8: 346. 1931. Figure 30.
Small trees to 10(-1 5) m tall, often with several trunks ca. 10 cm diam., leafy branchlets 4-12 mm thick, qua- drangular with rounded edges, densely appressed-pu- berulent or strigulose with short (ca. 0.3 mm) yellowish or brownish hairs; stipules apparently 4/node but united at the base (for 1-2 mm) and leaving a scar around the stem above the leaf bases, 6-12 (20-40) mm long, 3-6 mm broad, triangular-subulate, coriaceous, glabrous or sparsely puberulent on the abaxial surface. Leaves op- posite, petioles 1.6-7.5(-12) cm long, 2-5 mm thick, minutely (0. 1-0.2 mm) puberulent; leaf blades smaller beneath the inflorescences, 15-60 cm long, 7-22 cm broad, oblong to ovate-oblong or elliptic-oblong, apex acuminate or acute apex, tip to 2 cm long, base obtuse to rounded and truncate (rarely slightly decurrent on petiole), drying chartaceous to subcoriaceous, usually dark brown above, glabrous or subglabrous above and with the minor venation often impressed, minutely puberu- lent on the veins beneath, whitish canescent between the veins and the major and minor venation clearly outlined, 2° veins 1 5-22/side and loop-connected near the margin, proximal 2° veins arising at 90° to the midvein, 3° veins subparallel. Inflorescences (7-)14-35 cm long, (4-)6-15 cm broad, ovoid to ellipsoid in outline and with many short (4 cm) lateral branches, peduncle 2-6 cm long, to 6 mm thick, lateral branches usually a 5-flowered di- chasium (the distal branches with 2 flowers on the upper side), with short (1-3.5 cm) secondary peduncles, sparse- ly and minutely puberulent, flowers sessile or short (2- 4 mm) pedicellate, bracteoles 1-3 mm long, broadly ovate. Flowers sweet smelling and apparently opening in the evening, hypanthium and calyx tube 5-7 mm long, 3-5 mm diam., an elongate cup glabrous externally, the calyx lobes minute and 3-6 or obscure; corolla salverform, white, tube 32-55 mm long, 2-5 mm diam., sparsely and minutely puberulent externally, corolla lobes 6-7, 1 0- 1 4 mm long, with long (1mm) yellowish hairs inside near the mouth; stamens 6-7, filaments ca. 3 mm long and laminar, anthers 7-9 mm long, included; ovary 2- 3-locular, style 32-55 mm long, stigma with 2 oblong lobes 3-5 mm long. Fruits berries to 12 mm long, 8-1 1 mm diam., ellipsoid to subglobose, smooth and usually glabrous, becoming black; seeds 0.7-1 mm long.
Trees of open secondary vegetation in evergreen rain forest formations in both the Caribbean and southern Pacific lowlands, from near sea level to
about 800 m elevation in Costa Rica. Probably flowering throughout the year but with flowers and fruit collected mostly in May-September. This species ranges from Costa Rica and Panama along the Andes to Bolivia and in the adjacent Amazon Basin.
Isertia laevis is recognized by its large opposite leaves with whitish surfaces beneath, four persist- ing stipules at each node, solitary terminal inflo- rescences with long-tubed white flowers, subentire calyx tube, and subglobose berries. Panamanian material of this species was called Isertia hypoleu- ca by Croat (1978) and Dwyer (1980), but I. hy- poleuca Bentham is a different species occurring in Venezuela, the Guianas, and the Amazon basin.
Ixora Linnaeus
Shrubs or small trees, branchlets terete or angular, mostly glabrous; stipules interpetiolar (also slightly unit- ed above the petioles to form a very short intrapetiolar ridge), simple, usually broad at the base and acuminate, persisting. Leaves opposite and decussate or in whorls of 3, sessile or petiolate, laminae often coriaceous, ve- nation pinnate, domatia absent. Inflorescences terminal or less often axillary, cymose, thyrsoid or paniculate, branching often trichotomous, often with a rounded or flattened top (corymbose), flowers pedicellate and sub- tended by 2 bracteoles. Flowers bisexual and mono- morphic, hypanthium ovoid, calyx tube short, calyx lobes 4 (5), short or extended, corolla salverform, white to pinkish, red, scarlet, or yellow, corolla tube narrow, gla- brous or puberulent at the throat, corolla lobes 4 (5-8), linear-lanceolate to ovate, imbricate or convolute in bud and rotate at anthesis; stamens 4 (5-8), inserted in the throat or mouth of the tube, filaments short or none, anthers oblong to linear, dorsifixed, exserted or partly exserted; ovary 2-locular with 1 ovule in each locule, peltately attached to the middle of the septum, style filiform, stigmas 2. Fruits baccate, red becoming black, the pericarp fleshy or leathery, with 1-2 pyrenes; seeds concave-convex or plano-convex to subglobose.
A pantropical genus of ca. 400 species with the largest number of species in Africa and the Indo- Pacific area. The colorful inflorescences, narrow corolla tubes, stamens borne near the apex of the corolla tube, and two-seeded fruit help to char- acterize the genus. The flowers are protandrous in a distinctive manner: pollen is transferred from the clustered anthers into a concave area of the stigmatic head; elongation of the stigma disperses the pollen. Later, the stigma lobes open to expose their receptive surfaces for pollination. Four spe- cies are known to occur in Central America; two are native and two are widely cultivated as or- namentals.
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Key to the Species of Ixora
la. Flowers closely crowded together in capitate inflorescences; corolla tubes 25-45 mm long; plants cultivated for ornament 2
Ib. Flowers crowded in distal cymose groups in an open paniculate inflorescence; corolla tubes 3-8 mm
long; native wild species 3
2a. Flowers red or orange; leaves sessile or subsessile /. coccinea
2b. Flowers white; leaves petiolate /. finlaysoniana
3a. Fruits ca. 10 mm diam.; flowers sessile in small glomerules, peduncles 0.7-3 mm thick; leaf blades 1 2-26 cm long /. floribunda
3b. Fruits ca. 4 mm diam.; flowers in panicles and pedicellate, peduncles 0.3-1 mm thick; leaf blades 7-16 cm long /. nicaraguensis
Ixora coccinea L., Sp. PI. 110. 1753. Figure 31.
Shrubs 0.5-3(-5) m tall, much branched, leafy branch- lets 1-4 mm thick, glabrous, terete; stipules 0.5-1.5 mm long at the broad base and with a narrow tip 2-14 mm long, glabrous. Leaves opposite, sessile or subsessile with petioles to 2 mm long; leaf blades (2-)3-l 0(-1 6) cm long, ( 1-) 1 .5-4.5(-6) cm broad, oblong to elliptic-oblong, ovate- oblong or oblong-obovate, apex rounded to obtuse or acute and often with minute (0.5-1 mm) slender tip, base rounded and subtruncate or subcordate to obtuse, drying stiffly chartaceous or subcoriaceous, glabrous above and below. Inflorescences terminal, to 10 cm long, flowers closely clustered (subcapitate), peduncles 1-3 cm long, flowers usually sessile at the apex of short-stipitate triads (cymes). Flowers with hypanthium 1-1.5 mm long, tur- binate, calyx lobes 4, ca. 1 mm long; corolla red or or- ange-red, tube 25-45 mm long, only 0.5-1 mm diam., glabrous or subglabrous, corolla lobes 4, 10-15 mm long, 4-8 mm broad; stamens 4, anthers exserted and soon deciduous; style usually exserted. Fruits 8-1 5 mm diam. (rarely developed in cultivars).
Ixora coccinea is a widely cultivated species throughout Central America, from near sea level to about 1500 m elevation. The rounded clusters of brilliant reddish flowers with long slender tubes and the evergreen leaves make it an attractive or- namental. The species originated in India and is now grown throughout the tropics and subtropics. Several varieties and forms have been recognized; see F. R. Fosberg and H.-H. Sachet, Three culti- vated Ixoras. Baileya 23: 74-85. 1989. This spe- cies is called cruz de Malta, flor defuego, jazmin, jazmin rojo, and jazmin de coral.
Ixora finlaysoniana Wallich ex G. Don, Gen. hist. 3: 572. 1834. Figure 31.
Shrubs or small trees to 5 m tall, leafy branchlets 2- 6 mm thick, glabrous, becoming terete with age; stipules 2-7 mm long, triangular and acuminate or broad with 1 or 2 awns. Leaves opposite, petioles 4-20 mm long,
1-2 mm thick, glabrous; leaf blades 6-18 cm long, 2- 6.5 cm broad, narrowly oblong to narrowly oblong-ob- ovate, apex bluntly obtuse or rounded (acute), base grad- ually narrowed and obtuse or acute, drying subcoria- ceous, glabrous above and below. Inflorescences terminal, 5-10 cm long, to 10 cm broad and rounded, of densely congested cymose branches forming a capitate cluster, peduncles 0-3 cm long. Flowers glabrous, hypanthium 1-2 mm long, calyx lobes 3-4 mm long 1 .5-2 mm broad, petaloid in texture; corolla white, tube 20-30 mm long, 0.3-1 mm diam., lobes 4-6(-8) mm long, l-2(-3) mm wide; stamens 5, exserted, anthers narrow; stigma ca. 2.2 mm long, slender and narrowly 2-lobed.
Ixora finlaysoniana is widely cultivated in the tropics and planted at lower (0-1 000 m) elevations in Central America. The rounded inflorescences of many white flowers with long slender tubes, the lack of pubescence, the foliaceous sepal lobes, and the short-petiolate coriaceous oblong leaves char- acterize this species. Native of southeast Asia; this species is referred to as corona de la reina and mono de reina.
Ixora floribunda (A. Rich.) Griseb., Cat. PI. Cub. 134. 1866. Siderodendronfloribundum A. Rich, in Sagra, Hist. Cuba 1 1: 24. 1850. Figure 43.
Small trees to 1 5 m tall, short-shoots frequently pres- ent, leafy branchlets 2.2-6 mm thick, smooth and gla- brous, terete; stipules 4-10 mm long, united above the leaf bases to form a short (1-4 mm) sheath, triangular- subulate and often shifted to the side above the petiole (not strictly interpetiolar), glabrous abaxially. Leaves op- posite, petioles 10-25 mm long, 1.5-2.5 mm thick, gla- brous; leaf blades 12-26 cm long, 4-10 cm broad, oblong to elliptic-oblong, or obovate, apex obtuse to short-acu- minate, base obtuse to attenuate and decurrent on pet- iole, drying stiff-chartaceous to subcoriaceous, concol- orous, glabrous above and below, 2° veins 6-1 0/side, not loop-connected distally. Inflorescences terminal, 1-3, 5- 1 2 cm long, panicles with 2-3 pairs of opposite branches, primary peduncles 1-3 cm long, secondary peduncles equally long and densely puberulent with short (0.2-0.3
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mm) grayish hairs, flowers sessile and 3-12 in distal cymose clusters. Flowers with hypanthium ca. 1.5 mm long and 1.2 mm diam., turbinate, covered with short (0.2 mm) thin erect whitish hairs (and with some smaller glandular hairs); calyx tube 0.5-1 mm long, calyx lobes minute; corolla white, tube ca. 4 mm long, 0.3-0.6 mm diam. in proximal half, glabrous externally, white-villous within, lobes 4, 3-4 mm long, ca. 1 .5 mm wide, glabrous; filaments 0.6-1.5 mm long, anthers ca. 3.3 mm long, ca. 1.5 mm broad; stigma lobes ca. 1.5 mm long and un- equal. Fruits 6-10 mm long, subglobose, sessile, sparsely puberulent, drying red and smooth, persistent calyx tube 0.4-0.7 mm high and ca. 1.2 mm broad.
Trees of both lowland rain forest formations and seasonally deciduous formations, from 1 0 to 800 m elevation in Central America. Flowering in Jan- uary-February; fruiting in March-May in Central America. This species, apparently uncommon in Central America, ranges from Honduras and El Salvador to Colombia and some of the West In- dies.
Ixora floribunda is recognized by its stiff-gla- brous elliptic-oblong leaves, terminal inflores- cences of white salverform flowers, puberulent ca- lyx, and sessile red berries. Our collections come from the area between Canas and Bagaces and Monte Aguacate on the seasonally deciduous Pa- cific slope of central Costa Rica. The species has also been collected in the Caribbean rain forest formations of Honduras and Nicaragua. It has been called pah de Maria.
Ixora nicaraguensis Wernham, J. Hot. 50: 243. 1912. 7. rauwolfioides Standl., Trop. Woods 1 1: 25. 1927. Figure 43.
Shrubs or small trees to 10 m tall, often with many branches, leafy branchlets 1.5-3 mm thick, glabrous, te- rete, grayish; stipules 3-6(-8) mm long, 2-3 mm broad at the base, broadly triangular and acuminate to cuspi- date with a short awn to 3 mm long. Leaves opposite, petioles 3-9 mm long, 0.7-2 mm broad, glabrous, with lateral (or adaxial) ridges; leaf blades 6-1 3(-l 6) cm long, 2-5(-7) cm broad, elliptic-oblong, oblong or ovate-ob- long, apex abruptly narrowed and acute or short-acu- minate, base obtuse to somewhat attenuate and slightly decurrent on petiole, drying chartaceous, glabrous on both surfaces, 2° veins 6-12/side, often obscure and weakly loop-connected distally. Inflorescences 1-3 and terminal, 3-9 cm long and equally broad, open pyra- midal panicles with 2-3 pairs of opposite branches, pe- duncles 1-25 mm long, 0.3-1 mm thick, minutely (0.1- 0.2 mm) puberulent, flowers on slender pedicels 0-10 mm long in distal triads, distal bracts ca. 0.5 mm long. Flowers with hypanthium and calyx tube 1-1.7 mm long, minutely puberulent or glabrous, calyx lobes obsolete or minute (0. 1-0.5 mm); corolla white, glabrous externally, tube 3-6 mm long, ca. 1 mm diam., lobes 4, 3-4 mm
long, 1-2 mm broad, oblong and rounded distally; fila- ments 0.5-1 mm long, anthers 2-3 mm long, 0.3 mm thick; stigma ca. 1.5 mm long, exserted. Fruits 5-6 mm long, 4-5 mm diam., subglobose to ovoid, glabrous and red; pyrenes 5 mm long.
Plants of evergreen rain forest formations on both the Caribbean and southern Pacific slope of Costa Rica, from near sea level to 300 m elevation. Flowering in February-March and June-October; fruiting in March and July-November. This spe- cies ranges from Belize to eastern Panama.
Ixora nicaraguensis is recognized by its small white flowers on open thin-branched inflores- cences, slender corolla tubes, and small globose two-seeded fruit, borne on thin-branched infruc- tescences. This species may be mistaken for a Psy- chotria, but the corolla lobes are valvate in bud in Psychotria (and related genera).
Ladenbergia Klotzsch
Small to large trees, the bark with bitter substances; stipules usually large, interpetiolar or also united distally (intrapetiolar) and forming a ring around the stem above the leaf bases after falling, triangular to obovate, with colleters at the adaxial base, caducous. Leaves opposite (occasionally whorled), petiolate, often subcoriaceous, pinnately veined, some species with domatia. Inflores- cences terminal, paniculate with opposite branching or cymose, ebracteolate. Flowers bisexual, monomorphic, medium to large, fragrant, hypanthium turbinate to cy- lindrical, calyx tube cupular, calyx lobes short or long; corolla funnelform or salverform, sericeous externally, corolla lobes 5-6, valvate in bud, minutely papillose within and on the margins; stamens 5-6, filaments very short, inserted near the center of the tube, anthers linear, dorsi fixed, included; ovary 2-locular, placentas elongate, spongy and borne on the septum, ovules numerous in each locule and vertically imbricate, style slender, stig- matic lobes 2. Fruits elongated capsules, cylindrical or flattened, with septicidal dehiscence (but the septum very thin and dehiscence often appearing to be loculicidal). splitting from the top into 2 woody or coriaceous valves; seeds numerous, longitudinally imbricate and peltate, flattened and elongate, body oblong and surrounded with a thin flattened dentate or laciniate wing.
A genus of about 30 species ranging from Costa Rica to Bolivia; most of the species are Andean. The genus is similar to Cinchona and Condami- nea. The large salverform and sericeous flowers with long valvate corolla lobes, large stiff usually broad leaves, large broad stipules, and 2-valved capsules with winged seeds help to characterize the genus.
Paul Standley's original separation of Costa Ri-
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can material into three species appears to represent three valid morphological entities that are not sympatric, and intermediate collections are not
apparent. Nevertheless, the species are very sim- ilar, and they might prove to be three subspecies of a single species.
Key to the Species of Ladenbergia
la. Tufts of stiff hairs (0.5-1 mm long) usually present at the base of the petiole in young leaves; leaf
blades often acute at the base [corolla tubes 10-23 mm long, corolla lobes 11-17 mm long];
(1200-)! 500-2 100 m elevation L. valerii
Ib. Base of the petiole glabrous or with minute (0.2 mm) appressed hairs; leaf blades rarely acute at
the base; 0-1 500 m elevation 2
2a. Corolla tubes 15-20 mm long, corolla lobes 8-12 mm long; fruit 2-5.5 cm long; central highlands
1000-1400 m elevation L. brenesii
2b. Corolla tubes 25-50 mm long, corolla lobes 16-20 mm long; fruit 6-9 cm long; evergreen lowlands,
0-1000 m elevation L. sericophylla
Ladenbergia brenesii Standl., Pub. Field Mus. Nat. Hist., Bot. Ser. 18: 1323. 1938. Figure 29.
Trees, 5-25 m tall, leafy branchlets 4-12 mm thick, glabrous or sparsely puberulent at the nodes with short (0.1-0.3 mm) hairs, quadrangular, becoming terete and pale grayish; stipules 16-35 mm long, 4-16 mm broad, ovate to obovate, obtuse to acute at the apex, glabrous or with few slender ascending hairs ca. 0.5 mm long. Leaves with petioles 10-35 mm long, 1.8-3.5 mm thick, glabrous and drying dark; leaf blades 12-25 cm long, 5- 1 3 cm broad, broadly elliptic to elliptic-oblong or ellip- tic-obovate, apex abruptly narrowed and obtuse or bluntly acute, base obtuse or slightly rounded, drying subcor- iaceous, glabrous above, glabrous or with a few scattered hairs below, 2° veins (6-)7-10/side. Inflorescences 6-18 cm long, pyramidal, peduncles 2-7 cm long, glabrous, with opposite branches and bracts to 5 mm long, pedicels 0-3 mm long. Flowers with hypanthium 3-6 mm long, 2.5-5 mm diam., clavate-tubular, densely sericeous with yellowish brown ascending hairs, calyx tube ca. 1 mm long, calyx lobes l-2(-4) mm long, ca. 2 mm broad, broadly rounded, glabrous or sparsely puberulent distally and with a minutely ciliate edge; corolla white, tube 1 5- 20 mm long, 5-6 mm diam., densely sericeous, lobes 8- 12 mm long, 2-3.5 mm broad. Fruits (2-)3-6 cm long, 5-10 mm broad (to 16 mm when fully flattened), sub- terete and narrowly oblong before dehiscing, valves pu- berulent externally and lustrous within; seeds 12-15 mm long, 3—4 mm broad, surrounded by a thin translucent erose wing, body of the seed 3-4 mm long and ca. 2 mm broad.
Trees of wet evergreen cloud forest formations of the central highlands, 1000-1400 m elevation. Flowering in March-July; fruiting in July-Novem- ber. The species is endemic and ranges from the Cordillera de Tilaran to the western parts of the Cordillera de Talamanca.
Ladenbergia brenesii is recognized by its cloud forest habitat, large puberulent flowers, long nar-
row capsules splitting into valves that become flat, and broad glabrous leaves. Compare this species to L. valerii with which it can easily be confused. Also known as aquijilla and quina.
Ladenbergia sericophylla Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1324. 1938. Figure 29.
Trees, 8-35 m tall, trunks to 60 cm. dbh, leafy inter- nodes 5-12 mm thick, young stems quadrangular and drying dark, quickly becoming pale gray and terete; stip- ules 15-45 mm long, 10-22 mm broad, oblong or ob- ovate-oblong, rounded at the apex, minutely puberulent with thin appressed-ascending hairs 0.1-0.3 mm long. Leaves opposite, petioles 15-4 5 (-60) mm long, 2-3.5 mm thick, glabrous and drying very dark, reddish in life; leaf blades 14-24(-30) cm long, 8-16(-18) cm broad, very broadly elliptic to broadly elliptic-oblong or slightly obovate, apex abruptly narrowed and rounded or bluntly obtuse, base abruptly narrowed and obtuse or slightly attenuate, decurrent, drying stiffly chartaceous to sub- coriaceous, usually dark brown above, glabrous above, minutely (0. 1-0.3 mm) appressed-puberulent on the veins beneath and with larger (1 mm) hairs in the vein axils (domatia), 2° veins 5-8/side. Inflorescences 10-20 cm long, to 1 8 cm broad, pyramidal, peduncles 1-5 cm long, 4-8 mm thick, glabrous or minutely appressed-puber- ulent, lateral branches opposite and subtended by bracts 3-6 mm long, bracteoles subtending the flowers 1-2 mm long, rounded apically, pedicels 1-6 mm long. Flowers with hypanthium 5-7 mm long, 2-3 mm diam., tubular, densely sericeous with ascending yellowish hairs, calyx tube 2-4 mm long, cupulate, sparsely puberulent or gla- brous, calyx lobes 2-3 mm long, 2.5-3 mm broad, round- ed at the apex, glabrous and drying dark; corolla white or yellowish white, tube 25-50 mm long, 3-5 mm diam., densely sericeous, lobes 16-30 mm long, ca. 4 mm wide and narrowly oblong to lanceolate. Fruits 6-1 1 cm long, 10-14 mm broad, oblong-cylindrical and rounded-rect- angular in cross-section, straight or curved (falcate), mi-
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FIELDIANA: BOTANY
nutely appressed-puberulent, persisting calyx 3-6 mm long and 6-8 mm broad, narrowed at the base to form a pedicel 5-1 5 mm long; seeds 1 1-20 mm long, 4-5 mm broad with thin wing, body of the seed ca. 3 x 1.5 mm.
Tall trees of evergreen rain forest formations of the Caribbean lowlands and on the southern Pa- cific slope, 20- 1 000 m elevation. Flowering in Jan- uary-February, July, and September; fruiting in January-May. This species is only known from La Selva, the western parts of the General Valley, the mountains bordering the Pacific near Canas Gor- das, and the Osa Peninsula. While endemic to Cos- ta Rica, it may also occur in westernmost Panama.
Ladenbergia sericophylla is characterized by its large broad leaves with appressed hairs on the veins beneath, large broad stipules, long salverform se- riceous corollas, and large capsular fruit. The taller height of these trees may explain why our collec- tions of this species are so few.
Ladenbergia valerii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1324. 1938. Figure 29.
Small trees, 3-20 m tall, leafy branchlets 2-10 mm thick, quadrangular or terete in early stages, essentially glabrous but with stiff retrose or erect hairs ca. 0.5 mm long at the base of the petiole attachment; stipules 8-16 mm long, 5-8 mm broad, broadly elliptic or ovate, apex rounded, sparsely puberulent with appressed hairs or glabrous and with a few hairs at the base and along the midrib abaxially. Leaves opposite, petioles 6-25 mm long, 1.5-2.5 mm thick, glabrous or very sparsely ap- pressed-puberulent (except for tufts of longer hairs below the base), drying dark; leaf blades 6-17(-28) cm long, 3-8(-13) cm broad, broadly elliptic to elliptic-obovate or narrowly elliptic, apex abruptly narrowed and short- acuminate or bluntly acute, base obtuse to acute, drying stiffly chartaceous to subcoriaceous. glabrous or with a few thin hairs above, with thin straight hairs 0.3-0.7 mm long on the major veins beneath, 2° veins 5-9/side. In- florescences 7-20 cm long, pyramidal with opposite branching, peduncles 2-7 cm long, sparsely puberulent or glabrous, bracts ca. 3 mm long, distal flowers in triads and subtended by bracteoles 0.5-1 mm long, pedicels 0- 3 mm long and merging with the hypanthium. Flowers with hypanthium and calyx tube 5-8 mm long, 2-3 mm diam., densely appressed yellowish sericeous, calyx lobes 3-6 mm long, 2.5-3.5 mm broad at the base, broadly rounded to obtuse distally and becoming reflexed, gla- brous; corolla white or white with longitudinal pink stripes, salverform, tube 10-23 mm long, 1.5-4.5 mm diam., densely sericeous with ascending hairs, lobes 5, (11-)12-17 mm long, 2.5-4.5 mm broad, lanceolate to narrowly oblong, papillate-puberulent within, becoming recurved. Fruits 3-8(-l 1) cm long, 8-14 mm broad; seeds 10-14 m long, 2-4 mm wide, body of the seed 3 m long and 1.5 mm wide.
Trees of montane cloud forest formations from
(1200-)! 600 to 2100 m elevation. Flowering in January-September and November; fruiting in May-August. The species ranges from the Cor- dillera de Tilaran to the Cordillera de Talamanca (as far east as 83°04' W) and will probably be found in nearby Panama.
Ladenbergia valerii is recognized by its higher- elevation habitat, unusual tufts of stiff short hairs beneath the petioles on otherwise glabrous or gla- brescent stems, sericeous corolla with relatively long corolla lobes, and capsular fruit. This species is very closely related to L. brenesii of similar for- ests at somewhat lower elevations.
Lasianthus Jack
Herbaceous subshrubs (in Central America), shrubs or rarely small trees, glabrous or pubescent; stipules inter- petiolar, broadly triangular to lanceolate, persisting or deciduous. Leaves opposite and decussate, petiolate, usually acuminate at the apex, pinnately veined and of- ten with many arching secondary veins, without doma- tia. Inflorescences axillary, mostly sessile fascicles or cy- mose glomerules, (sometimes pedunculate and simple or branched), flowers sessile or subsessile, bracts small. Flowers bisexual (rarely unisexual and monoecious), ra- dially symmetrical, mostly small, sometimes heterosty- lous, hypanthium urceolate to ovoid or subglobose, calyx lobes 3-6, acute or rounded, persisting; corolla salver- form to funnelform, white, corolla tube densely hairy in the throat, corolla lobes 4-6, spreading or erect; stamens 4-6, filaments very short and borne in the throat, corolla lobes 4-6, anthers dorsifixed near the base, included or slightly exserted; ovary 4-12-locular, ovules solitary in each locule, erect from the base of the locule or septum, style short or long, stigmas 4-10, linear or lobed. Fruits usually succulent, blue to purple, black, or red, with 4- 1 2 pyrenes, the pyrenes 1 -seeded and 3-angled with flat sides, the dorsal side grooved, keeled or winged; seeds narrowly oblong.
A genus of about 1 50 species, of southeastern Asia, Malaysia, tropical Australia, and Africa. Two species are found in the West Indies, and one spe- cies in our area and South America. Our species with eight-locular ovary and style with eight stig- mas is unique among Central American Rubi- aceae. Our species was originally described as a new genus, Dressleriopsis Dwyer.
Lasianthus panamensis (Dwyer) Robbrecht, Ann. Missouri Bot. Gard. 69: 427. 1982. Dressleriop- sis panamensis Dwyer, Ann. Missouri Bot. Gard. 67: 154. 1980. Figure 7.
Herbaceous rhizomatous subshrubs to 0.6(-1) m tall, leafy stems 2-6 mm thick, hirsute with slender erect hairs
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to 2 mm long; stipules 4-7 mm long, 3-6 mm broad at the base, broadly triangular and hirsute along the midrib. Leaves with petioles 3-9 cm long, 1.5-2.5 mm thick, hirsute; leaf blades 1 0-1 9 cm long, 5-9 cm broad, oblong to elliptic-oblong, apex narrowed abruptly and short- acuminate, tip 0-8 mm long, base obtuse and rounded to subcordate-auriculate, often unequal at the petiole, drying thin-chartaceous, dark grayish brown, with scat- tered thin straight or crooked hairs 0.5-2 mm long on upper and lower surfaces, 2° veins 9-14/side and loop- connected near the margin to form an arcuate submar- ginal vein, often with a shorter (parallel) minor 2° vein between the major. Inflorescences axillary, dense fasci- cles of sessile flowers 1-2 cm broad, glomerulate or ver- ticellate, bracts ca. 2 mm long and difficult to see among the long hairs. Flowers monomorphic, with hypanthium ca. 2 mm long, subglobose, hirsute, calyx lobes 4, 3-4 mm long, 2-3 mm broad, with hairs to 1 mm long; corolla white, tube 3-4 mm long, lobes 5 (4, 6), 1.5-4 mm long, oblong; stamens 5 (4, 6), anthers ca. 1 mm long: ovary with 8 locules, style to 5 mm long, stigmas 8, oblong and radiate. Fruits berry-like, to 1 1 mm diam., globose, purple-black, pyrenes 8 or fewer, 3 mm long, with an oblique scar on the concave side.
Plants of wet evergreen forest formations on the Caribbean slope (at ca. 100 m elevation) and in the central highlands of Panama (to 1 000 m ele- vation). Flowering and fruiting in July. This spe- cies is known from the La Selva area, Heredia, and in Panama and Colombia.
Lasianthns panamensis is recognized by its slen- der hirsute stems to 1 m tall, long-petiolate oblong leaves with many secondary veins forming an ar- cuate submarginal vein, sessile fasciculate inflo- rescences, and fleshy berries with up to eight py- renes. No other Central American species of Rubiaceae has ovaries with eight locules. Super- ficially, these plants resemble some species of Hqffmannia and a few species of Psychotria with axillary inflorescences.
Lindenia Bentham
REFERENCE— S. Darwin, The genus Lindenia (Rubiacea). J. Arnold Arbor. 57: 426-449. 1976.
Small shrubs growing along rivers and streams, branchlets terete, glabrous or puberulent; stipules inter- petiolar, borne above the petiole bases, short, usually persisting. Leaves opposite short-petiolate; leaf blades narrow, often drying dark, chartaceous, pinnately veined, domatia absent. Inflorescences terminal, short and few- flowered, flowers often in triads or solitary, bracts and bracteoles present, pedicels short. Flowers bisexual, ra- dially symmetrical, large and showy, hyanthium elon- gate-turbinate, with 5 longitudinal ribs or angles, calyx with 5 elongate lobes, lobes equal or unequal, persisting in fruit; corolla narrowly salverform with a narrow elon-
gate tube, glabrous within, corolla lobes 5, convolute in bud and spreading at anthesis; stamens 5, sessile on the mouth of the tube, linear and exserted; ovary 2-locular, ovules very many, vertical, placentas longitudinally ad- nate to the septum, style slender and bifid. Fruits woody capsules, clavate to pyriform or obovoid, with persisting calyx lobes distally, 2-locular, splitting septicidally into 2 valves from the top; seeds numerous, angulate/rhom- boidal.
A genus of three species, with the other two species endemic in the western Pacific islands of Fiji and New Caledonia, respectively. The stream- side habitat, narrow leaves on thick stems, few terminal flowers, and very long corolla tubes make this genus quite distinctive.
Lindenia rivalis Benth., PL Hartw. 84. 1841. Fig- ure 15.
Small shrubs, 0.4-1. 2(-2) m tall, leafy branchlets 1.5- 6 mm thick, glabrous or minutely puberulent with thin ascending hairs ca. 0.2 mm long, older stems drying black; stipules 3-5(-10) mm long, 1.5-4 mm broad, the broad basal portion 2-3 mm high and with a narrow awn to 2(-5) mm long, glabrous, thin and brown. Leaves clustered near the ends of branchlets, petioles 2-10(-16) mm long, 0.4-1 .5(-2) mm broad, little differentiated from the leaf base; leaf blades 3-12(-17) cm long, 0.8-3(-4) cm broad, oblanceolate to very narrowly elliptic-oblong or narrowly elliptic, apex tapering gradually and acute, sometimes with a short (0.5-1 mm) spine-like tip, base tapering gradually and acute, decurrent on petiole, drying chartaceous to coriaceous, the margin often revolute, glabrous above, glabrous or puberulent beneath with thin erect hairs 0.2-0.3 mm long, 2° veins 6-8/side, not loop- connected. Inflorescences of solitary flowers or short (2- 1 0 mm) pedunculate triads (or clusters of up to 7 flowers), bracts difficult to see among the distal leaves 4-9 mm long, pedicels to 10 mm long, poorly differentiated from the flower base. Flowers nocturnal, hypanthium 5-9 mm long, 2-3.5 mm diam., puberulent, calyx tube minute, calyx lobes 10-1 7(-22) mm long, 0.7-2(-2.8) mm broad, narrowly oblong and acute, green; corolla white or white tinged with pink, tube 10-17 cm long, 2-3.5 mm diam.. puberulent externally with hairs 0.2-0.5 mm long, lobes 1 5-27(-35) mm long, 5-14(-16) mm broad, narrowly to broadly elliptic, acute at the apex; stamens sessile, an- thers ca. 10 mm long, 1-1.4 mm broad; style slender equalling or slightly exceeding the tube, stigmatic area ca. 10m long. Fruits to 4 cm long, body of the fruit 15- 25 mm long, 9-14 mm broad, with long persisting calyx lobes before dehiscence, broadly obovoid or pyriform, the woody valves twisting; seeds 1.5-2 mm long.
Small woody shrubs of stream sides and often growing on rocks next to the water, in deciduous forest areas of Guanacaste Province, 0-700 m el- evation (to 1300 m in Honduras). Flowering in February-October in southern Central America (primarily April-July); probably fruiting through-
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out the year (mostly August-March). The species ranges from northeastern Mexico and along the Pacific slope of Central America to central Pana- ma.
Lindenia rivalis is recognized by its small stiff narrow leaves on thick dark branches, short stat- ure and riverside habitat in seasonally deciduous areas, few terminal flowers, and extremely long narrow corolla tube. Opler's observation (Opler 945 F) that the flower is nocturnal would be con- sistent with the long narrow tube being an adap- tation for Sphingid moth pollination. Our only other Rubiaceae with such long/narrow corolla tubes have much larger leaves. Also known asjaz- mincillo and lirio de aqua.
Machaonia Humboldt & Bonpland
Shrubs or small trees, stems often armed with spines (leafless short-shoots), glabrous or puberulent, terete; stipules interpetiolar, triangular or subulate and often with a distal awn, persisting. Leaves opposite (rarely ter- nate or verticillate), sessile or petiolate, thin-textured, pinnately veined. Inflorescences terminal, solitary, pa-
niculate with opposite branching (or rarely umbeliform), bracteate, flowers in cymose or crowded distal clusters. Flowers bisexual and radially symmetrical, small, 4-5- parted, hypanthium turbinate or obovoid, slightly com- pressed laterally, calyx tube minute or cupulate, calyx lobes 4-5(-6), equal or unequal, persisting; corolla short- funnelform or short-salverform, white, corolla tube with long hairs in the throat, lobes imbricate in bud. rounded distally; stamens 4-5, filaments short or long, borne in the throat of the tube, anthers oblong, dorsi fixed and versatile, included or exserted; ovary 2-locular, with 1 ovule pendulous from the apex of each locule. style slen- der, with 2 stigmas. Fruits small and dry, capsule-like and splitting from the bottom into 2 elongate cocci (mer- icarps), pendulous for a short period from the apex to the persisting stipe-like central axis; seeds elongate and cylindrical.
A tropical American genus of about 30 species, found in Mexico, Central America, the West In- dies, and tropical South America. The genus is recognized by its small flowers with short corolla tubes, two-locular ovary with solitary pendulous ovules, and unusual capsule-like fruit. The sub- capitate clusters of distal flowers on an openly branching panicle, small thin leaves, and occa- sional presence of spines are also distinctive.
Key to the Species of Machaonia
la. Hypanthium/ovary of the flower with few small hairs; fruit 5-7 mm long (including calyx lobes), brownish and with few hairs often in rows; lowland evergreen Costa Rica M. martinicensis
Ib. Hypanthium/ovary of the flower whitish with a dense covering of short whitish hairs; fruit 4-5 mm long, yellow or whitish with many hairs; Belize and central Panama M. acuminata
Machaonia acuminata Humb. & Bonpl., PI. Ae- quin. 1: 101. 1806.
Shrubs or small trees, occasionally with spines to 4 cm long at the nodes, leafy branchlets 1.2-6 mm thick, glabrous to densely villose with hairs to 1 mm long, terete; stipules 2-4 mm long, triangular-subulate with a narrow awn apically, puberulent. Leaves often disti- chous, petioles 3-10 mm long, 0.6-1 mm broad, puber- ulent and often with the hairs restricted to adaxial side; leaf blades (3-)4-7(-10) cm long, 1.5^(-6) cm long, ovate to ovate-elliptic or broadly ovate, apex acute (ob- tuse) to short-acuminate and often slightly curved, base obtuse to rounded and subtruncate, drying thin-charta- ceous, glabrate above, with thin whitish hairs 0.3-0.7 mm long on the lower surfaces, 2° veins 3-6/side, tufted hairs rarely present in the leaf axils. Inflorescences 4-20 cm long, 3-10 cm broad, open panicles, peduncles 2-4 cm long and 1.5-3 mm thick, puberulent, bracts to 7 mm long and liner, peduncles of the lateral branches 5- 40 mm long, flowers closely crowded in subcapitate dis- tal groups of 7 or more, flowers sessile or subsessile. bracteoles ca. 1 mm long. Flowers with hypanthium ca. 2 mm long, turbinate or oblong, minutely puberulent
with thin ascending whitish hairs, calyx lobes 0.5-1 mm long, erect, rounded distally, puberulent; corolla white, 4-5 mm long, tube 1.5-3 mm long, sparsely puberulent or glabrous, lobes 1.5-2.5 mm long, ca. 1.1 mm broad, rounded at the tip; stamens exserted, anthers 0.3-0. 6 mm long; styles 1-3.5 mm long, stigmas 0.3-0.6 mm long. Fruits 4-5 mm long (including the persisting calyx), 1 .5- 2 mm broad, narrowly obovoid or turbinate, minutely (0.1-0.2 mm) puberulent with ascending whitish hairs, splitting at the top into 2 valve-like parts and later sep- arating at the base, persisting central axis 2.5-3.2 mm long.
Plants of lowland evergreen and partly decid- uous forest formations. Flowering in May-No- vember; fruiting in August-September and No- vember. This species is known from southern Mexico, northeastern Guatemala, Belize, and cen- tral Panama southward to Ecuador and Brazil.
Machaonia acuminata is recognized by its oc- casional spines, thin small ovate leaves, small flowers with whitish puberulent ovary, and small
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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fruit. This species has not been found in Honduras, El Salvador, Nicaragua, or Costa Rica. Its close similarity to M. martinicensis and peculiar distri- bution makes one wonder if the two are really different species.
Machaonia martinicensis (EXT.) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 193. 1940. Tertrea martinicensis DC, Prodr. 4: 481. 1830. M. ro- tundata Griseb., Fl. Brit. W. Ind. 348. 1861. M rotundata var. dodgei Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1326. 1938. Figure 37.
Shrubs or (rarely small trees or vines), l-4(-8) m tall, branches occasionally with spines to 35 mm long, leafy branchlets 1.6-5 mm thick, smooth glabrate; stipules 2.5-4 mm long, rounded or triangular at the apex and with a short narrow awn. Leaves opposite (rarely alter- nate), petioles 4-12 mm long, 0.5-1.2 mm broad, gla- brous or puberulent on the adaxial side; leaf blades 4- 9 cm long, 1.8-5 cm wide, ovate to ovate-oblong (ellip- tic-oblong), apex obtuse to acute or short-acuminate, base broadly obtuse or rounded and sometimes decur- rent on petiole, drying thin-chartaceous, glabrous above, glabrous below or with few short (0.2 mm) hairs beneath or with tufts of hairs (domatia) in the vein axils, 2° veins 5-7/side. Inflorescences 6- 11 (-15) cm long, to 14 cm broad in fruit, peduncles 2-6 cm long, ca. 2 mm thick, with whitish hairs 0.2 mm long, bracts 3-5 mm long and linear (or sometimes leaf-like), secondary branches with (secondary) peduncles 1 0-20 mm long, with soft whitish hairs 0.2-0.3 mm long, flowers sessile or subsessile in distal subcapitate clusters of 5-1 5. Flowers with buds 5- 6 mm long, hypanthium 1.5-2.5 mm long, sparsely pu- berulent, calyx lobes 0.7-1.2 mm long, broadly rounded distally; corolla white or pale greenish, tube 1.5-3 mm long, stiff, densely villous at the mouth within, sparsely puberulent externally, lobes 1-2 mm long; stamens 5, anthers ca. 0.6 mm long; style 3.5 mm long, stigmas 0.5- 1 .2 mm long. Fruits 5-7 mm long, 2-3 mm broad, nar- rowly turbinate-oblong, with 2 longitudinal sulci and separating at the base, surface reddish brown or brown, with few hairs (often in lines), persisting axis 4-5 mm long.
Shrubs usually growing near the ocean shore, in mangroves, or along rivers near the ocean in ev- ergreen forest areas of both the Caribbean and Pacific coasts. Flowering in March-April and June- August; fruiting in July-September. The species ranges from southeastern Nicaragua southward to Colombia and also in Jamaica.
Machaonia martinicensis is recognized by its restriction to near-shore environments, small thin- ovate leaves, occasional spines, small flowers in close clusters in an open panicle, and the unusual capsule-like fruit splitting from the bottom. The species has been collected along the Caribbean shore and on the Osa Peninsula in Costa Rica.
Macrocnemum P. Browne
Trees or shrubs, branchlets terete, glabrous or puber- ulent; stipules interpetiolar, oblong to obovate, caducous or persisting with the leaves. Leaves opposite, petiolate; leaf blades with pinnate venation, domatia often present. Inflorescences terminal or axillary, paniculate with op- posite branches, bracteate, flowers in distal cymose groupings, sessile or pedicellate. Flowers bisexual and radially symmetrical, calyx tube very short and cupulate, calyx lobes 5, small, persisting; corolla funnelform or salverform, corolla lobes 5, broadly imbricate in bud and spreading at anthesis, often wider than long, mi- nutely puberulent within; stamens 5, filaments short and villous, anthers oblong; ovary 2-locular, ovules many, peltate and vertically imbricate on the central placenta. Fruits capsular, bisulcate and dehiscing loculicidally into 2 valves; seeds flattened, narrowly elongate with thin wings.
A small genus of about 20 species, ranging from Central America to Colombia and in the West Indies. The broad rounded stipules, broadly over- lapping corolla lobes (in bud), bilocular capsules, and many small imbricate winged seeds help to distinguish this genus.
Macrocnemum glabrescens (Benth.) Wedd., Ann. Sci. Nat. Paris, ser. 4, 1: 76. 1854. Lasionema glabrescens Benth., Bot. voy. Sulph. 105. 1845. Figure 40.
Trees (rarely shrubs) (4-)8-25 m tall, bark brown and the trunk deeply fluted, leafy branchlets 2-7 mm thick, glabrous or minutely (0.1 mm) appressed-puberulent, angular in early stages but becoming terete; stipules (8-) 10-20 mm long, 5-10 mm broad, apex oblong-ob- ovate and rounded, with thin appressed hairs 0.1-0.2 mm long, leaving a scar around the stem above the node. Leaves often with the petioles unequal at the same node, petioles 5-20 mm long, 0.7-1 .8 mm broad, sulcate adax- ially; leaf blades 7-17(-21) cm long, 4-9 cm broad, ob- ovate to obovate-oblong, or ovate-oblong, apex abruptly narrowed and obtuse or short-acuminate apex, tip to 1 cm long, base gradually narrowed and obtuse or acute, drying thin- to stiff-chartaceous, very dark above, gla- brous above, glabrous or minutely appressed-puberulent on the veins beneath, usually with tufts of hairs ca. 1 m long in the vein axils beneath (= domatia), 2° veins 5- 9/side. Inflorescences terminal or axillary, solitary or 3 when the first branching node is subtended by a pair of leaves, 8-28 cm long, to 20 cm broad, paniculate with few opposite lateral branches to 1 2 cm long, peduncles to 1 8 cm long, bracts 3-6 mm long or leaf-like, lanceo- late, flowers in distal cymes or clusters of more than 10, sessile or with pedicels to 3 mm long, bracteoles 0.5-1 .5 mm long. Flowers with a hypanthium 3-4 mm long and ca. 2.4 mm diam., glabrous or minutely puberulent, ca- lyx tube minute, calyx lobes 0.3-1 mm long, broadly triangular; corolla bright rose pink, magenta or the tube becoming maroon, salverform, tube 6-10 mm long, 1 .3-
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FIELDIANA: BOTANY
2 mm diam., with 5 longitudinal ribs, glabrous or rarely sparsely and minutely puberulent, villous at the stamen attachment within, lobes 3-4 mm long, 4-5 mm broad, broadly ovate and rounded distally; stamens 5, filaments 2-4 mm long, villous below the middle, anthers 0.8-1 mm long; style 6-8 mm long, stigma lobes ca. 0.5 mm long, green. Fruits 9-16(-20) mm long, 2-4 mm broad (to 4 mm when split open), narrowly oblong-tubular, with obscure longitudinal ribs, valves opening but re- maining attached at both apex and base, persisting calyx ca. 2 mm broad and 1 mm high (together with disc); seeds 2-3 mm long, 0.5-1 mm broad, with a membra- naceous wing at opposite ends, body of the seed ca. 0.6 mm long.
Trees of evergreen or partly deciduous forest formations from near sea level to 400(-1000) m elevation. Flowering in December-April; fruiting in March-July in Costa Rica and Panama. The species ranges from the evergreen lowlands of southern Costa Rica, through Panama, to Colom- bia.
Macrocnemum glabrescens is recognized by its broadly rounded stipules, usually thin-obovate leaves (often on petioles differing in length at the same node), large open panicles with small bright pink flowers, broadly overlapping corolla lobes, and narrow capsules with opened valves remain- ing attached at the base and apex. This species is only known from near the Hitoy Cerere reserve, Dominical, and the Golfo Dulce region in Costa Rica, but the original description mentions Nicoya and it is possible that it grows in moist forest on the southern part of the Nicoya Peninsula. This species appears to be common in central Panama; see flower description in Croat ( 1 978, p. 8 1 1 ). This species resembles Ferdinandusa panamensis. Standley (1938) listed palo cuadrado as a common name.
Malanea Aublet
Shrubs, woody lianas, or trees, stems glabrous or pu- berulent, terete; stipules interpetiolar, simple, caducous. Leaves opposite, petiolate, entire, pinnately veined, with domatia. Inflorescences axillary, paniculate with short opposite or subopposite lateral branches resembling spikes (or with fasciculate flowers on the central rachis or lateral branches), flowers sessile or subsessile, bracteolate. Flow- ers bisexual and radially symmetrical, small, hypanthi- um turbinate to campanulate, calyx lobes 4, short; co- rolla funnelform to rotate, pale green to white, tube short, throat villose within, corolla lobes valvate or slightly imbricate, villous on the adaxial surface; stamens 4, fil- ament borne on the throat of the corolla, anthers dor- si fixed and partly exserted; ovary 2-locular, ovules sol- itary in each locule and pendulous from the apex of the
locule. Fruits fleshy, oblong or ovoid, 2-locular, exocarp thin-fleshy, endocarp woody or hard; seeds ellipsoid.
A genus of 20-30 species in tropical South America and the West Indies. Malanea colom- biana has been discovered along the Caribbean coast in Belize, in northern Nicaragua, and in southeastern Costa Rica. Malanea erecia Seem, (photo F), said to have been collected by Seemann in Panama, was not treated in the Flora of Pan- ama.
Malanea colombiana Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 66. 1930. Chomelia coclensis Dwyer, Ann. Missouri Bot. Gard. 67: 97. 1980.
Lianas or clambering shrubs with vining branches, leafy stems 1.7-8 mm thick, with closely appressed-as- cending straight hairs 0.4-0.8 mm long, glabrescent and lenticellate; stipules 6-18 mm long, 2.5-8 mm broad, oblong or obovate, with many ascending veins, ap- pressed-puberulent at the base and midrib, deciduous. Leaves with petioles (4-)7-22 mm long, 0.6-2 mm thick, appressed-puberulent; leaf blades 5-13 cm long, 2.5-7 cm broad, ovate-elliptic to ovate or oblong-elliptic, apex obtuse to short-acuminate or acute, base obtuse to slight- ly rounded and subtruncate, drying stiffly chartaceous, dark brown above, glabrous above or with appressed hairs along the midvein and widely scattered elsewhere, with appressed thin whitish hairs 0.3-1 mm long be- neath, often with tufts of hairs .in the vein axils beneath. 2° veins 5-7/side, distal secondaries to arcuate-ascend- ing, 3° veins numerous and closely parallel to form a straight or sinuous pattern between the secondaries. In- florescences 4-13 cm long, 3-8 cm broad, pyramidal with progressively shorter opposite branches, peduncles 2-6 cm long, 0.6-1 .8 mm thick, sericeous, lateral branch- es l-3(-4) cm long, bracts 1.5-4 mm long and linear, bracteoles ca. 1 mm long, flowers sessile or with pedicels to 1 .5 mm long. Flowers 3-4 mm long, hypanthium 0.5- 1 mm long, turbinate, calyx lobes 0.1-0.4 mm long, broadly triangular or poorly developed; corolla white or yellowish white, tube 1.5-2.5 mm long. ca. 1 mm diam. at mouth, lobes 4, 1-2 mm long, 1 mm broad at the base, villose within; anthers 0.5-0.8 mm long; style ex- serted, stigma lobes ca. 0.4 mm long. Fruits 5-6 mm long, 3-4 mm diam., oblong-cylindrical, becoming pur- plish red and drying black, persisting calyx very short (0.5 mm) and not elevated.
Plants of evergreen or partly deciduous vege- tation, 2-200 m elevation. Flowering in June in Costa Rica (Herrera 3081 CR, MO); fruiting in Au- gust in northern Nicaragua (Molina 14926 & 151125 EAP, F). The species ranges from Belize to Colombia.
Malanea colombiana is recognized by its very small flowers with four-lobed perianth, corolla lobes conspicuously villous within, fleshy oblong
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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fruit, unusual tertiary venation, and climbing hab- it. The tertiary veins of the leaf are many and parallel with inconspicuous transverse connec- tions. The Costa Rican collection has minor ve- nation that differs by having a more reticulate ar- rangement of 3° veins with sublineolate 4° veins. More material is necessary to confirm its place- ment under this name. The flowers resemble those of Elaeagia, Figure 39.
Manettia Mutis ex Linnaeus
Herbs or climbers, stems herbaceous or slightly woody, glabrous or puberulent; stipules interpetiolar, small, sometimes adnate to the petiole, triangular to laciniate, persistent. Leaves opposite and decussate, petiolate or subsessile; leaf blades often narrow, pinnately veined, domatia absent. Inflorescences axillary (rarely terminal), cymose to paniculate, fasciculate, umbellate or the flow- ers occasionally solitary, bracts and bracteoles present, flowers pedicellate. Flowers bisexual, monomorphic or distylous, radially symmetrical, small (in Central Amer- ica) to large, hypanthium turbinate, calyx lobes usually
4 (5, 8), short or long, with glands or teeth in the sinuses; corolla funnelform or tubular, white, pink, red, lavender, blue, or yellow, corolla lobes 4(-5), valvate in bud; sta- mens 4, filaments attached in the throat or at the mouth, anthers versatile, exserted or included; ovary 2-locular, ovules numerous and imbricate, vertical on axile pla- centas, style filiform, stigmas 2 clavate or bifid. Fruits thin-walled capsules, obovoid or turbinate to subglo- bose, biloculate and bisulcate, splitting septicidally from the apex into 2 valves; seeds compressed -discoid, usually with a thin wing surrounding the central seed.
A tropical American genus of about 100 species. The slender twining stems, smaller narrow leaves, small inflorescences, small flowers (in Central American species), small obovoid capsules break- ing into two valves, and winged or flattened seeds with erose margin help to distinguish these plants. This study has benefited from the herbarium an- notations of In-Cho Chung and David Lorence. Manettia luteo-rubra variety paraguariensis (Cho- dat) Chung (= M. inflata Sprague) is an ornamen- tal grown for its bright red flowers tipped with yellow.
Key to the Species of Manettia
la. Leaf blades narrowly lanceolate and often subsessile; seeds ca. 1 mm diam., orbicular with an erose margin (the wing minute or absent); flowers white, corolla tube 3-5 mm long M. barbata
Ib. Leaf blades rarely consistently lanceolate, petioles usually well developed; seeds 2-3 mm long, oblong or obicular with a thin expanded marginal wing; flowers white, pink, red, or magenta, corolla tubes 4-13 mm long 2
2a. Corollas white; capsules 4-6 mm long; leaf blades usually puberulent beneath; 1000-1600 m ele- vation M. flexilis
2b. Corolla brilliant red to magenta or pink; capsules 6-10 mm long; leaf blades usually glabrous beneath; 30-1 100 m elevation . . M. reclinata
Manettia barbata Oersted, Vidensk. Meddel. Kjobenhavn 1852: 47. 1853. M. stenophylla J. D. Smith, Bot. Gaz. 56: 58. 1913. Figure 1.
Slender vines to 3 m high, leafy internodes 0.7-2.5 mm thick, glabrous or subglabrous; stipules 0.5-3 mm, long, broadly triangular, glabrous. Leaves with petioles 1-5 mm long, 0.3-1.5 mm broad, glabrous; leaf blades 3-10 cm long, 0.4-3 cm broad, lanceolate to narrowly elliptic-lanceolate or linear-lanceolate, apex tapering gradually and long-acuminate or acute, base obtuse or slightly rounded, drying chartaceous, pale grayish green or grayish brown, glabrous above and below, 2° veins 4- 7/side. Inflorescences 10-15 mm long, axillary cymes of usually 3 flowers, peduncles only 1-4 mm long, bracts to 3 mm long, lanceolate, pedicels 2-5 mm long, brac- teoles 0.5-l(-2) mm long. Flowers with hypanthium 2- 3 mm long, 1 .2-2 mm diam., calyx lobes 1 .2-2 mm long, 0.7-1.2 mm broad, ovate; corolla white, tube 3-6 mm long, 2 mm diam., lobes 2-3 mm long, glabrous exter-
nally and villous within; anthers ca. 1 mm long; stigma ca. 0.5 mm long, oblong to lanceolate. Fruits 4-8 mm long, 3-7 mm broad, persisting calyx to 1.5 mm long and usually recurved; seeds 0.7-1 mm diam., orbicular, wing reduced to a dentate rim around the seed, body of seed ca. 0.6 mm diam., discoid.
Uncommon plants of evergreen forest forma- tions, from 500 to 1300(-2400?) m elevation. Flowering in November-January; fruiting in Jan- uary and March. This species is known only from central and southern Costa Rica.
Manettia barbata is recogized by its narrowly lanceolate leaves, twining habit, few-flowered and short-pedunculate inflorescences, short corolla tubes, smaller capsules, and circular-discoid seeds with little evidence of a wing.
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FIELDIANA: BOTANY
Manettia flexilis Brandegee, Univ. Calif. Publ. Bot. 6: 196. 1915. M. estrellae Standl., J. Wash. Acad. Sci. 15: 6. 1925. Figure 1.
Vines to 4 m high, leafy stems 0.7-2 mm long, pu- berulent with curled whitish hairs ca. 0.2 mm long; stip- ules 1-2 mm long, adnate to the petiole base, rounded distally or with short (0.3 mm) thick spike-like projec- tions. Leaves with petioles 2-10 mm long, 0.3-0.7 mm thick, puberulent; leaf blades (2.5-)3-7 cm long, 1-3 cm broad, oblong to ovate or lanceolate, apex acute to long- attenuate, base obtuse to acute, drying membranaceous or thin-chartaceous, dark green above, pale grayish green beneath, glabrous or puberulent above, short pilose be- neath with thin whitish hairs ca. 0.3 mm long, 2° veins 4-6/side, the upper secondaries strongly arcuate-ascend- ing. Inflorescences axillary, cymose or umbelliform flow- ers few (rarely 1), peduncles 1-7 mm long, bracts ca. 2 mm long, flowers closely crowded, pedicels 1-5 mm long. Flowers with hypanthium 2-3 mm long, densely puber- ulent with straight or curved multicellular hairs 0.2-0.5 mm long, calyx cup to 0.4 mm long, calyx lobes (2-)3- 4 mm long, 0.8-1.5 mm broad, lanceolate, with erect glands ca. 0.3 mm long in the sinsues; corolla white or rose red, tube 4-10 mm long, ca. 1.3 mm diam., short villose externally, glabrous within, lobes 4, 2-5 mm long, ca. 0.7 mm broad, ovate to triangular, densely villous externally; stamens 4, anthers ca. 0.7 mm long. Fruits 5-7 mm long, 4-6 mm broad, broadly obovoid to subglo- bose, rounded and truncated at the apex, with few scat- tered hairs; seeds 2-3 mm long, oblong and with a thin translucent wing, body of the seed ca. 1 mm long and oblong.
Plants of evergreen and partly deciduous forest formations in the central Cordilleras, from ca. 1 000 to 1 600 m elevation. Probably flowering and fruit- ing throughout the year. The species ranges from southern Mexico to western Panama.
Manettia flexilis is recognized by its smaller cap- sules, winged seeds, puberulent stems and leaves, and mid-elevation habitats. This species is infre- quently collected.
Manettia reclinata Mutis in I ... Mant. PI. 2: 558. 1771. Nacibea coccinea Aubl., Hist. pi. Guiane 96, t. 37, f. 1. 1775. M. coccinea (Aubl.) Willd. in L., Sp. PI. ed. 4, 1: 624. 1797. M. cuspidata Bertero in Spreng., Syst. Veg. 1:415. 1825. M. panamensis Duchass. & Walp., Linnaea 23: 753. 1850. M. costaricensis Wernham, J. Bot. 56, suppl. 1: 38. 1919. M. orbifera Wernham, loc. cit. 41. 1919. M, seleriana Loes., Verhand. Bot. Vereins Brandenb. 65: 107. 1923. Figure 1.
Vines to 4 m high, stems often with 4 narrow longi- tudinal ridges, leafy stems 0.7-2.5 mm thick, sparsely puberulent (rarely glabrous), often with a row of minute (0. 1-0.2 mm) hairs along the longitudinal ridges; stipules
1-2 mm long, broadly triangular, minutely puberulent. Leaves with petioles 2-14(-20) mm long, 0.6-1.2 mm broad, sparsely puberulent; leaf blades (2-)3- 10 cm long, l-3(-5) cm broad, narrowly ovate to ovate-elliptic or lanceolate, acute or acuminate at the apex, obtuse to acute at the base, leaves drying thinly chartaceous and dark green above, paler greenish gray beneath, usually glabrous above and below, with 4-6 major secondary veins strongly ascending on each side. Inflorescences ax- illary or terminal, usually solitary with 2-4 flowers per node, peduncles 0.5-2.5 cm long, puberulent, pedicels 10-25 mm long (to 35 mm in fruit). Flowers with hy- panthium 3-5 mm long, oblong, calyx tube to 1 mm long, calyx lobes 4-8, unequal. 4-7 mm long, 0.3-1.3 mm broad, linear to lanceolate; corolla red, rose red. deep pink, scarlet, or magenta, tube 6-13 mm long, 2- 2.5 mm diam., glabrate to densely minutely puberulcnt externally, with yellowish hairs in throat, corolla lobes 4 (5, 6), 2-4 mm long, 2.5-4.5 mm wide, ovate, puber- ulent externally and glabrous within; stamens 4 (5. 6), filaments ca. 1 mm long, anthers ca. 3 mm long; style ca. 1 1 mm long. Fruits 6-10 mm long, 6-9 mm broad, obovoid or subglobose, glabrous or pubescent with thin curled hairs to 0.5 mm long, with 8 longitudinal ribs (2 in the sulci), calyx lobes persisting and recurved; seeds 2-2.5 mm diam., thin-discoid and orbicular, with an erose translucent wing around the margin, body of the fruit 0.7-1 mm diam.
Plants of wet evergreen forest formations, 0- 1300 m elevation. Flowering in October-May; fruiting in December-May. The species ranges from southern Mexico and the West Indies into north- ern South America.
Manettia reclinata is recognized by its larger red or pink flowers, few flowers per node, usually gla- brous leaves, larger rounded capsules borne on long pedicels, flat orbicular seeds with thin winged margin, and vining habit. This species appears to be much more common than its congeners in Cos- ta Rica.
Mitracarpus Zuccarini
Annual or perennial herbs, erect or decumbent, stems tctragonous in cross-section, glabrous or puberulent; stipules adnate to the petiole base to form a sheath with 3-15 slender distal setae (awns), persistent. Leaves op- posite and decussate, subsessilc to short-petiolate, leaf blades usually narrow (linear to ovate), chartaceous, domatia absent. Inflorescences axillary or terminal, cap- itate or glomerulate with densely crowded flowers, the heads sometimes subtended by 4 leaf-like bracts, flowers sessile or subsessile. Flowers bisexual and monomor- phic, very small, hypanthium turbinate to subglobose. calyx tube short, calyx lobes 4(-5), unequal, persistent; corolla white, salverform or funnelform. tube usually with a ring of hairs within, glabrous or villous in the throat, corolla lobes 4 (3), valvate in bud; stamens 4, inserted on the throat, anthers oblong to linear, dorsi- fixed, included or exserted; ovary 2-3-locular, with 1
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ovule in each locule attached to a peltate placenta in the center of the septum, style short or long, with 2 short linear stigmas. Fruits thin-walled capsules, 2- or 3-locular with circumscissile or transverse dehiscence from below the middle (the upper portion breaking away with the persisting calyx and exposing the seeds), septum per- sisting with the basal part of the capsule; seeds ellipsoid to oblate or globose, ventral (adaxial) surface with 4 sulci radiating from a central area to give an X-like pattern demarking 4 broadly rounded areas, abaxial surface smooth.
A Neotropical genus of 30-45 species, with most in Brazil. The genus is very similar to Crusea, Diodia, and Spermacoce with its weedy-herba- ceous habit, broad stipules with distal awns, nar- rowly elliptic leaves, small flowers in verticillate heads, and two-locular capsules with two seeds. Mitracarpus is unusual in the circumscissile de- hiscence of the capsule and the X-like (cruciform) sulcus on the inner face of the seed.
Mitracarpus hirtus (L.) DC., Prodr. 4: 572. 1830. Spermacoce hirta L., Sp. PI. ed. 2: 148. 1762. S. villosa Sw., Prodr. 29. 1788. M. villosus (Sw.) Cham. & Schlend., Linnaea 3: 363. 1828. M. breviflorus Gray, PI. Wright. 2, 68. 1853. Fig- ure 4.
persisting calyx and top of fruit 2-3.5 mm long, body of the yellowish brown capsule ca. 1 mm long, 0.6-1 mm broad; seeds 0.5-0.8 long and ca. 0.6 mm wide, oblong- oblate, yellowish brown, with 4 sulci forming an im- pressed X on the adaxial face, smooth or pitted on the abaxial face.
Common weedy plants of open disturbed sites in both deciduous and evergreen formations, 0- 1200(-1400) m elevation. Flowering primarily in June-December in Costa Rica. The species is found in the southwestern United States and throughout tropical America; it has become established through much of tropical Africa and has been found in India, Burma, and the western Pacific.
Mitracarpus hirtus is recognized by its herba- ceous weedy habit, setose sheathing stipules, nar- row hispidulous leaves with few secondary veins, axillary heads of small congested flowers, and un- usual fruit and seed. It is found in both the Ca- ribbean and Pacific lowlands and the central up- lands. There has been considerable confusion regarding the nomenclature of this species and es- pecially the applicability of the names; we follow the recent annotations of C. Dennis Adams. These plants can be mistaken for species of Spermacoce, Diodia, and Hyptis (Labiatae).
Herbs to 0.6(-1) m tall, erect or spreading, stems usu- ally simple distally and with few branches mostly near the base, often slightly woody at the base, leafy stems 0.7-2.5 mm thick, with 4 longitudinal ridges and some- what quadrangular in cross-section, usually with curved thin hairs 0.2- 1.5 mm long and longer (1.5-2 mm) trans- lucent hairs at the nodes; stipules united to the petiole base to form a broad sheath l-3(-4) mm long, the straight distal margin of the sheath with 6-9(-l 3) linear setae 1- 5 mm long. Leaves subsessile, petioles 0-3 mm long, not clearly differentiated from the base of the blade; leaf blades 2-5(-8) cm long, 0.5-1. 5(-2) cm broad, narrowly elliptic to lanceolate, or narrowly elliptic-oblong, apex tapering gradually and acute, base cuneate, drying stiffly chartaceous, glabrescent or sparsely hispidulous above, sparsely hispidulous beneath with hairs ca. 0.3 mm long and along the margin, 2° veins 2-3/side, strongly as- cending. Inflorescences axillary or terminal, flowers densely crowded in capitulae or verticils 4-10 mm high and 8-15(-20) mm broad, bracts 2-4 mm long but dif- ficult to distinguish among the sepal lobes in the tightly congested heads, flowers sessile or subsessile. Flowers with hypanthium 1-2 mm long, glabrous beneath and hispidulous distally, calyx lobes 4, unequal, the 2 larger 1.5-2.5 mm long, lanceolate, thick centrally and with a hyaline margin; corolla white, glabrous or minutely hairy externally, tube 1.5-2 mm long, 0.3-0.9 mm diam., lobes 4, 0.6-1.1 mm long. 0.4-1 mm broad at the base, ovate or triangular; stamens 4, filaments very short or absent, anthers 0.4-0.7 mm long, usually exserted; style 1-1.6 mm long, stigma 0.3-0.5 mm long. Fruits with the top part coming off to expose the 2 locules and 2 seeds,
Morinda Linnaeus
Small to large trees or shrubs (lianas), stems terete or quadrangular in cross-section, glabrous or puberulent; stipules interpetiolar and sometimes slightly connate to form a short sheath above the petioles (intrapetiolar), entire to cuspidate or bifid, glabrous. Leaves opposite or 3/node (sometimes only 1 at a flowering node), petiolate, pinnately veined, often slightly succulent in life, usually with tufts of hairs (domatia) in the vein axils beneath. Inflorescences terminal or axillary, with 1 or 2 (rarely more) capitate heads on a common peduncle (the heads rarely sessile), large or small bracts sometimes present, flowers sessile in the capitulum with the basal parts usu- ally united. Flowers bisexual (rarely unisexual), radially symmetrical, usually distylous, hypanthium free or unit- ed with other flowers, calyx tube urceolate or hemi- spheric, often entire distally, calyx lobes usually minute or absent; corolla funnelform or salverform, corolla tube glabrous or puberulent in the throat, puberulent at the base within, corolla lobes 4-7, narrow and valvate in bud; stamens 4-7, filaments short and inserted in the throat of the corolla, anthers dorsifixed and versatile, included or exserted, connective often prolonged distal- ly; ovary 2- or 4-locular 1 ovule in each locule, ovules erect and attached below the middle or at the base of the septum, style slender, with 2 short or long style branches. Fruits fused into a fleshy syncarp made up of the united ovaries (or their bases), often large and turning white, pyrenes 1 -seeded or part of a 2-4-locular woody structure; seeds obovoid or reniform.
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A pantropical genus of 50-80 species; most of the species are native to the Old World. The flow- ering capitula with the basal parts of the flowers united, and the latter forming a fleshy largely syn- carpous fruiting capitulum, make this genus very distinctive among our Rubiaceae. Three species have been recorded from southern Central Amer-
ica; two additional native species (M. asperula Standl. and M. yucaianensis Greenm.) are found in northern Central America. There is consider- able variation within our species, and this can make their identification difficult. Current holdings of this genus are very limited; all the Costa Rican material is from near the seashore.
Key to the Species of Morinda
1 a. Leaf blades usually less than 5 cm broad, usually drying grayish or yellowish; branches often scandent M. royoc
Ib. Leaf blades more than 5 cm broad, usually drying very dark in color; branches never clambering or scandent 2
2a. Inflorescences usually solitary, with only 1 capitulum per peduncle, fruiting syncarps 3-1 2 cm diam.; largest leaf blades usually more than 20 cm long and 14 cm broad; only found near the ocean shore M. citrifolia
2b. Inflorescences 1-3/node, with 1-3 capitula per primary peduncle, fruiting syncarp 0.5-3 cm diam.;
largest leaf blades usually less than 20 cm long and 14 cm broad; lowland rain forests
M. panamensis
Morinda citrifolia L., Sp. PI. 176. 1753. Figure 19.
Shrubs or small trees. (l-)2-8(-12) m tall, trunks to 1 5 cm thick, wood yellow, branchlets quadrangular or terete, leafy stems 2.5-12 mm thick, glabrous; stipules 6-20 mm long. 5-14 mm broad, oblong to suborbicular- triangular and rounded distally, glabrous. Leaves well spaced along the stem, petioles 12-20 mm long, ca. 2 mm broad, glabrous, with slightly winged adaxial mar- gins; leaf blades 12-28(-40)cm long, 7-16(-24)cm broad, oblong to broadly elliptic-oblong or ovate-oblong, apex obtuse to acute (very short acuminate), base broadly obtuse to cuneate, drying chartaceous or thin-charta- ceous, often dark in color, glabrous above and below but with tufts of hairs in the vein axils beneath, 2° veins 6- 8/side. Inflorescences axillary and drying black, solitary or 2-3/node, capitulae 9-20 mm long, to 20 mm diam., oblong to subglobose, peduncles 10-22(-30) mm long, 1.2-2 mm thick, glabrous, flowers sessile and united at the base. Flowers united basally (hypanthia connate), calyx tube minute and truncated with a scarious margin: corolla white, slightly thickened, glabrous externally, tube 6-10 mm long, cylindrical, corolla lobes usually 5, 3-8 mm long, ca. 1.5 mm wide, obtuse, thick-fleshy; stamens 4-6, filaments slightly unequal, anthers to 5 mm long, becoming twisted; stigma to 5 mm long and 0.8 mm wide, erose. Fruits united into a syncarp. 4-1 2 cm diam., fleshy succulent and irregularly globose to oblate, white with green "eyes" ca. 8 mm diam. formed by the calyx and disc of individual flowers, pyrenes to 10 mm diam.; seeds 3.5 mm long.
Plants usually found near ocean beaches and lagoons along the Caribbean shore from Honduras to Panama, 0-20 m elevation. Flowering and fruit- ing throughout the year (Sanchez 1983). The orig-
inal range of this species was from India to the East Indies and northern Australia. It has become naturalized in a number of areas around the Ca- ribbean.
Morinda citrifolia is recognized by its large white Annona-\\ke syncarps, large opposite leaves on thick stems, and seaside Caribbean habitat. Her- barium specimens can be difficult to distinguish from M. panamensis. Yema de huevo is a common name; the fruits are edible.
Morinda panamensis Seem., Bot. voy. Herald 1 36. 1854. Figure 19.
Shrubs or trees. 3-25 m tall, branchlets quadrangular in cross-section, leafy stems 1.5-6 mm thick, minutely (0.1 mm) farinose puberulent. glabrescent; stipules ap- parently with the distal part tearing off to leave a cupulate base 1-3 mm long forming a shallow cup around the stem, at first broadly oblong (to 10 mm long and 9 mm broad) and covering the shoot apex. I^eaves with petioles 5-25 mm long. 1-2 m thick, glabrous or minutely (0.1 mm) puberulent; leaf blades 9-21 cm long, 4-13 cm broad, broadly elliptic to elliptic-oblong or ovate-oblong, apex bluntly short-acuminate with tip to 10 mm long, base obtuse or cuneate, drying thin-chartaceous, dark, glabrous above, glabrous or minutely (0.2-0.3 mm) pu- berulent beneath and with dense tufts of hairs (domatia) in the vein axils. 2° veins 4-7/side. central 2° veins 1-4 cm distant. Inflorescences 1-3/node, capitate, peduncles 4-35(-55) mm long, 0.7-1.5 mm thick, the primary pe- duncle simple or with 3-4 equal or unequal branches to 3 cm long, usually glabrous, each capitulum 4-10 mm
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long and 5-10 mm wide, oblong to globose, with 9 flow- ers or more, flowers sessile and united. Flowers united below, glabrous externally (rarely minutely puberulent) and drying black, hypanthium partly free distally for 1- 2 mm, calyx tube ca. 0.3 mm long and subentire; corolla white, tube 5-10 mm long, lobes 4, 3-6 mm long, 1-2 mm broad, narrowly oblong; stamens 4, anthers 2.5-3.5 mm long; stigmas 3.5-5 mm long. Fruit a syncarp, 15- 30 mm diam., subglobose, the calyx tube little (0-0.5 mm) elevated above the surface of the fruit and 2.5-4 mm diam.
Plants of wet evergreen lowland rain forest for- mations, 0-600 m elevation. Flowering in March- May in Central America; fruiting in July-Septem- ber. This species ranges around the Gulf of Mexico and the Caribbean from Florida (U.S.A.), Mexico, Central America, and the West Indies to Panama.
Morinda panamensis is recognized by its un- usual capitula of flowers with united ovaries, seeds imbedded within a globose syncarp, thin leaves drying very dark, and rain forest habitat. Some herbarium specimens can be very similar to spec- imens of M. citrifolia, and it is possible that there are intermediates in nature. An unusual collection from near Upala (Herrera 1783 CR, MO) is mi- nutely puberulent on all parts.
Morinda royoc L., Sp. PI. 176. 1753. Figure 19.
Shrubs, vines, or small trees, l-3(-7) m tall, often with scandent branches, leafy branchlets 1-5 mm thick, young stems with obscure longitudinal ridges and quadrangular or terete, minutely puberulent with thin hairs 0. 1 mm long; stipules l-2(-4) mm long, 3-5 mm wide at the base, triangular or with an awn 1-2 mm long, glabrous and drying yellowish brown. Leaves with petioles 4-14 mm long, 0.5-1 mm broad, with adaxial margins, gla- brous or minutely puberulent; leaf blades 4-1 1(-13) cm long, 1-4.5 cm broad, narrowly elliptic-oblong to oblan- ceolate or linear-oblanceolate, usually 4 times longer than broad, apex acute to short-acuminate, base gradually narrowed and acute or attenuate, drying stiffly charta- ceous, grayish or yellowish, with the margin often in- volute, glabrous above and below but often with barbate hairs in the leaf axils and along the midvein, 2° veins 3- 6/side, arcuate and weakly loop-connected near the mar- gin. Inflorescences solitary and axillary, the capitulae 4- 12 mm diam., usually oblong, sessile or with peduncles 0-7(-10) mm long, minutely puberulent, flowers sessile and united. Flowers united together in the lower half of their ovary, free portion of the hypanthium and calyx 1-3 mm long, calyx entire distally (or obscurely 5-lobed), minutely puberulent; corolla white, 6-8 mm long, tube ca. 5 mm long and 2 mm diam., cylindrical, minutely (0. 1 mm) puberulent externally, lobes 1-2 mm long; sta- mens 5-6, anthers ca. 2 mm long; style 3-5 mm long, stigmas ca. 1.5 mm long. Fruit a syncarp, 8-25 mm diam., irregularly globose, pyrenes 5 mm long.
Plants of the Caribbean lowlands (from pine savannas to evergreen forest formations) in Cen- tral America. Flowering in January-August. The species ranges from Florida (U.S.A.), Mexico, Central America, and the West Indies to northern South America.
Morinda royoc is recognized by its vining branches, smaller narrow leaves that rarely dry very dark, small syncarpous capitulae, and small stipules. We have seen no specimens from Costa Rica, but the species has been collected near Blue- fields, Nicaragua, and in central Panama. Most Central American collections are described as vines. Appunia guatemalensis can appear quite similar, but the branches are not vining and the fruit develop separately.
Mussaenda Linnaeus
Shrubs, erect or climbing; stipules interpetiolar, united and solitary or separate and paired (4/node). Leaves op- posite, petiolate, pinnately veined and often acuminate at the apex, domatia absent. Inflorescences terminal, cy- mose, bracts and bracteoles deciduous; a few flowers of the inflorescence with a single greatly expanded colorful and leaf-like expanded sepal lobe. Flowers bisexual and radially symmetrical (except for those flowers where 1 sepal lobe is greatly enlarged), calyx tube turbinate or ovoid, calyx lobes 5, 1 lobe greatly expanded in a few flowers of most species; corolla narrowly funnelform or salverform, glabrous or puberulent on the outer surface, corolla tube puberulent in the throat, corolla lobes 5; stamens 5, filaments very short, borne near the base or the upper part of the tube, anthers sagittate at the base; ovary 2-locular with many ovules, style 2-branched. Fruits usually fleshy and indehiscent (rarely dry and loculici- dally dehiscent); seeds small and ellipsoid.
A tropical Old World genus of about 200 spe- cies. The greatly enlarged and colorful sepal lobes on a few flowers of each inflorescence are found in most species of the genus and account for the ornamental appeal of the following species.
Mussaenda erythrophylla Schumach. & Thonn., Beskr. Guin. PI. 116. 1827. Figure 16.
Shrubs or woody climbers, 1.5-3(-8) m tall, leafy branchlets 2-6 mm thick, densely velutinous with yel- lowish hairs ca. 1 mm long, terete; stipules 4/node, 6- 1 1 mm long, ovate-triangular, glabrous on the inner face. Leaves with petioles 4-18 mm long, with hairs to 1.5 mm long; leaf blades 4-12(-15) cm long, 3-7 cm long, ovate to broadly ovate-elliptic or ovate-orbicular, apex short-acuminate, base obtuse or rounded and subtrun- cate, densely puberulent on both surfaces with slender
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hairs 0.5-2 mm long, 2° veins 5-7/side, arcuate-ascend- ing. Inflorescences ca. 10 cm long and 20 cm broad, at the ends of distal un branched leafy stems, with opposite or trichotomous branches, densely velutinous. Flowers densely pubemlent externally, calyx lobes 6- 1 2 mm long, 1-3 mm broad, red; the enlarged sepal lobe leaf-like, 4.5-6.5 cm long, 3-6.5 cm broad, on petioles 2-8 mm long, with ca. 7 palmate veins, bright red; corolla yellow to pink, tube 15-30 mm long, lobes 3-9 mm long, 4-7 mm broad, broadly ovate, whitish and papillate puber- ulent within.
Popular small shrubs or climbers grown for or- nament in parks and gardens. The enlarged leaf- like sepal lobes are bright crimson to deep red and give a very colorful effect. They flower throughout the year but usually do not produce fruits.
4 cm diam., peduncles 2-4 cm long, flowers many and closely congested. Flowers not conivent basally; corolla 7-8 mm long, narrowly funnelform, lobes 5, ca. 1.5 mm long; style long-exserted. Fruits tightly congested in the spherical heads, splitting into 4 parts, seeds minute.
The solitary pedunculate globose heads and larger drooping leaves distinguish this introduced species. It is fast-growing in open sites until it reaches about 10 m in height. Common names used in southeast Asia are kadam, kedam. and laran. The literature of this tree is to be found under Anthocephalus, now a synonym ofBreonia. See J. D. E. Fox, Anthocephalus chinensis, the La- ran Tree of Sabah. Econ. Bot. 25: 221-233. 1971.
Neoiamarckia Bosser
Nertera Banks & Solander Nomen conservandum
Trees; stipules interpetiolar, triangular, deciduous. Leaves opposite, petiolate, entire, domatia absent. In- florescences terminal, usually solitary, capitate and pe- dunculate, flowers densely congested. Flowers bisexual and radially symmetrical, calyx lobes small; corolla fun- nelform, corolla lobes 4—5, imbricate in bud; ovary 2- locular at the base, placentation axile with many vertical ovules; stigma fusiform. Fruits capsules, loculicidally de- hiscent from the apex, thin-walled; seeds 1-5 in each locule, small and angular.
A genus of two species of southeast Asia. One species is occasionally planted in Costa Rica.
Neoiamarckia cadamba (Roxb.) Bosser, Bull. Mus. Hist. Nat. Paris 4 ser. sect. B. Adansonia 6: 247. 1984. Nauclea cadamba Roxb., Fl. Ind., ed. 1, 2: 121. 1824. Anthocephalus cadamba (Roxb.) Miq., Fl. Ind. Bat. 2: 135. 1856. A. morindae- folius Korth., Verh. Nat. Gesch. 1 54, t.. 48. 1 842. A. indicus A. Rich., Mem. Soc. Hist. Nat. Paris 5: 238. 1834, nom. illeg. Cephalanthus chinensis a uctt.. non Lamarck; A. chinensis auctt.
Trees, 5-15(-30) m tall, fast-growing in early stages, leafy stems 2.3-10 mm thick, glabrous, quadrangular; stipules oblong, covering the buds, caducous. Leaves usually somewhat pendant, deciduous, petioles 22-55 mm long, 1-4 mm thick, glabrous or minutely puberu- lent, drying dark; leaf blades (7-) 13-3 4 cm long, (5-)6.5- 18 cm broad, ovate to ovate-oblong or oblong, apex abruptly narrowed and blunt or short-acuminate, base obtuse to rounded and truncate or subcordate, drying brownish, glabrous above, glabrous or minutely (0.05 mm) puberulent beneath, 2° veins (4-)8- 1 6/side, 3" veins subparallel. Inflorescences solitary, terminal on short lat- eral branchlets, each with a single globose capitulum 3-
Perennial herbs, creeping and repent, glabrous or sparsely puberulent, much-branched and often rooting at the nodes (often forming mats); stipules interpetiolar, small, partly united to the petiole bases, entire or with 2 teeth, persistent. Leaves opposite, very small, petiolate or sessile; leaf blades ovate to rounded, glabrous or pu- berulent, venation pinnate or subpalmate; domatia ab- sent. Inflorescences of solitary flowers, axillary or ter- minal, the flowers sessile or subsessile. Flowers bisexual, radially symmetrical, hypanthium ovoid to turbinate, calyx tube truncated and entire or slightly lobed; corolla broadly funnelform to tubular, corolla lobes 4-5, valvate in bud; stamens with thin filaments attached to the base of the tube, anthers basifixed, the connective apiculate. exserted; ovary 2-locular, with 1 ovule borne from the base of each locule, style deeply 2-branched and slender. Fruits fleshy drupes (rarely dry), rounded, with 2 plano- convex pyrenes.
A genus of 6- 1 2 species found in Australia, New Zealand, Malaya, Indonesia, South China, and some Pacific islands; in addition, a single species ranges from Mexico through the higher moist mountains of Central and South America to Chile.
Nertera granadensis (Mutis ex L.f.) Druce, Bot. Soc. Exch. Club Brit. Isles 1916: 637. 1917. Go- mozia granadensis Mutis ex L.f., Suppl. PI. 1 29. 1781. Nertera depressa Banks & Solander ex Gaert.. Fruct. et Sem. PI. 1 : 1 24. 1 788. Figure 3.
Creeping prostrate or pendant herbs to 5 cm tall and up to ca. 1 m long, terrestrial or low epiphytic, often forming mats, stems 0.3-1 mm thick and rooting at many nodes, often tetragonous in cross-section in life, glabrous or with a few scattered hairs; stipules 0.3-1 mm long, triangular, united at the base with the bases of the pet-
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ioles, persisting. Leaves quite variable on different plants, petioles 0.7-6(-9) mm long, 0.2-0.4 mm broad, gla- brous, sometimes sulcate above; leaf blades (1.5-)2.5- 8(-13) mm long, (1.2-)2-7(-12) mm broad, ovate-tri- angular to ovate-orbicular or ovate-deltoid, apex obtuse to rounded, sometimes with a minutely apiculate tip, base rounded and truncate to obtuse, often decurrent on petiole in larger leaves, drying thin-chartaceous but semisucculent in life, glabrous above and below (rarely puberulent), 2° veins 2-4/side, strongly ascending. Inflo- rescences absent, the solitary sessile flowers axillary to distal leaves, the flowers covered by surrounding leaves and very difficult to see in pressed herbarium material. Flowers minute (ca. 4 mm long), usually glabrous exter- nally, hypanthium 0.5-1 mm long, calyx lobes absent; corolla 1-3 mm long, greenish to yellow or white, tube 0.6-1 mm long, widely funnelform (subcampanulate), lobes ca. 0.5 mm long, glabrous; anthers 0.3-0.5 mm long. Fruits sessile, 4-7 mm diam., globose, bright or- ange, orange-red, or deep red, fleshy part often translu- cent, pyrenes 2-2.5 mm long and 1.2-2 mm wide, ob- long.
Plants of wet evergreen montane forest forma- tions, from (1000-)! 500 to 3400 m elevation in Central America. Fruiting throughout the year in Costa Rica. This species ranges from Mexico, through the high mountains of Central America and South America, as far south as Chile. This species, when interpreted in a broad sense, is said to be found throughout the range of the genus.
Nertera granadensis is a very distinctive species with its low-creeping and mat-forming habit, very small paired leaves, lack of pubescence, solitary little flowers, bright orange-red berries, and re- striction to higher-elevation wet forest habitats. Collections with larger (10 mm) leaves can look very different from those with smaller (4 mm) leaves. It is common in some montane rain forests, covering mossy banks and old logs. In Costa Rica it seems to be restricted to the central Volcanic Highlands and the Cordillera de Talamanca. The brightly colored berries and mat-forming ability have made this species valuable as a greenhouse or moist-area ornamental. These plants resemble small collections of Didymaea.
Oldenlandia Linnaeus
REFERENCES— E. E. Terrell, Synopsis of Olden- landia (Rubiaceae) in the United States. Phyto- logia 68: 125-133. 1990. E. E. Terrell and W. H.
Lewis, Overview and annotated list of North American species of Hedyotis, Houstonia, Olden- landia (Rubiaceae) and related genera. Phytologia 71: 221-243. 1991. D. A. Halford, Review of the genus Oldenlandia L. (Rubiaceae) and related gen- era in Australia. Austrobaileya 3: 683-722. 1992.
Annual or perennial herbs (rarely subshrubs), stems erect or decumbent, simple or branched, glabrous or puberulent; stipules interpetiolar, small, acute to acu- minate, often united to the base of the petioles to form a short sheath, with 1 -several awns. Leaves opposite and decussate, sessile or short-petiolate; leaf blades usually narrow, often with a stiff mucronate tip, domatia absent. Inflorescences axillary or terminal, open and branched panicles and cymes or the flowers solitary or fasciculate in the leaf axils, flowers sessile to long-pedicellate. Flow- ers bisexual, monomorphic or distylous, usually small, hypanthium turbinate to hemispheric, calyx lobes (3-)4(-5-8), equal, narrowly to broadly triangular; co- rolla rotate or salverform to funnelform, white to lav- ender, pink, or purple, tube cylindrical, the throat often puberulent, corolla lobes (3-)4(-5), valvate in bud; sta- mens (3-)4(-5), anthers dorsifixed and sessile or on short filaments inserted on the throat, included or exserted; ovary 2-locular, usually with many horizontal ovules on peltate placentas attached near the base of the septum, style filiform, stigmas 2, linear to subglobose. Fruits cap- sular, often papery, globose to oblong, usually with a loculicidally dehiscent apex (beak), later also septicidally dehiscent; seeds usually many (50-100), angular to subglobose, smooth to reticulate or alveolate, often be- coming viscid when moistened.
A pantropical and subtropical genus with prob- ably ca. 100 species but with problems regarding generic circumscription. Some authors have sug- gested placing this genus under Hedyotis or Hous- tonia, while others divide it into smaller genera; see the references cited above. Our plants of this genus are recognized by their delicate herbaceous habit, the very small or linear-lanceolate leaves, the axillary flowers or small few-flowered inflo- rescences, the minute flowers on filiform pedicels, and the broadly rounded thin-walled capsules with small seeds.
Many Costa Rican collections formerly identi- fied as O. corymbosa, O. herbacea, and O. land- folia are probably the same species: O. corymbosa. This conclusion is based on an overview of the material and following the keys and descriptions of Verdcourt in the Flora of Tropical East Africa (1976).
Key to the Species of Oldenlandia
la. Leaf blades less than 5 mm long, usually broadly ovate; plants often forming small mats (not yet collected in Costa Rica) O. callitrichoides
196
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1 b. Leaf blades more than 5 mm long, narrowly elliptic-ovate to lanceolate or linear; plants diffuse or foming loose mats 2
2a. Corolla 2-1 1 mm long; fruit with a beak 0.8-1 mm long (not yet known to occur in Costa Rica) O. herbacea
2b. Corolla 0.5-2 mm long; fruit with a small or well-developed beak 3
3a. Flowers pink or white marked with lavender, flowers often 2-3/peduncle; fruit globose to globose- oblate, rounded at the base but not saccate, with a beak 0.1-0.6 mm high O. corymbosa
3b. Flowers white, flowers usually solitary; fruit broader than long and distinctly rounded (saccate) at the base, with a beak ca. 1 mm long O. lancifolia
Oldenlandia callitrichoides Griseb., Mem. Am. Acad. 2, 8: 506. 1863. Oldenlandiopsis callitri- choides (Griseb.) Terrell & W. H. Lewis, Brit- tonia42: 185. 1990.
Prostrate herbs to 10 cm tall, much-branched and of- ten forming mats, leafy stems 0.1-0.5 mm thick (when dried), glabrous; stipules adnate to petiole base and form- ing a short sheath to 0.5 mm long, sheath entire distally or with a small appendage, glabrous. Leaves very small, petiole 0.5-2 mm long; leaf blades 1-3.5 mm long, 1-3 mm broad, rounded-triangular to ovate, apex bluntly obtuse or rounded, base broadly obtuse to rounded and subtruncate, somewhat decurrent on petiole, drying membranaceous, greenish, glabrous above and below (or with a few broad-based hairs ca. 0.2 mm long), 2° veins 3-4/side, strongly ascending. Inflorescences of solitary flowers in distal leaf axils, usually only 1 flower per node, pedicels 1-9 mm long, filiform, glabrous. Flowers 2-3 mm long, hypanthium 1-1.5 mm long, calyx lobes 0.2- 0.6 mm long. Fruits 2-3 mm long (including the calyx lobes), ca. 1 mm diam., distally truncated, with a few whitish raphides on the greenish surface, calyx lobes ca. 0.5 mm long.
Distinctive little plants usually found on sandy soil in open sunny situations, in tropical lowland sites. This species has the smallest leaves found among Central American Rubiaceae. In general aspect, these plants resemble Pilea hernariarioides (Sw.) Weddell and P. microphylla (L.) Liebm. This species is known from the Yucatan peninsula and from central Panama, but it has not been collected elsewhere in Central America. Terell and Lewis based a new genus on this distinctive species (see synonomy).
Oldenlandia corymbosa L., Sp. PI. 1 19. 1753. Fig- ure 3.
liptic-oblong, apex abruptly obtuse to acute and with a minute (0.2 mm) apiculate tip, narrowed to the base, drying chartaceous and with the margins usually revo- lute, dark above, glabrous or minutely scabrid above and below (or with the cystoliths appearing like minute tri- chomes), 2° veins usually obscure. Inflorescences axil- lary, with single flowers on slender pedicels or cymose- umbellate with 3 (2-5) flowers on a common filiform peduncle 2-8 mm long, both singular and umbellate in- florescences often present on the same stem or node, pedicels 2-6(-12) mm long, filiform and glabrous. Flow- ers very small, hypanthium 0.5-1 mm long, obconical to cupular, glabrous, calyx lobes 4, 0.7-1.5 mm long, narrowly triangular, corolla rotate, white or white marked with lavender, blue, or purple, tube 0.6-1 mm long, lobes 0.5-1.2 mm long; stamens 4, sessile on the distal half of the tube, anthers ca. 0.4 mm long. Fruits 1.7-2.7 mm diam., body of the fruit 1.2-2 mm long and truncated at the apex, raphides prominent, calyx lobes ca. 0.5 mm long, borne on slender pedicels 4-1 1 mm long; seeds ca. 0.3 mm long, ellipsoid to depressed conic, reticulate.
Plants of open weedy sites on sandy soils in evergreen and in seasonally deciduous formations, from near sea level to 1 500 m. Probably flowering and fruiting throughout the year. The species has been found in both the Caribbean and Pacific low- lands and in the Meseta Central. The species ap- pears to have originated in Africa but is now wide- spread in the tropics.
Oldenlandia corymbosa is recognized by its short weedy habit, almost linear subsessile leaves, mi- nute flowers, and rounded capsules. The senior author believes that this is probably the correct name for all the Central American material for- merly called O. corymbosa, O. herbacea, and O. lancifolia. Separation of species on the basis of inflorescence differences or whether or not the stip- ules are bifid appears to be trivial.
Herbs to 15(-30) cm tall, prostrate or erect, with few to many branches, leafy stems 0.2-1.3 mm thick, gla- brous or with a few hairs at the nodes; stipules with a sheath 0.3-2 mm long, with 2-5 slender unequal awns to 1.5 mm long. Leaves sessile or with petioles to 1 mm long; leaf blades (7-)15-30M5) mm long, 0.5-3.5(-6) mm broad, linear to linear-oblong or very narrowly el-
Oldenlandia herbacea (L.) Roxb., Hort. Bengal. 11. 1814. Hedyotis herbacea L., Sp. PI. 102. 1753.
Annual or perennial herbs, 5-60 cm tall, erect to spreading and decumbent, stems glabrous, with 4 lon-
BURGER & TAYLOR: FLORA COSTAR1CENSIS. RUBIACEAE
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gitudinal ridges; stipules forming a very short (0.1-0.5 mm) sheath, truncate and with a few awns to 0.3 mm long. Leaves sessile; leaf blades 6-55 mm long, 1-4 mm wide, linear to linear-lanceolate, apex acute, base cu- neate, drying chartaceous, glabrous or with a few short hairs along the margins. Inflorescences axillary, flowers 1 or 2/node, pedicels 3-30 mm long, filiform, glabrous. Flowers usually isostylous, hypanthium 0.5-1 mm long, ovoid, glabrous to papillate or puberulent, calyx lobes 0.5-2.5 mm long, narrowly triangular, scabridulous on the margins; corolla white or lavender, or the tube green and the lobes with purple marks, corolla tube 2-1 1 mm long, cylindrical, lobes 1-3 mm long, ovate. Fruits 2.2- 5 mm long, 1.5-2 mm diam.. subglobose to ovoid, drying pale yellowish, glabrous to puberulent, crowned by the dark calyx lobes, beak 0.8-1 mm long; seeds 0.2-0.4 mm long, ovoid to ellipsoid, reticulate, brown.
Weeds of open sunny sites. Originally from Af- rica but now naturalized in Asia and parts of the Americas. The above description is based on Verdcourt (1976). This species has not yet been recorded from Costa Rica. Central American ma- terial earlier placed under this name is likely to be Oldenlandia corymbosa or O. lancifolia.
Oldenlandia lancifolia (Schumach.) DC., Prodr. 4: 425. 1830. Hedyotis lancifolia Schumach. in Schumach. & Thonn., Beskr. Guin. PI. 72. 1 827. Manettia bocaturensis Dwyer, Ann. Missouri Hot. Gard. 67: 278. 1980.
Perennial (rarely annual) herbs to 90 cm long, pros- trate or creeping, usually much-branched near the base and with simple distal stems, leafy stems glabrous or rarely minutely scabridulous; stipules forming a sheath to 1 mm long, with 2-5 slender awns 0.5-1.5 mm long. Leaves sessile or subsessile; leaf blades 1 0-60 mm long, 2-12 mm broad, linear or linear-lanceolate to narrowly elliptic, apex acute, base cuneate, drying chartaceous, dark green above, glabrous on both surfaces but often minutely scabrid along the revolute margins, 2° veins 3- 5/side. strongly ascending but thin and difficult to see. Inflorescences axillary, of solitary flowers or several flow- ers on very short peduncles (or reduced lateral shoots), pedicels 5-30 mm long, filiform, glabrous or scabridu- lous. Flowers monomorphic, ca. 3 mm long, hypanthium ca. 0.8 mm long and 1.5 mm diam., cupulate, glabrous or with scattered short hairs, calyx lobes 1-1 .8 mm long, triangular, glabrous or scabridulous; corolla white (some- times tinged with pink or purple), tube ca. 1 mm long, lobes 1-2 mm long; stigma lobes 0.7-1 .4 mm long. Fruits 2-3 mm long (including the 1 mm tall beak), 3.2-5 mm in diam., depressed subglobose; seeds 0.3-0.4 mm long.
Herbs of both seasonally deciduous formations and lowland rain forest areas. The species is wide- spread in tropical Africa and has been introduced to parts of the South America and the West Indies. The preceding description follows that of Verd-
court (1976). Central American material earlier ascribed to this species is probably O. corymbosa. Likewise, Steyermark's use of this name in the Flora de Venezuela (1974, pp. 408-41 1), may be incorrect.
Osa Aiello
REFERENCE— A. Aiello, A re-examination of Portlandia and associated taxa. J. Arnold Arbor. 60: 38-126. 1979.
Small trees, stems slightly expanded at the nodes; stip- ules united (interpetiolar), small, with an acute central lobe. Leaves opposite, petiolate, attenuate at the apex, drying thin-chartaceous and grayish green, entire, pin- nately veined, domatia absent. Inflorescence of single axillary flowers, borne on pedicels continuous with the base of the hypanthium/ovary. Flowers large, radially symmetrical, glabrous, calyx with 6 long narrow lobes; corolla with a long tube and distally funnelform, corolla lobes 5, broadly triangular; stamens 5, anthers linear; ovary 2-locular, placentas borne on the septum, with ca. 1 0 ovules in each locule, style filiform. Fruits thin-walled capsules, ellipsoid, with 6 longitudinal ribs, apparently opening septicidally, the calyx lobes persisting distally; seeds biseriate, slightly compressed, lacking wings, not imbricate, with persisting funicle.
A monotypic genus known only from the Osa Peninsula of Costa Rica. No other species of Costa Rican Rubiaceae has such large flowers. This ge- nus is related to Portlandia and to a lesser extent to Hintonia.
Osa pulchra (D. Simpson) Aiello, J. Arnold Arbor. 60: 116. 1979. Hintonia pulchra D. Simpson, Phytologia 29: 277. 1974. Figure 15.
A small tree or slender treelet, 2.5-15 m tall, leafy stem 1 .5-6 mm thick, glabrous, slightly expanded below the node; stipules 1-3 mm long, the broad base 1-2 mm high with a narrowed acute tip 0.5-1.5 mm long, per- sisting. Leaves with petioles 4-1 2 mm long, 0.7-1 .5 mm thick, glabrous, poorly differentiated from the lamina base; leaf blades 12-19 cm long, 3-6 cm broad, elliptic- oblong or narrowly oblong, apex gradually narrowed and acuminate, base acute to attenuate, drying thin-charta- ceous and grayish green, glabrous above and below, 2° veins 6-8/side, 3° veins obscure, domatia lacking. Inflo- rescences of solitary flowers in the axils of distal leaves, pedicels ca. 1 5 mm long but merging imperceptibly into the flower base, ca. 1 mm diam., glabrous, drying black. Flowers glabrous externally, probably slightly pendu- lous, apparently homostylous, hypanthium ca. 6 mm long, obconic, calyx lobes 18-40 mm long, 0.5-2.5 mm broad; corolla long-tubular and funnelform distally (trumpet-shaped), white, tube 1 7-27 cm long, 4-5 mm
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diam. and gradually flaring to 80 mm diam. at the mouth, lobes ca. 1 5 mm long and 30—40 mm broad at the base, broadly obtuse; anthers more than 20 mm long, 0.7 mm thick. Fruits ca. 3 cm long and 1.5 cm broad, oblong- ellipsoid, dark brown, the sepal lobes persisting but breaking off; seeds ca. 6 mm long, testa tuberculate.
This species is known from only a few collec- tions in lowland rain forest near Rincon de Osa at ca. 50 m elevation on the Osa Peninsula. Flow- ering in January-February, with immature fruits in June and mature fruits in January.
Osa pulchra is distinguished by its very large trumpet-shaped flowers that are thin in texture. No other Costa Rican member of the Rubiaceae has so long a flower. The long narrow basal tube suggests pollination by a long-tongued sphingid moth. The flowers are reminiscent of those of or- namental species ofBrugmansia (formerly includ- ed in Datura, Solanaceae).
Palicourea Aublet
REFERENCE— C. M. Taylor, Revision of Pali- courea in Mexico and Central America. Syst. Bot. Monogr. 26: 1-102. 1989.
Shrubs or small trees, glabrous or pubescent, stems terete, trigonous or quadrangular; stipules interpetiolar and often also intrapetiolar to form a sheathing tube with 2 triangular lobes or awns on each side (4/node) or some- times elongated interpetiolarly into emarginate or bi- lobed apices with the intrapetiolar sheath poorly devel- oped, colleters present within at the base. Leaves opposite and decussate or rarely 3-4/node, petiolate or rarely ses- sile; leaf blades entire and often elliptic, apex acute to acuminate, base rounded to acute, drying chartaceous to subcoriaceous, pinnately veined, domatia present or ab- sent. Inflorescences terminal, paniculate with opposite or alternate branching, variable in form (from elongate and racemose to thyrsoid or broadly corymbose) with flowers usually in distal cymose groups, peduncles and branches of the inflorescences often colored red, orange, yellow, purple, or blue (rarely green), glabrous or pu- bescent, bracts and bracteoles usually present (in Costa Rica), flowers usually pedicellate. Flowers bisexual and usually distylous, radially symmetric or slightly bilat-
erally symmetric when the corolla is curved or gibbous on 1 side, glabrous or pubescent externally, hypanthium usually turbinate, calyx tube with 5 calyx lobes (rarely truncated or spathaceous); corolla yellow, orange, red, purple, or blue (rarely white or cream), tubular to fun- nelform or salverform, membranaceous to camose. the corolla tube straight or curved, usually expanded (gib- bous) near the base, glabrous or puberulent externally, usually with a ring of hairs within the lower half of the tube (more rarely with the hairs in the upper half), corolla lobes 5, valvate in bud; stamens 5, inserted in the middle or on the throat of the tube, anthers dorsifixed and bifid at the base, included or exserted; ovary 2(-6)-Iocular, with 1 erect ovule from a basal placenta in each cell, stigmas 2-branched. Fruits fleshy, exocarp bluish to blu- ish black or purplish black; pyrenes usually 2. hemi- spheric with ca. 5 longitudinal ribs on the rounded back, usually with a longitudinal sulcus on the flattened inner face.
A Neotropical genus of about 200 species, rang- ing throughout moist tropical vegetation from Mexico and the West Indies to southern Brazil and Paraguay. Most species are South American; there are 3 1 in Costa Rica. As in Hoffmannia. this genus has speciated profusely at middle and higher (1000-2800 m) elevations. These plants are rec- ognized by their colorful terminal inflorescences, corolla tubes often slightly bent or expanded at the base, ovaries with a single erect ovule in each loc- ule, and the fleshy fruit usually with longitudinal ribs when dried. The infructescences usually be- come purple regardless of their color during an- thesis. The stipules forming a short tube above the node and with two distal awns or lobes, and the leaves with many secondary veins are additional characteristics distinguishing many Costa Rican species.
Palicourea is separated from the closely related genus Psychotria by the ring of hairs in the interior of the lower half of the corolla tube, a tendency for the corolla tube to be inflated or gibbous near the base, the more colorful inflorescences, and the generally larger more colorful corollas. Despite these differences, dried fruiting material may be very difficult to separate.
Key to the Species of Palicourea
la. Plants of lowland formations, rarely found from as high as 1000 m elevation 2a
Ib. Plants of montane forest formations, from (900-)1000 to 3200 m elevation 5a
2a. Flowers lavender or white marked with purple; stipule forming a sheath to 8 mm long [leaves
with 9-15 pairs of secondary veins; from wet Caribbean slopes at 500-1000 m elevation]
P. copensis
2b. Flowers red, orange, or yellow; stipular sheath 0-2 mm long; wide-ranging species 3a
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3a. Leaves 3/node, stipular teeth 6/node, 8-1 2 mm long and persisting, leaf blades to 8 cm broad
P. triphylla
3b. Leaves 2/node, stipular teeth or lobes 4, 2-9 mm long, often deciduous, leaf blades to 16 cm
broad 4a
4a. Stipular lobes rounded; leaf blades 1 5-28 cm long, to 1 6 cm broad P. guianensis
4b. Stipular teeth acute; leaf blades 7-14 cm long, to 7 cm broad P. crocea
5a. Plants pilose with hairs 1.5-2 mm long; stipules united and bifid or emarginate on each side;
flowers yellow and pilose P. standleyana
5b. Plants glabrous to puberulent, the hairs not exceeding 1 mm in length; stipules 4-lobed, bifid or
emarginate; flowers white, yellow, blue, or purplish 6a
6a. Calyx teeth or calyx lobes regularly more than 1.5 mm long [often persisting in fruit and with slightly smaller dimensions; flowers usually blue, purple, or white (red to yellowish only in P.
macrocalyx and P. orosiana)] 7a
6b. Calyx teeth or calyx lobes 0.2-1 .5 mm long 1 7a
7a. Calyx lobes more than 3.5 mm long 8a
7b. Calyx lobes less than 3.5 mm long 1 5a
8a. Calyx spathe-like and splitting down 1 side (to 20 mm long); corolla tube 3-4 cm long;
leaves with 1 2-22 pairs of 2° veins P. spathaceae
8b. Calyx 5-lobed (not spathe-like); corolla tubes 8-26 mm long; leaves with up to 19 pairs
of 2° veins 9a
9a. Corolla 20-26 mm long, ca. 4 mm diam. [calyx lobes 4-14 mm long, lanceolate; bracts of the inflorescence to 25 mm long and lanceolate; secondary veins 12-18 pairs; 1400-
2200 m elevation] P. hammelii
9b. Corolla tubes less than 20 mm long and 4 mm diam lOa
lOa. Young stems minutely puberulent 1 la
lOb. Young stems glabrous 1 2a
1 la. Calyx lobes 10-14 mm long, corolla white; stipule lobes 7-20 mm long; leaves
with 15-19 pairs of 2° veins; western Panama P. bella
1 Ib. Calyx lobes 6-1 1 mm long, corolla yellow; stipule lobes 6-17 mm long; leaves
with 8-12 pairs of 2° veins; central Costa Rica P. orosiana
12a. Leaves with 17-20 pairs of 2° veins; stipule lobes 2-9 mm long [corolla white]; floral
bracts 12-16 mm long; central Costa Rica P. bellula
12b. Leaves with 7-14 pairs of 2° veins; stipule lobes 2-10 mm long 12a
1 3a. Floral bracts 10-20 mm long, persisting and enclosing the flowers; corollas white;
2600-2800 m elevation in the Talamanca mountains P. skotackii
13b. Floral bracts 6-10 mm long, deciduous; corollas white with yellow or blue; Cor- dillera de Tilaran and central volcanic highlands, 1400-2200 m elevation . . 14a 14a. Corolla white and blue; stipule lobes 2-4 mm long; inflorescence to 22 cm long and
narrowly thyrsiform P . albocaerulea
14b. Corolla white and yellow; stipule lobes 4-10 mm long; inflorescences to 15 cm long
and paniculate P. macrocalyx
1 5a. Young stems usually puberulent [flowers white; inflorescences narrowly thyrsiform; leaves
2-7 cm broad; 800-1 600 m elevation] P. lancifera
1 5b. Young stems glabrous 1 6a
16a. Leaves l-2(-3) cm broad, blades to 8(-10) cm long; corolla blue and white (compare also
1 7a below) P. salicifolia
1 6b. Leaves becoming larger and broader; flowers variously colored 1 7a
17a. Flowers purple to violet, blue, or white (if reddish then reddish purple or lavender) 18a
1 7b. Flowers yellow to orange or red-orange 24a
1 8a. Leaf blades not more than 9 cm long [0.7-3.5 cm broad]; young stems often slightly puberulent
19a
1 8b. Leaf blades usually becoming more than 9 cm long; young stems glabrous or puberulent beneath the inflorescence . . 20a
FIELDIANA: BOTANY
19a. Inflorescences blue, corolla white to violet or blue; leaves thick, petioles thick; 1800-
3200 m elevation p. adusta
19b. Inflorescences yellowish green, corolla white to yellow; leaves thin, petioles slender;
1 300-2300 m elevation p, montivaga
20a. Leaf blades l-3(-4) cm broad, to ca. 14 cm long 2 la
20b. Leaf blades usually becoming more than 4 cm broad, to 24 cm long 22a
2 la. Inflorescences thyrsiform and purple, to 16 cm long; calyx lobes ca. 0.4 mm long . . .
P. angustifolia
21b. Inflorescences paniculate and purple, to 8 cm long; calyx lobes ca. 1 m long
P. purpurea
22a. Corolla white, tube 3-6 mm long; calyx lobes ca. 0.3 mm long; leaves with 9-1 1 pairs of 2°
veins P. tilaranensis
22b. Corolla blue to purple, tube 5-10 mm long; calyx lobes 1-3 mm long; leaves with 11-21
pairs of 2° veins 23a
23a. Inflorescence purple, paniculate; leaf blades to 26 x 13 cm, with 11-19 pairs of 2° veins
P. discolor
23b. Inflorescence blue, thyrsiform; leaf blades to 18 x 8 cm, with 15-21 pairs of 2° veins ....
P. brenesii
24a. Stems and veins on the lower leaf surfaces densely puberulent with short stiff scurfy or slender
hairs; flowers puberulent P. vestita
24b. Stems glabrous to sparsely puberulent; flowers glabrous or puberulent 25a
25a. Stipule lobes/teeth 0.5-2 mm long; leaf blades 3-12 cm long and 1-3.5 cm broad 26a
25b. Stipule lobes/teeth 2-8 mm long; leaf blades usually more than 7 cm long and 3 cm broad . . 27a 26a. Flowers yellow or white with yellow and purple; leaves with 12-18 pairs of 2° veins; 1200-
1 600 m elevation P. garciae
26b. Flowers white and yellow; leaves with 6-9 pairs of 2° veins; wide ranging in Costa Rica;
1 1 00-2000 m elevation P. montivaga
27a. Inflorescence branches red to orange (to purple in fruit); corolla tube 6-10 mm long, orange;
(800-) 1200- 1700 m elevation P. padifolia
27b. Inflorescence branches yellow to greenish yellow; corolla tube 7-1 3 mm long, yellow; (800-)1000-
2300 m elevation 28a
28a. Corolla tube 7-1 1 mm long; stems glabrous; leaves with 8-13 pairs of 2° veins .... P. padifolia 28b. Corolla tube 1 1-13 mm long; stems puberulent or glabrous; leaves with 1 1-19 pairs of 2° veins
. P. lasiorhachis
Palicourea adusta Standl., J. Wash. Acad. Sci. 18: 279. 1928. Figure 53.
Small shrubs or rarely little treelets, 0.5-3(-6) m tall, leafy branchlets 0.7-6 mm thick, glabrous or with minute (0.1-0.5 mm) appressed hairs, becoming terete; stipules 2-6 mm long, united above the petiole and forming a very short (1-3 mm) tubular sheath, with 2 distal teeth 1-3 mm long. Leaves opposite, petioles 4-12(-22) mm long, 0.5-1 mm thick, glabrous (rarely minutely puber- ulent); leaf blades 3-7(-10) cm long. 1-3.5 cm broad, narrowly to broadly elliptic, elliptic-oblong, apex acu- minate with tip 5-10(-l 5) mm long, base broadly obtuse to acute and slightly decurrent on the petiole, drying thin- chartaceous to stiffly chartaceous, brown to greenish, glabrous above and usually with linear cystoliths visible, glabrous or with a few minute appressed hairs on the veins beneath, 2° veins 6-1 I/side and usually joining a vein at the edge of the leaf, intersecondaries present. Inflorescences 6-12 cm long, 4-9 cm broad, open py-
ramidal with opposite branching, yellowish to blue-gray, peduncles 1.5-5 cm long, 0.7-1.4 mm thick, puberulent with ascending hairs 0.2-0.4 mm long, bracts 1-4 mm long, linear-lanceolate, bracteoles 1-2.5 mm long, ped- icels 1-5 mm long. Flowers usually glabrous externally, hypanthium 1-1.5 mm long, turbinate, glabrous or sparsely puberulent (hairs to 1 mm long), calyx lobes 0.5-0.8(-1.5) mm long. ca. 1 mm broad at the base; corolla tubular, bluish purple, violet, or white marked with purple or violet, usually glabrous, tube 6-10.5 mm long. 1-2 mm diam. and slightly expanded at the base, corolla lobes 5, 1.5-3 mm long, minutely whitish pa- pillate along the inner margin; anthers 1.5-2 mm long. Fruits 3-4.5(-6) mm long and 4 mm diam., obovoid, becoming dark purple to deep blue and translucent, with longitudinal ridges when dry, persisting calyx ca. 0.7 mm high.
Plants of ridges and wet open areas in evergreen montane rain forest formations, ( 1 800-)2200-3 1 00
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m elevation. Probably flowering and fruiting throughout the year. These plants appear to prefer ridges and wet open areas. The species ranges from the Cordillera de Tilaran to western Panama.
Palicourea adusta is recognized by its high-al- titude habitat, small leaves, small calyx lobes, usu- ally bluish flowers (less often yellowish or purple). This species is similar to P. montivaga with white flowers and P. salicifolia with yellowish inflores- cences.
Palicourea albocaerulea C. M. Taylor, Syst. Bot. Monogr. 26: 15. 1989. Figure 53.
Shrubs or small trees, 2-7 m tall, leafy stems 1.2-4 mm thick, glabrous, quadrangular or rounded; stipules to 8 mm long, united at the base to form a sheath 1-4 mm long, apex truncated with 2 teeth 2-4 mm long. Leaves opposite, petioles 8-22 mm long, 0.6-1.2 mm thick, glabrous, often drying reddish brown; leaf blades 5-15 cm long, 2-5 cm broad, narrowly elliptic to nar- rowly oblong or elliptic-oblong, apex acuminate with tip 6-15 mm long, base acute (obtuse) to attenuate and slightly decurrent on petiole, drying chartaceous, gla- brous above, glabrous beneath or with a few short (0. 1- 0.3 mm) hairs along the midvein beneath, linear cys- toliths sometimes visible on the lower surface, 2° veins 7-1 I/side and merging with a marginal vein at the leaf edge. Inflorescences 1 1-22 cm long, 3-8 cm broad, nar- rowly pyramidal (thyrsoid) with opposite or alternate lateral branches, peduncles 1.5-5 cm long, 1.6-2.2 mm thick, glabrous and drying dark brown, bracts 6-10 mm long, linear and acute, pedicels 3.5-10 mm long, brac- teoles 2-6(-8) mm long, glabrous. Flowers glabrous ex- ternally, hypanthium ca. 1 .5 mm long, ca. 0.5 mm diam. and funnelform, calyx lobes 5, 3.5-9 mm long, 1-1.2 mm broad, narrowly oblong, often unequal; corolla tu- bular, white or white with blue, tube 8-1 1 mm long, 2- 3 mm diam., slightly expanded near the base, corolla lobes 1-2 mm long and 1 .5 mm broad at the base; anthers ca. 2 mm long. Fruits 5-8 mm long (not including the calyx lobes) and 5-10 mm diam., ovoid or globose, blu- ish black, persisting calyx lobes ca. 7 mm long.
Plants of lower montane cloud forest forma- tions, from 1450 to 1650 m elevation. Flowering in September-December; fruiting in June. This species is only known from the Cordillera de Ti- laran and adjacent areas in Costa Rica.
Palicourea albocaerulea is recognized by its re- stricted range, generally glabrous parts, long inflo- rescences, long calyx lobes, and white corollas that turn blue with age. The narrow leaves and long slender petioles are also distinctive. This is our only species in which the flowers change from white to blue. Compare this species with P. lancifera with shorter calyx lobes and flowers that do not turn blue.
Palicourea angustifolia H.B.K., Nov. gen. sp. 3: 367. 1819. P. lanceolata Oersted ex Polakows- ky., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 17. 1852.
Shrubs or small trees, 2-5(-10) m tall, leafy branchlets 1.3-4 mm thick, glabrous or rarely sparsely puberulent, terete; stipules to 10 mm long, the tubular sheath 2-4 mm long and truncate but with slender awns 3-6(-8) mm long, glabrous, persisting. Leaves opposite, petioles 2-9(-13) mm long, 0.6-1.2 mm thick, glabrous; leaf blades 5-15(-23) cm long, l-4(-5) cm broad, lanceolate to narrowly elliptic or very narrowly oblong, apex grad- ually narrowed and acute or acuminate, tip 4-14(-25) mm long, base acute and slightly decurrent on petiole, drying chartaceous, glabrous above, glabrous beneath or with short (0.4 mm) thin whitish hairs along the veins, 2° veins 10-14/side. Inflorescences 3-14(-16) cm long, 3-6(-8) cm broad, a narrowly pyramidal (thyrsoid) pan- icle, peduncles 1.5-3(-6) cm long, 1-2 mm, thick, gla- brous or minutely appressed-puberulent, lateral branch- es opposite, subopposite or alternate, purple to magenta, bracts 2-5 mm long, pedicels 0-3 mm long, bracteoles ca. 1 mm long. Flowers glabrous or minutely (0. 1 mm) papillate puberulent externally, hypanthium ca. 1 mm long, calyx lobes ca. 0.4 mm long, triangular; corolla slender funnelform, rose red to purple or fuchsia, tube 9-14 mm long, 1.5-2.5 mm diam., expanded near the base, corolla lobes 5, 1.2-2.5 mm long; anthers 1.7-2.5 mm long, included. Fruits 4-6 mm long and 5 mm diam., globose to ovoid, becoming purple and drying black, puberulent or glabrate, persisting calyx ca. 0.5 mm long.
Plants of evergreen montane forest formations on both the Caribbean and Pacific slopes, from (1400-)! 600 to 2000(-2300) m elevation. Flow- ering and fruiting throughout the year but with most flowering collections made in January-Sep- tember. The species ranges from central Costa Rica to Peru.
Palicourea angustifolia is recognized by the nar- row leaves, truncated stipule sheath with long awns, narrow inflorescences, and slender funnelform purple corollas with relatively short lobes. This species resembles P. padifolia and P. purpurea but differs in the relatively narrower inflorescences, shorter pedicels, and thinner, more funnelform co- rollas that are usually minutely puberulent at the base. In addition, P. padifolia has yellow corollas and red or orange inflorescence branches.
Palicourea bella (Standl.) Dwyer, Ann. Missouri Bot. Gard. 67: 299. 1980. Psychotria bella Standl., J. Wash. Acad. Sci. 18: 185. 1928.
Shrubs or small trees to 5 m tall, leafy stems ca. 4 mm thick and quadrangular, glabrous or with small (0. 1-0.3 mm) hairs; stipules 10-20 mm long, to 15 mm broad.
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with thin leaf-like texture, united above the petioles, oblong, rounded and bilobed to emarginate. Leaves clus- tered distally, petioles 8-20 mm long, 1.2-2 mm thick, puberulent; leaf blades 9-22 cm long, 3-8.5 cm broad, elliptic to elliptic-obovate or oblong-elliptic, apex acu- minate, base gradually narrowed and acute, drying char- taceous, dark greenish brown, glabrous or with short (0.3 mm) hairs on the veins and smaller hairs between the veins on both surfaces, 2° veins 12-19/side. Inflores- cences solitary, 8-20 cm long, 7-10 cm broad, broadly corymbiform, peduncles 5-13 cm long, to 2 mm thick and sparsely strigulose, bracts 10-26 mm long, ovate- oblong, translucent brown and sparsely hirtellous, sec- ondary peduncles 1 5-25 mm long, bright pink to purple, pedicels 2-8 mm long, obscured by the large bracts and bracteoles to 8 mm long. Flowers with hypanthium ca. 2 mm long, glabrous or with conspicuous thin yellowish hairs ca. 0.7 mm long, the calyx lobes 6-14 mm long, 3-6 mm broad, ovate to ovate-elliptic, drying thin trans- lucent with scattered short (0.2 mm) hairs; corolla tu- bular, white to pink, carnose, tube 14-20 mm long and 2-3 mm diam., dilated at the base, corolla lobes 4-5 mm long, ovate; anthers ca. 3.5 mm long. Fruits ca. 6 mm long and 6 mm diam., ellipsoid, glabrous or sparsely puberulent.
Plants of evergreen montane forest formations at (1500-)2000-2700 m elevation. Flowering in April. This species is only known from Chiriqui Province in Panama and a collection from the Caribbean slope of central Costa Rica at 1500 m.
Palicourea bella is recognized by the broadly two-lobed stipules, corymbiform inflorescences with broad thin bracts, and the large thin calyx lobes. In life, both inflorescences and calyx are red- violet to bright pink in color. The similarity in morphology, texture, and vestiture of bracts and calyx lobes is an interesting example of heterotopy. This species is similar to P. bellula (q.v.) as well as to P. hammelii C. M. Taylor and P. ochnoides Dwyer of western Panama. Grayum 7046 (CR, MO) from Volcan Barva is tentatively placed here.
Palicourea bellula C. M. Taylor, Syst. Bot. Mono- gr. 26: 24. 1989. Figure 49.
Shrubs or slender treelets, to 2.5 m tall, leafy stems 2.5-5 mm thick, glabrous, quadrangular and drying dark, grayish in age; stipules 5-1 1 mm long, 7-10 mm broad, united above the petioles for 2-3 mm to form a broad tube, with 2 rounded lobes 2-4 mm high, thin in texture and glabrous. Leaves opposite, petioles 4-18 mm long, 1.5-2.5 mm thick, glabrous, sulcate adaxially near the base; leaf blades 7.5-14 cm long, 3.5-7 cm broad, broad- ly elliptic to slightly elliptic-obovate, apex acuminate, tip ca. 5- 1 0(- 1 5) mm long, base obtuse (acute) and slight- ly decurrent on petiole, drying stiffly chartaceous, green- ish, glabrous above, with thin whitish hairs 0.3-1 mm long along the sides of the midvein and on some 2° veins beneath, 2° veins ( 1 0-) 1 7-20/side. Inflorescences 1 or 3,
9-14 cm long, 9-11 cm broad, rounded panicles, red- violet to fuchsia, peduncles 3-8 cm long, 1-3 mm thick, glabrous, bracts 12-16 mm long, 3-4 mm broad, lan- ceolate, bracteoles 5-12 mm long and 3-5 mm broad, narrowly ovate, pedicels 4-12 mm long. Flowers gla- brous externally, hypanthium ca. 1 mm long and 1.2 mm diam.. drying dark and glabrous, calyx tube 2-3 mm long and broadly cupulate, calyx lobes 5. 4-7 mm long, 2-3 mm broad, ovate, red-violet to fuchsia; corolla white, tubular, carnose, tube 12-15 mm long and 2.5 mm diam.. corolla lobes 2-3 mm long; anthers ca. 3 mm long. Fruits not seen at maturity.
Plants of open sites in montane rain forest for- mations near the continental divide at 1 900-2200 m elevation. Flowering in January-February, May, and November. This species is only known from between Volcan Viejo (Alajuela) and the south- eastern slope of Volcan Barva near the upper part of Rio Patria (Heredia Province) Costa Rica.
Palicourea bellula is distinguished by its colorful glabrous inflorescences with conspicuous pink to purple bracts and calyx lobes, tubular white co- rollas, leaves with many closely parallel secondary veins, broadly rounded stipule lobes, and restrict- ed cloud forest habitat. This species is similar to P. bella, but the calyx and corolla are usually small- er and stiffer than in P. bella. Palicourea hammelii and P. ochnoides are similar but lack the broad imbricate calyx lobes.
Palicourea brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1333. 1938. P. talamancana Standl. & L. O. Williams, Ceiba 1: 250. 1951. Figure 52.
Shrubs, 1-4 m tall, erect or clambering, often with only a few distal branches 2.5-5 mm thick, glabrous or sparsely strigillose with hairs 0.5-1 mm long, drying dark, quadrangular; stipules united to form a tubular sheath 3-6 mm long, truncated distally and with 2 prominent teeth (on each side) 4-7(-10) mm long and 1-2 mm broad at the base, usually glabrous, persisting. Leaves opposite, petioles 1.5-3 cm long, 1.2-2.2 mm thick, gla- brous; leaf blades 9-1 8 cm long, 3.5-8 cm broad, broadly elliptic to elliptic-obovate or elliptic-oblong, apex abruptly narrowed and short-acuminate, base obtuse to acute, drying stiffly chartaceous, brownish or yellowish, gla- brous above, glabrous or sparsely puberulent beneath, 2° veins 15-21 /side. Inflorescences 8-15 cm long, 4-9 cm broad (at broadest part), pyramidal or elongate thyr- soid panicles, often blue throughout, peduncles 2-5 cm long, 1.5-3 mm thick, sparsely to densely puberulent with thin hairs ca. 0.2 mm long, bracts 5-8 mm long, ca. 2 mm broad, lanceolate, pedicels 0-6(-8) mm long, bracteoles 1-5 mm long. Flowers minutely puberulent and glabrescent distally, hypanthium ca. 1 mm long and 0.8 mm diam., calyx tube broadly cupulate, calyx lobes 1-2.5 mm long, broadly triangular and obtuse; corolla
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blue, tubular, glabrous near the base and densely ap- pressed-puberulent distally or glabrous throughout, tube 8-1 1 mm long, ca. 2 mm diam.. narrowed near the mid- dle, corolla lobes 5, 3-5 mm long; anthers ca. 2.5 mm long. Fruits 4-6 mm long, 4-6 mm diam., globose, gla- brate.
Plants of evergreen montane rain forest for- mations, from 1800 to 2600(-3200) m elevation. Flowering and fruiting in February-June. The spe- cies has only been collected in the northern part of the Meseta Central (near Zarcero and Palmira) and along the Interamerican Highway in the west- ern part of the Cordillera de Talamanca in Costa Rica.
Palicourea brenesii is distinguished by the larger leaves with many secondary veins, bright "china blue" thyrsoid panicels, and small blue to violet flowers with conspicuous calyx lobes. The usually glabrous stems contrast with the puberulent pe- duncle and inflorescence branches. The striking bright blue color of the inflorescence is similar to that found in species of Faramea. The disjunct collections from the Cordillera de Talamanca have smaller corollas and stipule lobes to 10 mm long. This species is very similar to P. discolor after the flowers have faded.
Palicourea copensis (Dwyer) C. M. Taylor, comb, nov. Psychotria copensis Dwyer, Ann. Missouri Bot. Gard. 67: 365. 1980. Figure 52.
Shrubs or small treelets, 2-5 m tall, leafy stems 2-5 mm thick, with curved yellowish or whitish hairs 0.4- 0.9 mm long, glabrescent; stipules 6-12 mm long, united to form a broad tube to 10 mm long, 4-8 mm broad (to 14 mm broad below the inflorescences), bilobed with a distal sinus 1-6 mm deep, glabrous but with a ciliolate margin, persisting. Leaves with petioles 10-55 mm long, 1.3-2.3 mm thick, glabrous or sparsely puberulent be- neath; leaf blades 9-23 cm long, 3.5-9 cm broad, elliptic- obovate to elliptic-oblong, apex narrowly acuminate and 5-13 mm long, base cuneate and slightly decurrent on petiole, drying chartaceous, dark green above (pale green beneath), glabrous above or with a few hairs along the major veins, sparsely pubescent beneath with crooked whitish hairs 0.4-1.5 mm long, 2° veins 9-14/side and arcuate-ascending distally. Inflorescences terminal or pseudoaxillary, 1 or several per node, 2-7 (-1 1) cm long, 3-6(-l 2) cm broad, pyramidal with few opposite branch- es, peduncles 8-26 mm long, ca. 1 .5 mm thick, pubescent or glabrous, bracts 3-7 mm long, flowers in distal cymose or irregular clusters. Flowers with hypanthium ca. 1 mm long, obconic, glabrous or pubescent, calyx yellowish, calyx lobes quite variable, 1-3 mm long, oblong and obtuse; corollas lavender or white marked with purple, salverform with a distinctly gibbous base, tube 7-12 mm long, 2-4 mm diam., puberulent, corolla lobes 3-5 mm
long; anthers ca. 3 mm long. Fruits 6-7 mm long, 4-5 mm diam., obovoid, becoming blue or violet.
Plants of the very wet Caribbean escarpment, between 400 and 1 000 m elevation. Flowering and fruiting in March-May and November. The spe- cies is known from the Caribbean slopes of Volcan Barva and the P. N. Braulio Carrillo area in central Costa Rica. It was originally described from El Cope in Code Province, Panama.
Palicourea copensis is recognized by its white or lavender flowers, variable sepal lobes, smaller thick-branched inflorescences, large persisting stipules, puberulence of usually curved hairs, and lower montane habitat. This is a rarely collected species that appears to prefer creek margins.
Palicourea crocea (Sw.) Roem. & Schult., Syst. Veg. 5: 193. 1819. Psychotria crocea Sw., Prodr. 44. 1788. Figure 50.
Shrubs or small trees, 1-5 m tall, leafy stems 1-4 mm thick, glabrous or minutely puberulent, terete; stipules united to form a short (0.5-2 mm) truncated tube with narrow distal teeth 1-4 mm long, glabrous, deciduous. Leaves opposite, petioles 6-12(-20) mm long, 1-1.5 mm thick, usually glabrous; leaf blades 7-14(-19) cm long, 2.5-6(-7) cm broad, elliptic, narrowly elliptic, lanceolate, to elliptic-ovate or elliptic-oblong, apex acute to acu- minate, base obtuse to acute and slightly decurrent on the petiole, drying membranaceous to thin-chartaceous, greenish, usually glabrous above, with short (0.3 mm) thin hairs along the sides of the midvein beneath, 2° veins 8-12/side. Inflorescences 6-15 cm long, 2.5-7 cm broad, nararow pyramidal to rounded with opposite or subopposite (alternate distally) branching, reddish to or- ange or pink, peduncles 1.5-1 1 cm long, 1-2 mm thick, glabrous or sparsely and minutely puberulent, bracts 1- 10 mm long, linear-lanceolate and deciduous, pedicels 4-8(-12) mm long, slender, bracteoles ca. 1 mm long and deciduous. Flowers distylous, glabrous or minutely puberulent externally, hypanthium ca. 1 mm long and 0.7 mm diam., calyx lobes 0.3-0.5 mm long, broadly ovate; corolla tubular, dark red to yellow-orange or pink (rarely bluish green), tube 5-9 mm long, 1 .5-3 mm diam., inflated near the base, corolla lobes 1-2.5 mm long; an- thers ca. 2.7 mm long. Fruits 4-6 mm long and 4 mm diam., subglobose or ovoid, becoming dark blue or black, longitudinally ribbed when dry, persisting calyx minute.
Plants of secondary growth in evergreen rain forest areas often along streams and paths in the Caribbean lowlands, from near sea level to 600 m elevation. Flowering and fruiting throughout the year (flowering mostly in April-August in Central America). The species ranges from central Mexico and the West Indies to Paraguay.
Palicourea crocea is recognized by its smaller
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stature and lowland habitat, short stipular tube, colorful thyrsoid inflorescences, minute calyx lobes, and orange or red corolla tubes. This species is often found in swampy areas and other sites with poor drainage. This species is sometimes confused with P. padifolia, of higher elevations and with longer stipule sheaths, with P. guianensis, with longer stipule lobes, and with species ofHamelia.
Palicourea discolor K. Krause, Bot. Jahrb. Syst. 54: Beibl. 119: 40. 1916. P. macrosepala K. Krause, loc. cit. 41. 1916. P. panamaensis Standl., Ann. Missouri Bot. Gard. 25: 839. 1938. Figure 52.
Shrubs or small trees, 1-4(1-6) m tall, leafy stems 3- 6 mm thick, glabrous and quadrangular; stipules united to form a broadly tubular sheath 2-6 mm long, truncated apically and with 2 narrow teeth 4-7 mm long, glabrous and persisting. Leaves opposite, petioles l-3(-5) cm long, 1.7-2.7 mm thick, glabrous; leaf blades 10-26 cm long, 5-13 cm broad, elliptic to broadly elliptic-oblong, acute to short-acuminate, the tip 5-10(-15) mm long, obtuse or acute at the base, drying stiffly chartaceous, dark brown or greenish above, glabrous above, sparsely puberulent with inconspicuous (0.05-0.2 mm) thin whitish hairs on the veins beneath, 2° veins 11-19/side, united distally with a vein at the leaf edge. Inflorescences solitary or the basal branches subtended by leaves and apparently 3, 12-22(-30) cm long, 10-1 5(-24) cm broad, pyramidal with opposite or subopposite branches, bright purple in life, peduncles 2-14 cm long, 2.5-4 mm thick, glabrous, bracts 5-8 mm long, 1 mm wide, linear-lanceolate, ped- icels 1-8 long, glabrous, bracteoles absent. Flowers gla- brous externally, hypanthium 0.5-1 .3 mm long and 0.3- 1 mm diam.. calyx lobes 1.5-5 mm long, acute to Un- gulate, very variable in some collections; corolla purple or white flushed with purple, carnose, tube 7-13 mm long, 1-2 mm diam., corolla lobes 1-3.5 mm long. Fruits 4-5 mm long and 4-5 mm diam., globose to globose- ellipsoid, with longitudinal ribs (dried), calyx persisting or deciduous.
Plants of forest interiors and shaded thickets in evergreen montane forest formations, from 1 1 00 to 1900(-2600) m elevation. Flowering and fruit- ing in February-August. The species ranges from the upper Rio Grande de Orosi and Moravia de Chirripo in the Cordillera de Talamanca of Costa Rica southward into the Chiriqui Highlands of western Panama.
Palicourea discolor is recognized by its relatively large leaves and inflorescences, purple flowers and inflorescence branches, and variable calyx lobes to 5 mm long. This species is similar to P. brenesii with bright blue inflorescences and smaller corol- las and P. purpurea with smaller calyx lobes and smaller leaves.
Palicourea garciae Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 195. 1940, non P. garciae Steryerm. 1971. Psychotria copeyana Standl. & L. O. Williams, Ceiba 1: 251. 1940. Figure 53.
Shrubs or small trees, 1.5-5 m tall, leafy branchlets 1.5-3.5 mm thick, glabrous or with short (0.3 mm) as- cending yellowish hairs, quickly becoming glabrescent and terete, drying dark; stipules united to form a tubular sheath 2-4 mm long, each side with 2 short ( 1 mm) distal lobes and a small ( 1 mm) central sinus, glabrous or mi- nutely appressed-puberulent. Leaves opposite, petioles 5-15 mm long, 0.6-1.2 mm thick, glabrous; leaf blades 5-12(-15) cm long, 2-4(-4.5) cm broad, narrowly ellip- tic, narrowly elliptic-ovate to narrowly oblong or lan- ceolate, apex acuminate with tip 7-15 mm long, base acute to obtuse and slightly decurrent on petiole, drying stiffly chartaceous, much darker above than beneath, glabrous above or with hairs along the midvein, with few or many minute (0.05-0.2 mm) hairs along the major veins beneath and usually glabrescent, 2° veins 12-18/ side, arising from the midvein at near 90°. Inflorescences solitary, 4-12 cm long, 6-12 cm broad, open pyramidal or rounded corymbiform with opposite (subopposite) branching, green or flushed with purple, peduncles (l-)2.5-4 cm long, very sparsely to densely puberulent with short (0.2 mm) thin hairs, bracts 2-5 mm long, linear-subulate, pedicels 1-5 mm long, bracteoles ca. 1 mm long. Flowers glabrous externally, hypanthium 1- 1.5 mm long, 0.8-1 mm diam., calyx lobes 5, 0.7-1.5 mm long, acute; corolla funnelform and gibbous near the base, white or flushed with yellow or purple distally (grayish to purple near the base), tube 6-8 mm long, constricted in the middle (1-1..5 mm diam.) and gibbous near the base, corolla lobes 2-3 mm long and 1 mm broad at the base; anthers 2-2.5 mm long. Fruits 4-5 mm long, 3-4 mm diam., globose to ellipsoid, becoming pale blue at maturity, with longitudinal ribs, persisting calyx ca. 1 mm long.
Plants of montane rain forest formations, from 1200 to 1600(-2000) m elevation. Flowering in March and May-August; fruiting in May-June and August. The species ranges from central Costa Rica to Colombia.
Palicourea garciae is recognized by the small narrow leaves with many secondary veins, the small corolla tubes strongly bent below the middle and inflated at the base, and the small pale blue fruit. The very short tubular sheath and bifid interpetio- lar part of the stipule is noteworthy. Psychotria copeyana was based on a specimen ( Williams <&. Allen 16482 F) with immature inflorescences from El Copey. Vegetatively this species resembles sev- eral smaller-leaved species of Psychotria, but the strongly gibbous flowers are very different.
Palicourea guianensis Aubl., Hist. pi. Guiane 1: 173, t. 66. 1775. Psychotria palicourea Sw., Fl. Ind. Occ. 1797. Figure 50.
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Shrubs or small trees, 2-6(-10) m tall, leafy stems 1.5- 8 mm thick, glabrous or minutely puberulent, quadran- gular, often drying dark and contracted beneath the node; stipules free or united for 1-2 mm at the base, lobes, 5- 8(-12) mm long, lobes 2-4 mm broad, blunt, glabrous and drying dark. Leaves opposite, petioles 8-24(-38) mm long, 1.3-3 mm thick, glabrous; leaf blades (12-) 15-28 cm long, (5-) 7-1 8 cm broad, broadly ovate-elliptic to ovate-oblong, elliptic-oblong or ovate, apex acute or acu- minate, base obtuse to rounded and subtruncate, drying thin-chartaceous, concolorous, glabrous or minutely (0. 1 mm) papillate-puberulent above and below, 2° veins 10- 15/side. Inflorescences usually solitary, 7-18 cm long, 4-12 cm broad, pyramidal or thyrsiform with many closely crowded alternate or subopposite branches, red- orange when flowering and later turning purple, pedun- cles 3-12 cm long, 1.8-3.2 mm thick, minutely puber- ulent with thin hairs ca. 0. 1 mm long, bracts absent or 0.5-7 mm long and often adnate to the primary branches, pedicels 2-5(-8) mm long, minutely puberulent. Flowers monomorphic, minutely puberulent externally, hypan- thium ca. 1.5 mm long and 0.8 mm diam., tubular, calyx lobes 0.2-0.6 mm long, broadly obtuse; corolla yellow or orange, tube 8-25 mm long and 2-3 mm diam. at base, contracted in the middle and a slightly inflated distally, corolla lobes 1-2 mm long, triangular; ovary 2- 6-locular. Fruits 4-7 mm long, 4-5 mm diam., ovoid to ellipsoid, becoming purple violet and drying black with prominent longitudinal ribs, persisting calyx and disc ca. 1 mm high.
mm thick, glabrous or minutely puberulent, drying dark; leaf blades 8.5-20 cm long, 2.5-8.5 cm broad, elliptic to elliptic-obovate, apex acuminate with tip 5-15 mm long, base gradually narrowed and acute or cuneate, slightly decurrent on petiole, drying stiffly chartaceous, dark olive green above, glabrous above or puberulent on the midvein, puberulent beneath with crooked hairs 0.05- 0.3 mm long, 2° veins 12-18/side. Inflorescences 4-14 cm long, to 1 3 cm broad, open pyramidal, pink to purple but drying dark, peduncle 15-35 mm long, 1.5-3 mm thick, glabrous or very sparsely minutely puberulent, proximal bracts 18-35 mm long, ca. 5 mm broad, lan- ceolate, distal bracteoles 6-20 mm long, pedicels 4-15 mm long. Flowers glabrous externally, hypanthium ca. 2 mm long and 1 mm diam., calyx 1 3-20 mm long, pink to purple, calyx lobes 4-1 2 mm long, lanceolate, unequal; corolla white, tubular, tube 20-27 mm long, ca. 4 mm diam., corolla lobes 3-6 mm long, triangular. Fruits 8- 10 mm long, 7-8 diam., ellipsoid.
Palicourea hammelii was originally described from collections made between 1 400 and 2200 m elevation in Chiriqui, Panama. It has been recently collected on the Fila Matama at 1 600 m elevation on the Caribbean slope of the Cordillera de Tala- manca (Herrera & Chacon 2781 CR, MO). This spe- cies resembles P. bella with ovate calyx lobes and P. bellula with a shorter corolla tube.
Plants of open secondary formations in ever- green and partly deciduous forest areas, from near sea level to 900 m elevation. Flowering and fruit- ing occur throughout the year; flowering mostly in March-July in Costa Rica. The species ranges from southern Mexico and the West Indies southward to southern Brazil and Bolivia.
Palicourea guianensis is recognized by its low- land habitat, the wide glabrous or very minutely puberulent thin-textured leaves, the large rounded stipule lobes, the conspicuous densely flowered yellow or orange inflorescences, and yellow or or- ange puberulent corollas. Most of our collections come from the Caribbean lowlands and from the Golfo Dulce area. Central American material has two-locular ovaries, but specimens from eastern South American may have three-or four-locular ovaries. Palicourea guianensis resembles P. padi- folia (q.v.).
Palicourea hammelii Taylor, Syst. Bot. Monogr. 26:43. 1989.
Small trees to 5 m tall, leafy stems 3-7 mm thick, glabrous or minutely (0. 1 mm) appressed-puberulent, drying dark; stipules 6-12 mm long, with a sheathing base 3-5 mm long with 2 lobes (4/node) 3-7 mm long, persisting. Leaves with petioles 13-35 mm long, 1.3-2
Palicourea lancifera Standl. & L. O. Williams, Cei- ba 1: 249. 1951. Figure 53.
Shrubs, 1.5-3(-5) m tall, leafy stems 1.3-4 mm thick, glabrous to (less often) densely puberulent with crooked yellowish hairs 0.1-0.3 mm long, quadrangular at first, glabrescent; stipules united to form a tubular sheath 3- 5 mm long, apex truncated with 2 narrow teeth 2-5 mm long on each side. Leaves opposite, petioles 7-25(-30) mm long, 0.4-1.2 mm thick, glabrous or minutely pu- berulent; leaf blades 5-1 5(-l 8) cm long, 2-5(-7) cm broad, elliptic to narrowly elliptic-oblong, apex acuminate with tip 6-20 mm long, base acute and decurrent on petiole, drying thin-chartaceous or chartaceous, dark greenish or brownish above, glabrous above, minutely (0. 1-0.4 mm) strigulose on the veins beneath, 2° veins 8-12/side and connecting with a vein along the leaf edge. Inflorescences solitary, 7-17(-21)cm long, 2-6(-9)cm broad, narrowly pyramidal with many short branches, green or marked with yellow, peduncles 1-4 cm long, 1-2 mm thick, gla- brous to densely puberulent, bracts 5-8 mm long, nar- rowly lanceolate, flowers often alternate along the lateral branches, bracteoles 1.5-4 mm long, pedicels 2-7(-17) mm long. Flowers glabrous or sparsely puberulent ex- ternally, hypanthium 1-1.5 mm long, ca. 1 mm diam., calyx lobes 1.5-3 mm long, often unequal, acute or rounded distally; corolla white or flushed with green or yellow, tubular, tube 7-1 1 mm long, ca. 2 mm diam. and expanded at the base, corolla lobes 1-2 mm long; anthers 2-2.5 mm long. Fruits 4-5 mm long and 4-5 mm diam., ovoid or globose-oblate, persisting calyx ca. 1.5 mm high.
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Plants of very wet lower montane cloud forest formations, from 800 to 1 700 m elevation. Flow- ering June-November; apparently fruiting throughout the year. This species ranges from the Reserva Forestal de San Ramon and Zapote de San Carlos (Alajuela) eastward to Moravia de Chirripo (Limon) and the western part of the Gen- eral Valley (San Jose).
Palicourea lancifera is recognized by its narrow long-petiolate leaves, narrowly thyrsoid greenish inflorescences, slightly larger calyx lobes rounded distally, and white or pale yellow corolla tubes. This species resembles P. albocaerulea with longer calyx lobes and P. leucantha J. D. Smith of Mexico and northern Central America.
Palicourea lasiorrhachis Oersted, Vidensk. Med- del. Dansk. Naturhist. Foren. Kjobenhavn 39. 1 852. P. veraguensis Dwyer, Ann. Missouri Bot. Card. 67: 318. 1980. Figure 51.
Shrubs or small treelets, l-5(-7) m tall, leafy stems 1.8-5 mm thick, glabrous to pubescent with stiff yellow- ish curved hairs to 0.5 mm long; stipules united and forming a broad tube 2-9 mm long, truncated and with narrow teeth 2-5(-8) mm long, glabrous or minutely puberulent, persisting. Leaves opposite, petioles 7-24(-32) mm long, 1-2 mm thick, glabrous or puberulent; leaf blades 6-17(-26) cm long, 2-8(-10) cm broad, elliptic to narrowly ovate-elliptic or elliptic-oblong, apex taper- ing gradually and acuminate, tip 4-13(-20) mm long, base acute to obtuse, drying chartaceous, dark grayish green to brownish above, glabrous above, sparsely to densely puberulent along the major veins beneath with short (0.3 mm) yellowish or longer (1 mm) whitish as- cending hairs (rarely glabrous), 2° veins (8-)l 1-14(-18)/ side and joining a vein along the leaf edge distally. In- florescences 5-1 5(-20) cm long. 4-1 3 cm broad, narrow- ly pyramidal with 4-8 pairs of opposite or subopposite lateral branches from the central rachis, greenish to bright yellow, peduncles 1.5-7(-10)cm long, 0.7-2.8 mm thick, sparsely to densely puberulent with stiff multicellular hairs 0.3-1 mm long, bracts 2-9 mm long, linear-su- bulate, pedicels 1-6 mm long, minutely puberulent, yel- low to bright green, bracteoles 0.5-2 mm long. Flowers sparsely to densely pubescent externally (often glabres- cent), hypanthium 0.5-1.5 mm long, ca. 1 mm diam. distally, obconic, calyx lobes 0.3-1.2 mm long, obtuse: corolla yellow to greenish, tubular-funnelform, tube 6- 15 mm long, narrowed in the middle, ca. 2 mm diam. at the base and 3 mm diam. distally, corolla lobes 1-2.5 mm long, ca. 1.5 mm broad at the base, triangular; an- thers 1.5-2 mm long or 2.5-3 mm long in short-styled form. Fruits 4-5 mm long and 3-4 mm diam., obovoid, with prominent longitudinal ribs, puberulent or glabres- cent, blue or black.
Understory plants of very wet montane rain for- est formations, from (1000-)! 300 to 2600 m el-
evation. Flowering throughout the year (mostly in May-June); probably fruiting throughout the year. This species ranges from the Cordillera de Tilaran along the continental divide and Cordillera de Ta- lamanca to western Panama.
Palicourea lasiorrhachis is recognized by the of- ten narrow leaves, usually puberulent beneath, the minutely puberulent inflorescences (rarely gla- brous) with yellow pedicels, the small calyx lobes, narrowly funnelform yellow corolla tubes, and small obovoid fruit. This common species is quite variable and includes individual collections that differ greatly from each other in regard to leaf size, leaf shape, puberulence, and aspects of flowers and inflorescences. A group of collections from near El Empalme have thicker stems; stifler, more ob- long leaves; secondary veins arising at almost 90° angles; and dense puberulence. This species may be difficult to distinguish from P. vestita (q.v.). It is often confused with P. padifolia, which has red- orange inflorescence branches and more tubular corollas. Also compare P. montivaga (smaller plants with white flowers) and P. adusta (with blue flowers).
Palicourea macrocalyx Stand!.. J. Wash. Acad. Sci. 18: 278. 1928. Figure 49.
Small shrub, subshrubs, or little treelets. 0.5-2(-6) m tall, leafy branchlets 1 .5^4(-10) mm thick, glabrous, flat- tened or quadrangular but quickly becoming terete: stip- ules with a broad thick sheath 2-6 mm long, truncated and with narrow acute teeth 4-10 mm long and 1-3 mm broad, glabrous and persisting. leaves opposite, petioles 7-30(-45) mm long, 0.9-2.2 mm thick, glabrous; leaf blades 5-15(-18) cm long, 2-6(-9.5) cm broad, elliptic- obovate to obovate-oblong or elliptic (rarely broadly ob- long), apex acuminate with tip 4-13 mm long, base ob- tuse to acute (subtruncate). drying stiffly chartaceous to subcoriaceous. olive green or brownish above, glabrous above and below or with small (0.3-0.5 mm) hairs along the midvein, 2° veins 9-1 3/side. Inflorescences 3-8(-l 5) cm long. 2-10 cm broad, pyramidal, yellow or marked with purple, peduncles 1-8.5 cm long, 1-1.8 mm thick, glabrous, bracts ca. 6 mm long, pedicels 1-8 mm long, glabrous, bracteoles 1-4 mm long. Flowers glabrous ex- ternally, hypanthium ca. 1 mm long, narrowly obconic. calyx lobes 3-6 mm long, 1-2 mm broad, narrowly tri- angular to ovate, green or yellow and sometimes marked with purple: corolla tubular, yellow or greenish yellow, tube 8-13 mm long, ca. 2 mm diam.. slightly narrower in the center, corolla lobes 2-3 mm long, obtuse; anthers 1-2 mm long. Fruits 5-6 mm long, 4-5 mm diam., el- lipsoid, becoming blue, longitudinally ribbed (when dried), with persisting calyx to 6 mm long.
Plants of the very wet cloud forests along the continental divide, from 1300 to 2200 m eleva-
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tion. Probably flowering throughout the year; fruiting in April-May and August-October. En- demic to Costa Rica, and collected primarily in the Cordillera de Tilaran, but ranging to near San Isidro de Heredia on the western slopes of Volcan Irazu.
Palicourea macrocalyx is distinguished by the stiff slightly obovate leaves with many secondary veins, lack of pubescence, yellow inflorescences sometimes marked with purple, the larger imbri- cate greenish yellow calyx lobes, and yellow co- rollas. The name of this species should not be confused with P. macrosepala Krause, a synonym of P. discolor. This species is very similar to P. bellula, which has longer corolla tubes.
Palicourea montivaga Standl.. J. Wash. Acad. Sci. 18:279. 1928. Figure 53.
Shrubs, 1-3 m tall (rarely slender treelets to 6 m), leafy stems 0.9-6 mm thick, glabrous or with a few thin hairs ca. 0.3 m long distally, terete; stipules with a tubular sheath 0.5-4 mm long, truncated distally and with tri- angular or linear teeth 0.5-2.5 mm long, glabrous and persisting. Leaves opposite, petioles 6-14(-25) mm long, 0.4-1.1 mm thick, glabrous, slightly sulcate above; leaf blades 3-7(-9) cm long, l-2.5(-3) cm broad, lanceolate or elliptic-lanceolate to narrowly elliptic-oblong, apex tapering gradually and acute or acuminate, tip to 1 5 mm long, base acute (obtuse), drying chartaceous, dark green or brown above, glabrous above, glabrous or with a few thin hairs 0. 1 -0. 5 mm long on the veins beneath, 2° veins 6-9/side. Inflorescences 2-10 cm long, 2-8 cm broad, pyramidal with open opposite branching, peduncles 8- 45 mm long, 0.3-1.2 mm thick, glabrous or minutely puberulent, inflorescence branches usually bright yellow (green), bracts 2-3.5 mm long, linear, pedicels 1-6 mm long. Flowers usually glabrous externally, hypanthium 0.5-1 mm long, ca. 1 mm diam. distally, obconic, calyx lobes 0.4-1 mm long, acute or obtuse; corolla white or yellow, tube 5-9 mm long, 1-1 .3 mm diam., corolla lobes 1-2 mm long; anthers ca. 1.5 mm long. Fruits 4-5 mm long, 3-4 mm diam., obovoid, becoming longitudinally ribbed.
Plants of montane rain forest formations, from (1 1 00-) 1300 to 2300(-2900?) m elevation. Prob- ably flowering throughout the year, but mostly in April-August; fruiting April-November. The spe- cies is endemic and ranges from Volcan Tenorio southward along the continental divide to the western slopes of Volcan Irazu.
Palicourea montivaga is recognized by its small narrow leaves with long tips, glabrous parts, small yellowish inflorescences, small white or yellowish corollas, and montane habitat. This species is closely related to P. adusta with smaller bluish flowers and P. lasiorrhachis with relatively larger
yellowish floral parts and shorter leaf tips. The plant attributed to this species from Panama (Dwyer, 1 980) is P. lasiorrhachis. Material of P. padifolia can also be confused with this species.
Palicourea orosiana C. M. Taylor, Syst. Bot. Monogr. 26: 61. 1989. Figure 51.
Shrubs, 2-8 m tall, leafy stems 1 .8-4 mm thick, stri- gulose with stiff straight hairs to 1 mm long; stipules with a short (1.5-3 mm) sheath (difficult to see under the strigulose hairs) and narrow distal teeth 6-17 mm long, covered with long (1-2 mm) thin hairs. Leaves opposite, petioles 4-12 mm long, 1-1.9 mm thick, puberulent in early stages with straight or curved ascending hairs; leaf blades 7-14 cm long, 2.5-4.5 cm broad, elliptic-oblong to narrowly elliptic, apex acuminate with tip 6-10(-13) mm long, base acute, drying stiffly chartaceous, yellow- ish, upper surface glabrous, glabrous on 2° veins and interveinal areas beneath but with straight stiffhairs 0.5- 1.5 mm long on the sides of the midvein, 2° veins 8-1 3/ side. Inflorescences solitary, 7-1 1 cm long, 7-8 cm broad, pyramidal, yellowish, peduncles 3.5-6.5 cm long and 1 .5 mm thick, strigulose, bracts and bracteoles 3-7 (-1 3) mm long, ca. 1 mm broad, yellowish and resembling the calyx lobes, pedicels 2-6 mm long. Flowers greenish yellow, hypanthium ca. 1.3 mm long and 1 mm diam., densely pubescent, calyx lobes 5-1 1 mm long, 0.8-1 . 1 mm broad near the base, narrowly triangular, acute, sparsely pu- bescent with thin whitish hairs ca. 0.5 mm long, drying yellow; corolla tubular, yellow, tube ca. 10 mm long, with whitish ascending hairs, corolla lobes ca. 2 mm long; anthers ca. 2.5 mm long. Fruits not seen.
Plants of the wet cloud forests of the Caribbean slopes and continental divide, at 1200-2100 m elevation. Flowering in May-June. This species has only been collected above the upper Rio Gran- de de Orosi near Tapanti, Cartago, in Costa Rica; it is also known from western Panama.
Palicourea orosiana is distinguished by its pu- berulent yellowish inflorescence and flowers, larg- er yellowish bracts and calyx lobes, and the long stipule lobes. The pubescence along the midvein on the underside of the leaf is also unusual. This species is similar to P. lasiorrhachis, which has much smaller calyx lobes, and to P. macrocalyx, which has glabrous inflorescences and broadly im- bricate calyx lobes.
Palicourea padifolia (Willd. ex Roem. & Schult.) C. M. Taylor & Lorence, Taxon 34: 669. 1985. Psychotria padifolia Willd. ex Roem. & Schult., Syst. Veg. 5: 189. 1819. Psychotria mexicana Willd. ex Roem. & Schult., loc. cit. 192. 1819. Palicourea costaricensis Benth. ex Oerst., Vi- densk. Meddel. Dansk. Naturhist. Foren. Kjo-
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benhavn 1852: 38. 1853. Palicourea subrubra Polak., Linnaea 41: 571. 1877. P. galeottiana sensu Standley, not M. Martens (see Taylor & Lorence, 1985, cited above). Figure 51.
Shrubs or small trees, (l-)2-7(-10) m tall, leafy stems 1.5-5 mm thick, glabrous (rarely hirsutulous), quadran- gular but becoming terete; stipules with a tubular sheath 1-4 mm long, truncated distally and with narrow awns 2-10 mm long, glabrous. Leaves opposite (rarely 4/node), petioles 4-1 6(-22) mm long, 0.6-1 .8 mm thick, glabrous; leaf blades 6-16(-24) cm long, 2-6(-8.5) cm broad, nar- rowly elliptic-oblong to narrowly oblong, elliptic-ovate or lanceolate (rarely elliptic-obovate), apex acute to acu- minate with tip 4- 1 2(-20) mm long, base acute to obtuse, drying stiffly chartaceous, glabrous and with short (0.2 mm) cystoliths often visible above, glabrous beneath except for thin whitish hairs 0.3-0.9 mm long on the major veins beneath or only along the midvein, 2° veins 8-14/side. Inflorescences (5-)7-18 cm long, 4-14 cm broad, broadly pyramidal, reddish purple to salmon red or orange (yellow), peduncle 1.5-5 cm long, 1.2-2 mm thick, glabrous or sparsely puberulent, bracts ( l-)2-6(-8) mm long and linear-subulate, bracteoles 0.5-2 mm long, pedicels l-6(-10) mm long. Flowers glabrous or sparsely puberulent externally, hypanthium 0.7-1 mm long, 0.5- 1 mm diam., calyx lobes 0.5-1 mm long, triangular, obtuse; corolla tubular to slightly funnelform, carnose, orange or yellow, tube 8-15(-18) mm long and 1.5-2.5 mm diam., narrower in the middle, corolla lobes 2-3 mm long, ca. 1.5 mm broad at the base, obtuse; anthers 2.5-4 mm long. Fruits 4-6(-10) mm long, 4-6(-10) mm diam., ovoid to ellipsoid or globose, longitudinally ribbed, persisting calyx ca. 1 mm high.
Plants of evergreen lower montane forest for- mations and in moist sites in deciduous forest for- mations, (800-)1000-2000(-2400) m elevation. Flowering and fruiting throughout the year (mostly flowering in December-August and fruiting in Jan- uary-August). The species is quite common in Costa Rica and ranges from the Cordilleras de Guanacaste and Tilaran, around the Caribbean side of the central volcanic chain, through the Cor- dillera de Talamanca. The species ranges from eastern Mexico to Panama.
Palicourea padifolia is recognized by its usually narrow leaves, short-tubular stipule sheaths, col- orful inflorescences with (usually) reddish pedi- cels, minute calyx lobes, narrow yellowish (or or- ange) corolla tubes, and montane habitats with high rainfall. This is the most commonly collected species of Palicourea in Central America. This spe- cies is easily mistaken for P. angustifolia. P. crocea, and P. purpurea, but those species have purplish flowers. Palicourea lasiorrhachis is similar but has yellow inflorescence branches and corollas. Pali- courea padifolia is very closely related to P. thyr- siflora (Ruiz & Pavon) Roem. & Schult. of Ecuador
and Peru and may be conspecific, but neither of the two species has been collected in Colombia or the eastern half of Panama. Fruiting plants of this species may be confused with species of Psy- chotria, but the inflorescences are more colorful in P. padifolia.
Palicourea purpurea C. M. Taylor, Syst. Bot. Monogr. 26: 71. 1989. Figure 52.
Shrubs or small trees, (l-)3-6(- 10) m tall, leafy stems 1-4 mm thick, glabrous, stipules with sheaths 1-4 mm long, truncated distally and with slender teeth 2-5 mm long, glabrous and persisting. Leaves opposite, petioles 6-25(-35) mm long, 0.6-1.3 mm thick, glabrous; leaf blades 7-19 cm long, 2-7 cm broad, narrowly elliptic to elliptic-oblong, apex acuminate with tip 5-15 mm long, base acute and slightly decurrent on petiole, drying char- taceous, dark brown above, glabrous above and below or with a few thin white hairs along the side of the mid- vein beneath, 2° veins 6-10/side. Inflorescences 5-1 2 cm long, 4-1 1 cm broad, pyramidal, purplish to deep lav- ender in color, peduncles 2-6 cm long, 1 .2-1 .8 mm thick, glabrous, bracts 2-7 (-10) mm long, linear-subulate, bracteoles 1-3 mm long, pedicels 0-1 1 mm long. Flowers glabrous externally, hypanthium 0.6-1 mm long and 1 mm diam. at apex, obconic, calyx lobes 0.3-1 long and 1 mm broad at the base, triangular; corolla tubular and carnose, purple to lavender, pink, or white, tube 11-18 mm long and 2-4 mm diam.. slightly expanded at the base, corolla lobes 2-4 mm long; anthers 3.5-4 mm long. Fruits 4-6 mm long and 4-5 mm diam., globose, drying black and with longitudinal ridges.
Plants of montane evergreen forest formations, (750-)1 200-2800 m elevation. Flowering in March, May-July, and December; fruiting in January- September and December. The species is found on the Caribbean slope and continental divide of the Central Volcanic chain and in the area above Rio Grande de Orosi and in western and central Panama.
Palicourea purpurea is distinguished by its lon- ger purplish corollas, purple inflorescence branch- es, and globose fruit. It is very similar to P. padi- folia, but that species has yellow or orange corollas and red or orange inflorescence branches, and ovoid or slightly flattened fruit. Palicourea discolor is also similar but has larger leaves, inflorescences, and calyx lobes. Compare P. angustifolia also.
Palicourea salicifolia Standl.. J. Wash. Acad. Sci. 18: 280. 1928. P. austinsmithii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1333. 1938. P. caerulescens Suesseng., Bot. Jahrb. Syst. 72: 286. 1942. Figure 53.
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Shrubs, 0.5-4 tall, leafy stems 1-4 mm thick, glabrous and quadrangular, young stems drying black, pale gray in age; stipules with short (1-2 mm) tubes truncated and with narrow teeth 1-2 mm long, glabrous. Leaves op- posite, petioles 3-9(-15) mm long, 0.4-0.8 mm thick, glabrous; leaf blades 4-8(-10) cm long, l-2(-3) cm broad, narrowly elliptic to lanceolate or lanceolate-oblong, apex usually tapering gradually and acute or acuminate, base acute and slightly decurrent on petiole, drying stiffly chartaceous, glabrous above and below (rarely minutely puberulent along the mid vein), 2° veins 9-13/side. In- florescences 2-6 cm long, 3-6(-8) cm broad, pyramidal, green or marked with blue or purple, peduncles 3-18 mm long, glabrous and usually drying black, bracts 4-7 mm long, 1 mm broad at the base, pedicels 0-8 mm long, bracteoles ca. 3 mm long. Flowers glabrous exter- nally, hypanthium ca. 1.3 mm long and 1 mm diam. distally, calyx lobes 2-3(-4) mm long, ca. 1 mm broad, oblong and rounded distally; corolla funnelform, car- nose, white and often flushed with blue or purple, tube 6-13 mm long, ca. 2 mm diam., narrowed in the middle, corolla lobes 3-6 mm long, to 2 mm broad at the base; anthers 2-4 mm long. Fruits 6-8 mm long and 5-7 mm diam., obovoid or ellipsoid, flattened, becoming deep blue (pedicels also blue), with strong longitudinal ribs (dried).
Plants of evergreen montane forest formations, from 1 500 to 2700 m elevation. Flowering in Jan- uary-September; fruiting in December-March and May-August. The species is known from Monte- verde and the northern part of the Meseta Central (near Zarcero), from the Cerro de Carpintera, and in the Cordillera de Talamanca.
Palicourea salicifolia is recognized by the small narrow leaves tapering at both apex and base, small inflorescences that often dry black, slightly en- larged calyx lobes, and funnelform white corollas marked with blue or purple. The fruit have sharply denned longitudinal ribs. Two collections from be- low 2000 m (Dryer 768 & 965 from Monteverde and Stork 1 175 from Carpintera) have dried pale greenish and the inflorescences are little-branched. Because of its small leaves, this species resembles P. adusta and P. montivaga with shorter corolla tubes and calyx lobes.
Palicourea skotakii C. M. Taylor, Syst. Bot. Monogr. 26: 76. 1989. Figure 49.
Shrubs, 1.5-2 m tall, leafy stems 1.5-5 mm thick, glabrous, quadrangular or flattened; stipules 6-8 mm long, tubular sheath 3-8 mm long, broadly bilobed dis- tally with obtuse lobes ca. 2 mm long and 3 mm broad, glabrous, becoming coriaceous. Leaves opposite, petioles 10-24 mm long, 1-1.6 mm thick, glabrous; leaf blades 7-13(-17) cm long, 3-5-5. 5(-6.5) cm broad, elliptic to elliptic-oblong or elliptic-obovate, apex abruptly nar-
rowed and acuminate, base acute and slightly decurrent on petiole, drying stiffly chartaceous to subcoriaceous, dark greenish above, glabrous above and below, 2° veins 9-14/side and joining a vein along the leaf edge distally, parallel minor 2° veins usually present. Inflorescences 4-10 cm long, 5-1 1 cm broad, usually 3-branched, rounded to pyramidal, peduncle 2.5-4 cm long and 3 mm thick, glabrous, bracts 1 5-30 mm long, pale green (dried), bracteoles to 1 5 mm long, ovate and imbricate around the flowers, persisting. Flowers glabrous exter- nally, calyx lobes 2-6 mm long, lanceolate to ligulate, usually unequal; corolla funnelform, carnose, white, tube ca. 10 mm long, corolla lobes ca. 6 mm long; anthers ca. 2.5 mm long. Fruits 6-7 mm long, 5 mm diam., ellipsoid to slightly obovoid.
Plants of high montane rain forest formations at 2700-2800 m elevation. Flowering in March and June. Endemic; like the type (Taylor & Skotak 4756 DUKE), all collections are from along the In- teramerican Highway in the western part of the Cordillera de Talamanca.
Palicourea skotakii is distinguished by its high- altitude habitat, stipules with broad lobes, lack of pubescence, large greenish ovate persisting bracts and bracteoles, and white corollas. This species resembles P. bella\ it also resembles Psychotria chlorochlamys and its allies.
Palicourea spathacea C. M. Taylor, Syst. Bot. 9: 226. 1984. Figure 49.
Shrubs or small trees, 2-8 m tall, leafy stems 2-8 mm thick, quadrangular, with thin brownish hairs to 0.5 mm long but soon glabrescent; stipules 5-12 mm long, 4-8 mm broad, bilobed, puberulent near the base and along the midrib abaxially. Leaves opposite, petioles 8-20 mm long, 1-2.2 mm thick, appressed pubescent with thin ascending hairs or glabrescent; leaf blades 12-28 cm long, 4.5-14 cm broad, elliptic-obovate to broadly ellip- tic, obovate or elliptic-oblong, apex short-acuminate with tip 8-15 mm long, base narrowed gradually and acute or cuneate, slightly decurrent on petiole, drying stiffly chartaceous, dark green or brown above (much paler beneath), glabrous above, with thin brownish ascending hairs to 1 mm long on the midvein beneath and shorter (0.1-0.3 mm) hairs on the minor venation and surfaces, 2° veins 12-22/side (and usually with a weaker 2° be- tween them). Inflorescences solitary, drooping, 10-23 cm long, 8-16(-20) cm broad, open broadly pyramidal or hemispheric, peduncles 3-9 cm long, 1.5-3.5 mm thick, puberulent near the base, bracts 1 5-20 mm long and 5 mm broad, bracteoles 8-15 mm long, ca. 4 mm broad, pink to purple (white), glabrous pedicels 12-26 mm long, ca. 0.5 mm thick and black when dried. Flow- ers glabrous externally, hypanthium ca. 1.5 mm long and 1.2 mm diam., cylindrical and drying black, calyx tube spathe-like and 1 5-20 mm long by 5-8 mm diam., split- ting open along 1 side, pink to purple; corolla tubular, camose, white, tube 3-4 cm long, 4-5 mm diam., straight
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or slightly curved, corolla lobes 4-7 mm long; anthers 4.5-5 mm long. Fruits ca. 8 mm long (not including the persisting calyx), ca. 7 mm in diam., dark purple but drying black and with longitudinal ribs.
Plants of the very wet lower montane rain forest formations on the Caribbean slope of the Cordil- lera de Talamanca, at 1300-1700 m elevation. Flowering in March, April, and November-De- cember; fruiting in April-May, July-August, and December. This species is only known from the upper drainage area of Rio Grande de Orosi near Tapanti, Cartago Province.
Palicourea spathacea is a striking species; the bright pink or purplish bracts, bracteoles, and large spathaceous calyx make the inflorescences es- pecially conspicuous. The long white corollas, large leaves with many secondary veins and cuneate bases, and stipules with a single obtuse apex on each side are further distinctions. The calyx is in- flated before anthesis and splits for half to three- quarters of its length as the corolla emerges; there are no calyx lobes.
Palicourea standleyana C. M. Taylor, Syst. Hot. Monogr. 26: 81. 1989. Psychotria brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1347. 1938, not Palicourea brenesii ; Standl., 1938. Figure 54.
Shrubs or small trees, 2-7 m tall, leafy branchlets 3- 6 mm thick, densely pilose with thin yellowish hairs 1-
3 mm long; stipules (6-)8-18 mm long, 3-8 mm broad, with a tubular sheath 4-8 mm long, acute or with 2 short (1-2 mm) lobes, sparsely pilose. Leaves with petioles 1-
4 cm long, 0.7-1.7 mm thick, pilose with straight or slightly curved hairs to 2 mm long; leaf blades 9-23 cm long, 3-9 cm broad, broadly elliptic, elliptic-ovate or elliptic-obovate, apex acuminate to caudate-acuminate with 8-15(-20) mm long, base obtuse to acute, drying stiffly chartaceous, dark brown above, densely pilose above and below with hairs 1-1.5 mm long, 2° veins (8-) 10-1 7/side, connected distally to make a slightly ar- cuate submarginal vein. Inflorescences terminal, 3 or 1 with 3 main branches from a short (0-1 5 mm) common peduncle, 7-12 cm long, to 14 cm broad, open trichot- omous panicles, primary (basal) lateral branches 3-7 cm long, pilose, bracts 4-1 1 mm long, ca. 1.5 mm broad, lanceolate, flowers in close distal clusters subtended by ovate pilose bracts, pedicels 0-5 long. Flowers puberu- lent externally, hypanthium ca. 2 mm long, calyx lobes 2-4 mm long but difficult to see among the hairs; corolla tubular-funnelform, camose, deep lemon yellow to greenish yellow, tube 10-12 mm long, 1.5-2 mm diam., expanded at the base and with hairs to 2 mm long, corolla lobes 3-5 mm long; anthers 2.5-3 mm long. Fruits 6-8 mm long (not including the 3-mm-long calyx), 5 mm diam., ellipsoid, pilose, becoming blue.
Plants of very wet cloud forest formations along the Caribbean slope and continental divide, 1 100- 1600 m elevation. Flowering in December-July; fruiting in January-June. The species has been col- lected in the Cordilleras de Guanacaste and Ti- laran, north of San Ramon (Alajuela), near Ta- panti (Cartago), in Chiriqui, Panama, and Narino Province, Colombia.
Palicourea standleyana is recognized by the prominent pubescence on all parts, the large tu- bular stipules, stiff leaves with many secondary veins and submarginal veins, open three-branched inflorescences, and bright yellow corollas. This species is easily confused with Psychotria pilosa, but that species has much smaller corollas and smaller fruit and the corolla tubes lack the internal ring of the hairs that define Palicourea.
Palicourea tilaranensis C. M. Taylor, Syst. Bot. Monogr. 26: 84. 1989. Figure 49.
Small shrubs, ca. 1.5 m tall, leafy stems ca. 3 mm thick, quadrangular and glabrous, young stems drying black; stipules 5-8 mm long, with a very short (0.5-1.2 mm) tubular sheath, stipule lobes to 4 mm long, broadly obtuse to deeply bilobed, persisting. Leaves opposite, petioles 13-40(-65) mm long, 0.7-1.7 mm thick, gla- brous; leaf blades 8-16 cm long, 3.5-6 cm thick, ovate to ovate-elliptic or ovate-oblong, apex acuminate with tip 7-14 mm long, base obtuse, drying stiffly chartaceous, dark greenish brown, glabrous above and below, 2° veins 8-1 I/side. Inflorescences 12-15 cm long. 6-9 cm broad, pyramidal panicles with opposite branching, peduncles 4-6 cm long, 1 .5-2 mm thick, glabrous, bracts 8-1 3 mm long, 2-4 mm broad, lanceolate, bracteoles ca. 5 mm long and 2 mm broad, flowers sessile or on pedicels to 4 mm long within the persisting bracteoles. Flowers gla- brous externally, hypanthium 0.5-1 mm long, calyx lobes 0.3-1.5 mm long, triangular; corolla tubular, carnose, white, tube 3-7 mm long, ca. 1.8 mm diam., slightly narrowed in the middle, corolla lobes ca. 2 mm long. Fruits not seen.
Plants of evergreen lower montane rain forest formations, at about 15500-1800 m elevation. Flowering in February and June. This species is known only from the Monteverde Nature Reserve, in the Cordillera de Tilaran.
Palicourea tilaranensis is recognized by its re- stricted range, large and persisting bracts and brac- teoles, small calyx lobes, and white corollas. It resembles Psychotria palicoureoides and its allies.
Palicourea triphylla DC., Prodr. 4: 526. 1 830. Psy- chotria triphylla (DC.) Muell.-Arg. in Mart., Fl.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
21
Bras. 6(5): 233. 1&&1. Palicourea longibracteata Bartlingex DC., Prodr. 4: 527. 1830. Palicourea parviflora Benth., Bot. voy. Sulph. 107. 1844. Figure 50.
Shrubs or herbaceous subshrubs, l-3(-5) m tall, erect and usually unbranched (rarely clambering), leafy inter- nodes 2.3-7 mm thick, glabrous or sparsely pubescent on the new growth and nodes with short (0.1-0.3 mm) thin hairs, terete; stipules united for only 1-2 mm at the base or free, stipule lobes 6/node. 6-12(-15) mm long and ca. 2 mm broad at the base, narrowly triangular and acute, glabrous abaxially and ciliolate along the edge. Leaves 3/node (rarely 2 or 4), petioles 3-7(-l 5) mm long, 1.4-2 mm broad, minutely pubescent but glabrescent; leaf blades 7-20 cm long, 3-8 cm broad, narrowly ellip- tic-oblong to lanceolate, narrowly elliptic or oblanceo- late, apex tapering gradually and acuminate with tip 5- 1 5 mm long, base cuneate to acute (obtuse) and slightly decurrent on petiole, drying thinly to stiffly chartaceous, minute (0.05-0.2 mm) thin hairs along the major veins above, with thin whitish hairs 0.1-0.3 mm long on the veins and surfaces beneath, 2° veins 7-1 l(-15)/side. In- florescences 6-20 cm long (to 26 cm in fruit), 5-10 cm broad near the base, pyramidal to narrowly thyrsoid- cylindrical, red to orange or orange-yellow, peduncles 5- 12(-18) cm long, 1.5-3 mm thick, puberulent with thin whitish hairs, bracts 5-20(-30) mm long linear-subulate, pedicels 1-5 mm long, puberulent. Flowers distylous, minutely puberulent, hypanthium ca. 1 mm long and 0.5 mm diam., tubular, calyx lobes 0.2-1 mm long, broadly triangular; corolla tubular or slightly funnelform, yellow or reddish distally, tube 8-14 mm long, 1.5-3 mm diam., inflated at the base, corolla lobes 1-2 mm long; anthers 2.5-3.5 mm long. Fruits 4-5 mm long, 3- 5 mm diam., ovoid to globose with prominent longi- tudinal ridges (dried), becoming blue, purple, or black at maturity, glabrous or puberulent, calyx ca. 0.7 mm high.
Plants of open wet or poorly drained sties in evergreen or partly deciduous forest formations in the Caribbean lowlands and on the Pacific slope of central and southern Costa Rica and in Panama, from near sea level to 700 m elevation, but to 1 300 m on the semideciduous Pacific slope. Flowering and fruiting throughout the year (but flowering mostly in April-July). The species ranges from the Caribbean side of central Mexico and Cuba to Brazil and Bolivia.
Palicourea triphylla is recognized by its ternate leaves, puberulence of short thin hairs, long per- sisting stipules with six teeth per node, orange- yellow inflorescences with persisting bracts, small corolla lobes, and gibbous corollas with multicel- lular hairs. The species is often found in open swampy sites.
Palicourea vestita Standl., J. Wash. Acad. Sci. 18: 277. 1928. Figure 51.
Shrubs or small treelets, 1-4 m tall, leafy stems 1.5- 5 mm thick, densely hirsutulous with yellowish hairs 0.3-0.9 mm long; stipules with a tube 3-6 mm long, with narrow teeth 3-8 mm long, puberulent, persisting. Leaves opposite, petioles 6-1 8(-30) mm long, 0.8-1 .6(-2.2) thick, densely puberulent with yellowish hairs; leaf blades (6-)8.5-19 cm long, (1.7-)2.5-7 cm broad, elliptic to elliptic-oblong, elliptic-obovate or lanceolate, apex acu- minate (acute) with tip to 1 cm long, base acute to cuneate (obtuse), drying stiffly chartaceous, often yellowish green, glabrous above or with thin hairs along the midvein, pubescent along the major veins beneath with thin yel- lowish hairs 0.4-1.2 mm long, 2° veins 12-20/side and with minor short parallel 2° veins between them. Inflo- rescences 5-12 cm long, 5-8 cm broad near the base, thyrsiform pyramidal, yellowish, peduncles 2-8 cm long, 1-3 mm thick, densely pubescent, bracts 4-10 mm long and 1.5 mm broad, bracteoles 1-4 mm long and per- sisting, pedicels 2-5 mm long. Flowers sparsely and mi- nutely puberulent externally, hypanthium 1-1.5 mm long, obconic, calyx lobes 0.7-1.5 mm long, acute; corolla bright yellow, tube 7-13 mm long, ca. 2 mm diam., slightly inflated at the base, corolla lobes 1.5-3 mm long; anthers 2.1-2.5 mm long. Fruits 5-8 mm long, 3.5-6 mm diam., obovoid, persisting calyx ca. 0.7 mm high.
Plants of evergreen montane wet forest forma- tions of the Cordillera de Talamanca, from 1 200 to 2600 m elevation. Flowering in April-August; fruiting in June-August, October, and December. The species is known only from Costa Rica and western Panama.
Palicourea vestita is recognized by the short yel- lowish hairs on most plant parts, the leaves with many secondary veins, the elongate thyrsiform in- florescences, and the bright yellow corolla tube. This species is distinguished from the very similar but more common P. lasiorrhachis by the more numerous secondary veins and longer calyx lobes.
Pentagonia Bentham Nomen conservandum
Shrubs or small trees (rarely large trees), main stems often unbranched or with few lateral branches, stems thick, flattened, quadrangular or terete in early stages, glabrous or puberulent, often with conspicuous rounded lenticels; stipules interpetiolar, free, large, triangular, usually early deciduous. Leaves opposite and decussate, usually very large, sessile or petiolate, the petioles some- times with auriculate (leafy) developments at the base; leaf blades entire or pinnatind, usually drying coriaceous, the minor venation lineolate-parallel and often with 2 distinct orientations in the same leaf area (parallel with the tertiary veins and at right angles to the tertiary veins),
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FIELDIANA: BOTANY
domatia absent. Inflorescences axillary to distal leaves, usually short-pedunculate, cymose or corymbose to glo- merulate, generally with fewer than 25 flowers, bracts developed or minute, flowers subsessile or pedicellate. Flowers bisexual and monomorphic, radially symmet- rical, often large, usually densely pubescent externally, hypanthium conical to turbinate or campanulate, calyx tube well developed, calyx lobes 5-6 or spathaceous, lobes equal or unequal, often with glands inside at the base; corolla tubular to funnelform, carnose. white to yellow or red, glabrous within the throat, villous at the stamen attachment, corolla lobes 5-6, valvate in bud, short; stamens 5-6, filaments borne on the middle of the tube, equal or unequal, usually villous at the base, an- thers dorsifixed, included; ovary 2-locular, ovules many on expanded elongate placentas borne on the septum, stigmas subcapitate or branched. Fruits baccate, fleshy or becoming hard and nut-like when dry, usually globose and the surface often lenticellate-muricate, 2-locular, the large calyx often persisting on the mature fruit; seeds many, angular.
A genus of about 20 species, ranging from Gua- temala into northern and western South America. The very large leaves with distinctive minor ve- nation, the small axillary inflorescences with larger crowded flowers, the stiff pubescent tubular co- rollas, and the globose indehiscent fruit with large persisting calyx distinguish this genus. The very large leaves are correlated with a lack of lateral branches on a solitary vertical trunk in many spe- cies. The leaves are often reddish or purplish be-. neath, making the plants even more striking. The genus is seriously in need of revision. The large leaves often very variable in form and size, the compact inflorescences resulting in crushed or hid- den flowers, and a paucity of collections account for the fact that the species are still poorly under- stood.
Key to the Species of Pentagonia
la. Leaf blades entire, never lobed distally; Caribbean lowlands and central cordilleras 2a
Ib. Leaf blades conspicuously pinnatind or pinnately lobed; Pacific evergreen lowlands 6a
2a. Leaves and stems with hairs ca. 1 .4 mm long, leaves sessile, the leaf blade slightly auriculate near the stem P. hirsula
2b. Leaves and stems with hairs less than 0.5mm long, leaf blades petiolate and never with expanded auriculate tissue near the base 3a
3a. Calyx lobes usually less than 3 mm long; inflorescences usually open and branched, with the branches easily seen at anthesis P. costaricensis
3b. Calyx lobes usually more than 4 mm long; inflorescences compact, the branches of the inflo- rescence short and usually difficult to see (except in fruit) 4a
4a. Bracts of the inflorescence minute or undeveloped; calyx often absent at the apex of the mature
fruit; leaves obtuse to rounded/subtruncate at the base; Guatemala to Costa Rica
P. donnell-smithii
4b. Bracts of the inflorescence conspicuous in early stages, more than 10 m long and 4 mm broad; calyx persistent on the mature fruit; leaves rarely rounded or subtruncate at the base; south- ernmost Costa Rica and Panama 5a
5a. Leaves subsessile with a thick petiole less than 2 cm long, lamina long-cuneate at the base; floral bracts 5-10 mm long P. wendlandii
5b. Leaves with well-developed petioles more than 3 cm long, lamina obtuse to cuneate at the base;
bracts 10-18 mm long P. macrophylla
6a. Leaf blade long-decurrent (at its base) on the petiole . . P. linajita
6b. Leaf blade not at all decurrent on the petiole P. gymnopoda
Pentagonia costaricensis (Standley) W. Burger & C. M. Taylor, comb. nov. Nothophlebia costari- censis Standl., Contr. U.S. Natl. Herb. 17: 438. 1914. Figure 14.
Trees, 5-18 m tall, leaf stems 6-12(-22) mm thick, glabrous or subglabrous, flattened-quadrangular in early stages, often with dark round lenticels ca. 1 mm diam.;
stipules 2-6(-8) cm long, lanceolate, glabrous or mi- nutely appressed-sericeous. Leaves with petioles (2.5-)4- 7 cm long. 2.5-7 mm thick, glabrous or subglabrous and drying brown; leaf blades (2 l-)27-l 00 cm long, ( 1 2-)l 6- 50 cm broad, broadly elliptic-obovate to very broadly obovate, apex abruptly narrowed and broadly obtuse, base obtuse and only slightly decurrent (rarely slightly auriculate at the petiole as in Gomez- Laurito 9321 CR, F), margin entire, drying coriaceous, glabrous above and
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
213
below or with minute sericeous hairs on the veins be- neath, 2° veins 8- 11 /side, the minor venation closely parallel in 2 different directions (but both systems not always apparent). Inflorescences 3-12 cm long, 3-15 cm broad, often dichotomously branched, the primary pe- duncle 5-20 mm long, secondary branches 5-20 mm long, major branches lacking subtending bracts, ultimate branches usually with 3-flowered cymes, striate and gla- brous with elongate (0.4-2 mm) lenticels, pedicels 0-7 mm long. Flowers with hypanthium 2-4 mm long and 2-3 mm diam., obconic, minutely puberulent or gla- brous externally, calyx tube 4—7(-12) mm long, 4.5-6 mm diam., cupular to short-tubular, with parallel ve- nation, rose-colored, calyx lobes 0.5-2 mm high, 2-4 mm broad, broadly rounded to obtuse or obscure, vari- able and often unequal; corolla cream white or yellow, tube 12-20 mm long, 3-5 mm diam., minutely yellowish puberulent externally, corolla lobes 3-6 mm long; an- thers white. Fruits (based on Hammel& Grayum 14292) 10-20 mm diam., globose with a persistent calyx tube 4-5 mm high and 4 mm diam. at the top, calyx lobes often obscure.
Trees of evergreen lowland rain forest forma- tions, 5-600(-900) m elevation. Flowering in April-May and November; fruiting in May, July- September, and November. The species is known only from the Caribbean lowlands, from northern Costa Rica to western Panama.
Pentagonia costaricensis is recognized by the very large leaves, unusual minor venation, branched inflorescences, short calyx lobes, and smaller fruit. Standley erected the genus Nothophlebia for this unusual species but admitted (Standley 1938, p. 1329) that it might be referred to Pentagonia. A collection from 1300-1400 m near Las Alturas (Almeda et al. 6699 F) with larger calyx lobes and glabrous corolla tube is tentatively placed here.
Pentagonia donnell-smithii (Standl.) Standl., J. Wash. Acad. Sci. 17: 170. 1927. Watsonamra donnell-smithii Standl., Contr. U.S. Natl. Herb. 17: 442. 1914. Figure 14.
Small trees or treelets, 2-7(-12) m tall, often with a single stem, leafy stems 5-22 mm thick, at first ap- pressed-sericeous but quickly becoming glabrous; stip- ules 2.5-9 cm long, 6-25 mm broad, lanceolate to nar- rowly ovate, with appressed-ascending sericeous hairs 0.2-0.5 mm long externally. Leaves often reddish be- neath when young, petioles (2-)3-l 2 cm long, 1 .5-6 mm thick, appressed-puberulent with thin ascending hairs ca. 0.2 mm long; leaf blades (15-)26-90 cm long, (9-) 13- 50 cm broad, broadly elliptic to broadly elliptic-oblong or broadly ovate, apex bluntly obtuse or rounded, base rounded and subtruncate to cuneate-obtuse, margin en- tire and without lobes, drying chartaceous to subcoria- ceous, pale to dark grayish green, glabrous above, sparse- ly to moderately puberulent with thin ascending hairs on the veins beneath, 2° veins 10-14/side, minor ve-
nation closely parallel. Inflorescences 2.5-5 cm long, to 8 cm broad (including both inflorescences of the node), primary peduncle 1-2 cm long and 2-3 mm thick branches of the inflorescence usually short and obscured by the flowers, bracts 0-2 mm long, pedicels 0-6 mm long. Flowers with hypanthium 3-7 mm long, ca. 3 mm diam. distally, calyx tube 6-10 mm long, ca. 6 mm diam., minutely ascending sericeous, calyx lobes 4-8 mm long, 3—4 mm broad, broadly rounded distally, corolla yellow or white, corolla tube 1 5-30 mm long, 3-5(-6) mm diam. and broadest near the base, with short thin hairs ca. 0.2 mm long, corolla lobes 5, 5-10 mm long, acute. Fruits 16-40 mm diam., globose or globose-ovoid, outer wall hard and 2-3 mm thick, with small (0.5 mm) tuberculate lenticels, calyx deciduous or less often persisting on the mature fruit; seeds 3-5 mm long, ellipsoid, orange.
Trees of evergreen rain forest formations on the Caribbean slope and lowlands of Costa Rica, from near sea level to 900 m elevation. Flowering in March-July and October-November in Costa Rica; probably fruiting throughout the year. The species ranges from Guatemala southward along the Ca- ribbean to central Costa Rica.
Pentagonia donnell-smithii is recognized by the very large leaves with unusual minor venation, the small compact inflorescences lacking developed bracts, the prominent calyx lobes, stifFcorolla tubes, and usually globose fruit. The mature fruit in most Costa Rican collections are lacking the calyx, but the calyx persists regularly in Guatemala and Hon- duras. Standley (1938) suggested that this species be submerged in P. macrophylla. But while the two species appear to be closely related, P. macro- phylla has a red calyx, white corolla, and large distinctive bracts subtending the branches of the inflorescence in early stages. No such bracts are seen in P. donnell-smithii. Also, this species does not appear to occur in eastern Costa Rica, where intergradation might be expected.
Sterile material of these species, and of P. cos- taricensis and P. wendlandii, may not be separable, since there is considerable variation within each species. For example, a dwarf treelet 1 m tall with very short (2 cm) petioles and slightly obovate leaves resembling the leaves off. wendlandii was collected near Puerto Viejo de Sarapiqui (A. Ji- menez 3424 CR). This collection lacks the bracts that are found in both P. macrophylla and P. wend- landii and is therefore placed here under P. don- nell-smithii.
Pentagonia gymnopoda (Standl.) Standl., J. Wash. Acad. Sci. 17: 171. 1927. Watsonamra gym- nopoda Standl., Contr. U.S. Natl. Herb. 1 7: 444. 1914.
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FIELDIANA: BOTANY
Small treelets, 1-6 m tall, usually with a vertical un- branched stem, leafy stems 6-16 mm thick, somewhat flattened in early stages but becoming terete, glabrous, grayish; stipules 4-6 cm long, 2-3.5 cm broad, broadly triangular to lanceolate, apparently glabrous or minutely appressed-puberulent. Leaves with petioles 5-12 cm long, 2.7-5 mm thick, glabrous or minutely (0. 1 mm) puber- ulent; leaf blades 28-70(-100) cm long, 30-45(-70) cm broad, ovate to oblong in general outline but deeply pinnatitid to the distal part of the blade, with 5-7 major oblong lobes, the lobes 5-25(-35) cm long and 2-7(-8) cm broad, apex acute to acuminate, base truncate to acute, drying chartaceous, dull grayish green above, gla- brous above, minutely (0.1-0.2 mm) papillate-puberu- lent on the veins beneath, 2° veins 5-9/side. Inflores- cences closely crowded in the leaf axils, 2-5 cm long, subsessile and partly covered by the broad stipules or with a peduncle up to 1 cm long, bracts to 3 cm long, flowers closely crowded and apparently sessile. Flowers with hypanthium ca. 10 mm long, densely appressed- puberulent, calyx tube 6-10 mm long and ca. 6 mm diam., calyx lobes 5, 18-15 mm long, 5-8 mm broad, stiff and parallel veined; corolla white to yellow or pale green, tube ca. 20 mm long, puberulent externally, co- rolla lobes ca. 10 mm long; stamens 5, filaments 8-10 mm long and unequal, anthers ca. 3 mm long; style ca. 9 mm long. Fruits 15-20 mm diam., globose to ovate or pyriform, the persisting calyx to 22 mm long and 6 mm diam.
Poorly known plants of evergreen lowland rain forest formations. Flowering in January-Febru- ary; fruiting in January-February. The species ranges from the Carara Biological Reserve (84°35'W) southeastward along the Pacific low- lands to central Panama.
Pentagonia gymnopoda is recognized by the very large deeply pinnatifid leaves, small axillary inflo- rescences, congested flowers, and sessile fruit. The broad stipules often cover part of the inflorescence. Currently known from only about four collections in Costa Rica. Compare the closely similar P. ti- najita.
Pentagonia hirsute Standl., J. Wash. Acad. Sci. 17: 170. 1927.
Probably small treelets, young branches ca. 10 mm thick, hirsute; stipules not seen. Leaves sessile and entire; leaf blades ca. 60 cm long and 27 cm broad, broadly obovate or broadly elliptic-obovate, apex short-acumi- nate, lamina gradually narrowed below the middle and cuneate but merging with a 3-cm-broad basal region that is slightly expanded (auriculate) and cordate-clasping at the stem, the auricles ca. 2 cm wide on each side, drying chartaceous, the upper and lower surfaces with numerous thin straight hairs 0.7-1 .8 mm long, 2° veins 14-1 7/side, with 1-3 branches distally and obscurely loop-connected near the margin. Inflorescences poorly known, small, with crowded sessile flowers in the leaf axils. Flowers
poorly known, hypanthium densely hirsute, calyx ca. 24 mm long (including both tube and lobes), membranous, hirsute with whitish hairs. Fruits unknown.
Pentagonia hirsuta is a poorly known species from about 500 m elevation near Tsaki in the Talamanca valley, Limon Province, on the Carib- bean slope of easternmost Costa Rica. The type and only collection (Tonduz 9415 F, us) was made in March 1 895. The long hairs on stems and leaves are very unusual within Pentagonia. The leaf shape resembles that of P. wendlandii in the long base but not in its shape; P. wendlandii seems to have more narrowly obovate leaves not as auriculate at the base (and without the long hairs).
Pentagonia macrophylla Ik-nth.. Hot. voy. Sulph. 105, t. 39. 1845. Watsonamra macrophylla (Benth.) Kuntze, Rev. gen. pi. 302. 1891. W. pubescens Standl., Contr. U.S. Natl. Herb. 17: 441. 1914.
Trees or shrubs to 5(-10) m tall, often unbranched. leafy twigs 6-12 mm thick, minutely (0.1-0.3 mm) pu- berulent, quadrangular or becoming terete; stipules 3-7 cm long, 12-15 mm broad at the base, triangular and acuminate, glabrous or puberulent. Leaves with petioles 3-9(-12) cm long, 2.5-5 mm thick, minutely puberulent or glabrescent; leaf blades 22^45(-65) cm long 1 1-28(-37) cm broad, elliptic to elliptic-obovate or very broadly elliptic, apex gradually narrowed and acute, base grad- ually narrowed and obtuse or acute, drying subcoria- ceous, grayish, glabrous above, minutely puberulent be- neath, 2° veins 9-13/side, minor venation parallel. Inflorescences axillary, ca. 5 cm long (including the flow- ers), dense corymbs of 3-7 flowers, sessile or with short (ca. 1 cm) peduncles, bracts 10-18 mm long. 6-9 mm broad, oblong-ovate, apex obtuse, red, with many par- allel veins, minutely sericeous or glabrous with a ciliolate margin, pedicels 0-4 mm long. Flowers ca. 3 cm long, hypanthium 6-8 mm long, glabrous or minutely seri- ceous, calyx lobes 5, 6-12 mm long (becoming 20 mm long in fruit), to 7 mm broad, usually red; corollas yellow, sparsely to densely sericeous externally, tube 14-40 mm long, 3-10 mm diam., corolla lobes 5, 4-7 mm long, ovate with acute apices; stamens differing in size, anthers 3.5-8 mm long; style 20-25 mm long. Fruits 1 5-28 mm diam., globose beneath the persisting calyx, red or or- ange; seeds ca. 4 mm long.
Plants of lowland evergreen forest vegetation, at 10-900 m elevation in Panama. Flowering mostly in late April-September in central Panama (Croat, 1978). The species ranges from near the Panama border in Limon Province to Colombia.
Pentagonia macrophylla is distinguished from its congeners by the larger bright red bracts and red calyx lobes. We have not seen material of this
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
215
species collected in Costa Rica, but the species was studied near BriBri (Limon) by Lucinda McDade (see Oecologia 68: 218-223. 1986).
Pentagonia tinajita Seem., Bot. voy. Herald. 134. 1854. P. alfaroana Standl., J. Wash. Acad. Sci. 17: 171. 1927. Figure 14.
Small unbranched treelets, l-3(-4) m tall, stems at first with 4 prominent ridges or flattened soon becoming terete, leafy stems 5-12 mm thick, glabrous; stipules 3- 5.5 long, 1.2-2 cm broad, ovate-oblong, obtuse or bifid with striate parallel venation, puberulent along the mid- rib. Leaves with petioles 0-1 1(-20) cm long (variable in length due to the decurrent lamina base), 2.5-4.5 mm thick; leaf blades 30-80(-100) cm long, 22^10 cm broad, pinnately lobed and ovate in general outline (from a cuneate base), the sinuses 5-15 cm deep near the base and becoming more shallow distally, pinnatifid in the lower VB of the lamina, sinuses usually rounded, usually obtuse at the apices, abruptly narrowed and cuneate at the base with long-decurrent lamina margins running down to the petiole or to the leaf base, leaves drying thin-chartaceous and dark green above (paler green be- neath), with few short (0.2 mm) hairs on the upper sur- faces, with more frequent short (0.3-0.4 mm) thin hairs on the lower surfaces, 2° veins 6-12/side, minor veins 0.2-0.4 mm apart. Inflorescences sessile or subsessile, densely fasciculate, bracts to 2 cm long and 3-4 mm broad, with parallel longitudinal veins and difficult to distinguish from the calyx lobes. Flowers with hypan- thium ca. 5 mm long, calyx tube 10-15 mm long, calyx lobes 4-8 mm long, 3-4 mm broad, with the same stiff texture brown color and parallel venation as the bracts; corolla tube 2-2.5 cm long, 4 mm diam. at the base and 8 mm distally, stiff, corolla lobes 6-8 mm long. Fruits 10-20 mm long and 1 2-1 5 mm diam., globose or ovoid, the persisting calyx 10-20 mm long (tube and lobes).
Poorly known plants of lowland rain forest for- mations, 0-200 m elevation. Flowering in Janu- ary; fruiting in February and April. The species ranges along the Pacific lowlands of central Costa Rica to western Panama.
Pentagonia tinajita is recognized by the large thin pinnatifid leaves with decurrent lamina base, congested axillary inflorescences, and smaller fruit. Pentagonia alfaroana (based on Standley 40194 F, us) was said to differ because the larger leaf lobes are also pinnatifid, but leaf lobing appears to vary greatly and does not appear to be a sound basis for separating a species. Compare P. gymnopoda, which may be conspecific.
Pentagonia wendlandii Hook., Bot. Mag. pi. 5230. 1861. Watsonamra wendlandii (Hook.) Kuntze, Rev.gen.pl. 320. 1891.
Shrubs or small treelets to 3 m tall, main stems usually unbranched and with a cluster of leaves at the top, leafy stems 8-20 mm thick, with 4 prominent ridges, minutely appressed-puberulent or glabrous; stipules 2.5-6.5 cm long, lanceolate to ovate-oblong, minutely appressed- puberulent. Leaves entire and subsessile, petioles 5-15 mm long, 6-10 m thick; leaf blades 50-100 cm long, 25-50 cm broad, narrowly obovate to obovate-oblong, apex broadly obtuse or rounded, gradually narrowed be- low the middle, base cuneate and slightly auriculate, leaves drying stiffly chartaceous or subcoriaceous and brown, glabrous above, minutely (0.1 mm) puberulent on the veins beneath or glabrescent, 2° veins 14-16/side. Inflo- rescences to 5 cm long, with ca. 6-15 flowers, peduncles ca. 5 mm long, bracts 6-10 mm long, ca. 5 mm broad, oblong and brown, with whitish sericeous hairs along the midrib, flowers subsessile (or with pedicels to 1 0 mm long, fide Dwyer). Flowers (from Dwyer, 1980) with a calyx tube ca. 10 mm long, campanulate, purplish red, stiff, puberulent, with numerous glands at the base with- in, calyx lobes 5, slightly unequal, 5-10 mm long, 1 calyx lobe usually short and acute; corolla yellow, tube ca. 25 mm long, narrowly cylindrical, glabrous to puberulent externally, corolla lobes 4-7 mm long, 2.2-2.6 mm wide, oblong to triangular; stamens 5, filaments to 1 6 mm long, unequal, anthers ca. 4 mm long, oblong; style ca. 18 mm long, stigmas 3.5 mm long. Fruits to 45 mm long, oblong rotund, drying black, the persisting calyx to 1 5 mm long.
Pentagonia wendlandii occurs in Caribbean lowland rain forest formations, from near sea level to ca. 500 m elevation. The species ranges from Bocas del Toro Province to central Panama, but we have seen no flowering or fruiting material from Costa Rica. (A leaf associated with Gomez- Laurito 9321 and collected near BriBri may be this species, but the flowering sheet appears to be P. costari- censis.) The species appears to be closely related to P. macrophylla Benth. of central Panama but differs in the long-cuneate leaf base with a very short petiole. See the discussion under P. donnell- smithii.
Pentas Bentham
Herbs or shrubs, stems erect or clambering, puberulent in most species, from a fibrous or woody rootstock; stip- ules interpetiolar with 2-many awns or narrow lobes. Leaves opposite or in whorls of 3-5, petiolate; leaf blades entire and usually narrowly ovate to lanceolate, pin- nately veined, without domatia. Inflorescences terminal or axillary to the distal leaf pair, usually much-branched and cymose. Flowers bisexual, radially symmetrical, monomorphic, dimorphic, or trimorphic, calyx lobes 5, equal or unequal with 1-3 longer than the others; corolla usually salverform with a narrow tube, throat pilose within, corolla lobes 5, ovate to oblong or lanceolate; stamens usually included, anthers usually linear and with short filaments; ovary 2- or 3-locular, with many ovules in each locule. Fruits capsules, usually ovoid and lon- gitudinally ribbed, beaked, splitting open at apex into 2
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or 4 valves; seeds minute, tetrahedral or subglobose. with reticulate testa.
A genus of ca. 40 species of Africa, Arabia, and Madagascar. The following species is a popular garden ornamental with bright red, rose, lilac, or white flowers.
Pentas lanceoiata (Forssk.) Deflers, Voy. Yemen, 142. 1889. Ophiorhiza lanceoiata Forssk., F. Aegypt.-Arab. 42. 1775. Figure 31.
Herbs or subshrubs with erect or clambering stems, 0.5-2 m tall, stems 1.5-5 mm thick, puberulent with crooked whitish hairs ca. 0.5 mm long; stipules 2-9 mm long, with a short base and 3-9 slender setae, persisting. Leaves opposite, petioles 0-5(-l 5) mm long, puberulent; leaf blades (3-)4-13(-18) cm long, (l-)2-6 cm broad, ovate-lanceolate to lanceolate or elliptic, apex acute, base acute and slightly decurrent on petiole, drying stiffly chartaceous, sparsely to densely puberulent above and below, 2° veins 6-9/side and strongly ascending. Inflo- rescences terminal or with the distal leaf pair subtending the lateral branches, 3-9 cm long and equally broad, subcapitate or corymbose and hemispheric, puberulent. Flowers often trimorphic (style exserted and anthers in- cluded, anthers exserted and style included, both anthers and style included), calyx tube 1-3 mm long, calyx lobes 5, usually unequal with the longer 4-12 mm long; corolla brilliant rose red to lilac or white, tube 14-25(-40) mm long, ca. 0.5 mm diam. but enlarged in the distal 2-6 mm and 3-6 mm diam., throat filled with erect whitish hairs, corolla lobes 3-10 mm long, acute, ovate-lanceo- late to ovate-oblong, acute.
A popular ornamental grown in gardens around the world. In Central America plants with deep red or pinkish red corollas are most common. It has become naturalized in Colombia. Compare Ixora coccinea, another garden favorite.
Pentodon Hochsteter
Annual or short-lived herbs, glabrous; stipules united to the petiole bases and forming a short truncated sheath with minute distal lobes. Leaves opposite, sessile or short- petiolate, entire with obscure pinnate venation, domatia absent. Inflorescences terminal or axillary and I/node, racemose with 1-4 distal nodes and 1-4 long- pedicellate flowers at each node or few flowered and irregular (as in ours). Flowers bisexual, distylous, glabrous externally, calyx with 5 small equal lobes; corolla funnelform, tube glabrous or with hairs at the throat within, corolla lobes 5; stamens 5, included or exserted, subsessile from near the base of the tube or between the petal lobes; ovary 2-locular, with many ovules in each locule borne on peltate placentas from the septum. Fruits capsules, lo- culicidally dehiscent; seeds many, small, black.
A genus of probably two African species, with one now introduced into pans of the Americas. These plants are distinguished from the closely related Oldenlandia by the five-parted flowers.
Pentodon pentandrus (Schumach. & Thonn.) Vatke, Oest. Bot. Zeitschr. 25: 231. 1875. Hed- yotis pentandra Schumach. & Thonn.. Kongel. Dansk. Vidensk. Selsk. Naturvidenske, Math. Am. 3: 71. 1827. Oldenlandia pentandra (Schu- mach. & Thonn.) DC., Prodr. 4: 427. 1830. H. halei Torr. & Gray, Fl. N. Amer. 2: 42. 1841. P. halei (Torr. & Gray) Gray, Syn. Fl. N. Amer. l.pt. 2: 28. 1884.
Herbs with weakly erect or decumbent stems 10-30 cm high, stems 0.8-1.8 mm thick, glabrous and slightly succulent; stipule sheath 0.5-1.5(-3) mm long, entire or with 1-3 short (0.3-2 mm) subulate lobes. Leaves with poorly defined petioles 0.5-10 mm long (in ours); leaf blades 1.5-4(-8) cm long, 0.4-2.5 cm broad, narrowly ovate-elliptic to elliptic (elliptic-lanceolate to linear-lan- ceolate), apex obtuse to acute, base obtuse to cuneate and decurrent on the petiole, drying membranaceous or thin-chartaceous (often translucent), greenish and gla- brous on both surfaces, 2° veins 3-4/side but difficult to see. Inflorescences 2-5 cm long, of (l-)3-5 irregularly arranged flowers in leaf axils or at apex (elongate race- mose inflorescences not seen in North American mate- rial), peduncles 0-8 mm long, pedicels 2-5 mm long. Flowers glabrous externally, hypanthium 2-3 mm long, calyx tube 0.3-1 mm long, calyx lobes 0.5-1 .5 mm long, acute; corolla white in ours, tube 1.5-4 mm long, 2-3 mm diam., corolla lobes 5, 1-3 mm long, 0.7-2 mm broad, ovate-triangular. Fruits 2.5-4 mm long, 2-3 mm diam., obconic to oblong; seeds 0.3 mm long.
Plants of moist depressions in open sunny sites, often along streams. Collected in central Nicara- gua as early as 1869 and recently found along Rio San Juan, and in Belize. Originally in tropical Af- rica, Texas to Florida, and Cuba.
Pentodon pentandrus is recognized by its deli- cate habit with small thin ovate-elliptic leaves, small few-flowered and irregularly arranged inflo- rescences, small five-parted flowers, and small capsules with many seeds. The North American material (as represented by P. halei from the southern U.S.A. and illustrated in Correll & Cor- rell, 1982, fig. 615) differs greatly from typical Af- rican collections (Verdcourt, 1976, p. 264, fig. 38). Measurements in parenthesis in the description represent African material. Nevertheless, the spe- cies varies greatly in Africa, and the North Amer- ican material probably represents an early atypical introduction (Verdcourt, 1976, p. 263). Compare Sipanea biflora.
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Pogonopus Klotzsch
Trees or shrubs; stipules interpetiolar, free, small, del- toid to narrow, deciduous with the leaves. Leaves op- posite, petiolate; leaf blades usually thin, entire, pin- nately veined, domatia absent. Inflorescences terminal, paniculate and subcorymbose-cymose, pedunculate, bracteolate, 1 or 2 flowers of the inflorescence usually with a greatly enlarged sepal lobe forming a broad pet- iolate leaf-like and colorful "petal." Flowers bisexual, radially symmetrical (except for those with a single great- ly enlarged sepal lobe or where the tube is curved), hy- panthium turbinate, calyx tube short, calyx lobes 5, den- tate and deciduous (each inflorescence usually with 1 or 2 flowers with a greatly expanded calyx lobe); corolla cylindrical, straight or curved, barbate in the throat, co- rolla lobes 5, short, valvate in bud; stamens 5, inserted near the base of the corolla tube, filaments slender and glabrous, anthers versatile dorsifixed above the middle, exserted; ovary 2-locular, ovules many on placentas lon- gitudinally adnate to the placenta, style slender with 2 linear or oblong branches. Fruits capsular, subglobose to oblong-ovoid, 2-locular, areolate at apex, loculicidally bivalvate; seeds many, horizontal, crowded.
A neotropical genus of three species; one species ranges from Mexico to South America and is grown as an ornamental. The greatly enlarged and bright- ly colored laminar sepal lobe of one or two flowers in each inflorescence is distinctive (but compare Mussaenda erythrophylld).
Pogonopus speciosus (Jacq.) Schum. in Mart. Fl. Bras. 6, pt. 6: 265. 1889. Macrocnemum spe- ciousum Jacq., Hoit. Schoenbr. 1: 19,1.43. 1797. Macrocnemum exsertum Oersted, Vidensk. Meddel. Kjobenhavn 45. 1852. P. exsertusOer- sted, Amer. Centr. 17. 1863. Figure 16.
Shrubs or small trees, 2-6(-l 2) m tall, leafy stems 1.5- 5 mm thick, minutely (0.2-0.4 mm) appressed-pubes- cent and glabrescent in age, the older stems with con- spicuous rounded to linear whitish lenticels 0.5-1 (-2. 3) mm long; stipules 1-3.5 mm long, 3-4 mm broad and with a rounded or cuspidate tip to 1 m long. Leaves with petioles 2-15 mm long, 1-2 mm broad, appressed-pu- bescent, often with lateral wings continuous with the lamina margins; leaf blades 7-18(-25) cm long, 3- 7.5(-9.5) cm broad, elliptic-obovate or elliptic-oblong (smaller leaves near the inflorescence usually ovate), apex acuminate with tip 5-15 mm long, base tapering grad- ually and cuneate-acute, decurrent on the petiole, drying thin-chartaceous, glabrous above, sparsely pubescent on the veins beneath with thin whitish hairs 0.2-0.4 mm long, 2° veins 4-10/side, the minor venation forming a small reticulum on both surfaces. Inflorescences termi- nal or axillary, 4-18 cm long, peduncles to 5 cm long, main axis usually with opposite branches, minutely ap- pressed-pubescent, flowers usually in compact distal cy- mose gruoups, bracteoles 2-4 mm long, lanceolate to linear, pedicels 0-3(-5) mm long. Flowers with hypan-
thium 3 mm long, 2-3 mm diam., conical-turbinate, sparsely and minutely puberulent, drying dark, calyx lobes 0.6-2 mm long, 0.2-0.3 mm broad, expanded calyx lobe with a petiole 8-20 mm long and lamina 2-4(-6) cm long and 1.5-3.5(-5) cm broad, broadly ovate or ovate- orbicular, bluntly obtuse to rounded, dark red to bright red or rose red, palmately veined; corolla tubular, rose red to dark red, tube 1 2-28 mm long, straight or curved, 3-6 mm diam. distally, densely and minutely pubescent with ascending hairs, corolla lobes 3-5 mm long, 2-3 mm broad at the base; stamens with filaments 20-23 mm long, anthers ca. 2.5 mm long; style 30-35 mm long, stigmas 2-2.5 mm long. Fruits 6-9 mm long, 5-7 mm diam., oblong-urceolate, minutely (0. 1 mm) puberulent, with 2 longitudinal sulci, abruptly truncated at apex, the linear calyx lobes often persisting.
Ornamental plants grown in both evergreen and deciduous environments in Central America, from near sea level to about 500 m elevation. Probably flowering principally in the rainy season: August- December. The species grows wild in southern Mexico, the Guanacaste lowlands, and central Panama and in Colombia and Venezuela.
Pogonopus speciosus is recognized by the bright red or rose petaloid expansions of the calyx in a few flowers of each inflorescence. This species is not known from the wild in Nicaragua, El Sal- vador, or Honduras; it is possible that it was brought to Mexico as an ornamental in pre-Co- lumbian times. Compare Mussaenda erythrophyl- la, another ornamental with bright red petaloid sepal lobes.
Posoqueria Aublet
Trees, treelets, or shrubs, branches soon becoming terete, glabrous or pubescent, usually thick; stipules in- terpetiolar, free, large, triangular, usually early decidu- ous. Leaves opposite and decussate, petiolate; leaf blades usually large, entire, often coriaceous, usually with domatia. Inflorescences terminal, usually few-flowered and short (but the flowers very long), corymbose to cy- mose or umbellate, pedunculate, bracts absent or min- ute, flowers pedicellate. Flowers bisexual and mono- morphic, (4-)5(-6)-parted, radially symmetrical or bilaterally symmetrical because of the curved corolla tube, hypanthium little differentiated from the pedicel, calyx tube short, calyx lobes short-dentate, usually persisting, often auriculate at the base, with glands within; corolla long-salverform or long-tubular, white, corolla tube long and narrow, glabrous externally, glabrous or glandular- papillate within, corolla lobes imbricate or contorted in bud, rotate or reflexed; stamens unequal or subequal, short and inserted on the mouth of the tube, glabrous or pilose, anthers basifixed and sagitate, linear-oblong, ex- serted, (the pollen is said to be released explosively in some spp.); ovary 2-locular or incompletely 1-locular, ovules many in each locule and erect on stipitate bila- mellar placentas borne on the septum, style filiform, as
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FIELDIANA: BOTANY
long as the tube, stigmas short and bind. Fruits baccate or drying hard and indehiscent. globose to ovoid, 1 - or 2-locular, often large; seeds large, hard, rounded and obtusely angled to flattened, imbedded in a gelatinous pulp.
A Neotropical genus of 1 2- 1 6 species, with most of the species in South America; a single species ranges as far north as southern Mexico. The genus is distinctive because of the larger stiff leaves, large stipules, the long white flowers, and the globose
indehiscent fruit with large seeds imbedded in a fleshy pulp. The corolla tubes are usually more than 10 cm long and less than 1 cm diam. The flowers are strongly fragrant at night when they open but yellowish and odorless by morning; these are probably pollinated by long-tongued moths. In some species the stamens are held asymmetrically and under tension over the mouth of the corolla tube until they are disturbed, when they snap for- ward. Compare Borojoa and Tocoyena.
Key to the Species of Posoqueria
la. Corolla tubes (20-)30-38 cm long; leaf blades (10-) 18-46 cm long leaf blades usually puberulent with very thin hairs beneath (but difficult to see, except along the larger veins), usually slightly rough and minor venation obscure beneath (uncommon) P. grandiflora
Ib. Corolla tubes 8-20 cm long; leaf blades 10-20(-24) cm long, essentially glabrous beneath .... 2a
2a. Leaf blades drying dull above with the tertiary veins not easily seen, the minor venation not raised;
fruit dull yellow to orange, ellipsoid to ovoid, with pericarp leathery and 4—10 mm thick
P. coriacea
2b. Leaf blades lustrous above with tertiary veins easily seen and sometimes slightly elevated; fruit orange and globose, with pericarp hard and brittle, 1-3 mm thick P. latifolia
Posoqueria coriacea M. Martens & Galeotti, Bull. Acad. Brux. 11: 240. 1844.
Small trees or shrubs, 3-8(-20) m tall, branchlets 2- 7 mm thick, glabrous; stipules 8-12(-28) mm long, ovate, ovate-oblong to ovate-lanceolate or suborbicular, acute, glabrous. Leaves with petioles 6-22 mm long, glabrous; leaf blades (6-)10-22(-27) cm long, (3-)5-12(-18) cm broad, narrowly elliptic to ovate or narrowly obovate, apex obtuse to abruptly acute, base cuneate to rounded and subtruncate, drying coriaceous, 2° veins 4-8/side. Inflorescences 2-4 cm long and 2-3 cm broad (not in- cluding the corolla lengths), cymose-corymbose, with 7- 25 flowers, peduncles 1-2 cm long, glabrous or minutely puberulent, pedicels 5-10(-23) mm long, glabrate. Flow- ers with hypanthium + calyx 6-8 mm long, subulate glands present within the calyx, calyx lobes 5, unequal, 0.5-1.2 mm long, rounded; corolla greenish white, gla- brous externally, long-tubular, tube 9-20 cm long, 1.5- 3 mm diam. in the lower half, pilose within the throat and at the base of the lobes, corolla lobes 5, 15-25 mm long, unequal, rounded distally; anthers 7-10 mm long. Fruits 7-10 cm diam., ellipsoid to ovoid, brown, peri- carp leathery, 4-10 mm thick; seeds 10-12 mm diam., angular, aril white.
Plants of lowland rain forest formations, from near sea level to 1 100 m elevation (to 1500 m in South America). Flowering January-March; fruit- ing in March-April, June-August, and October- November. The species ranges from Mexico to Brazil.
Posoqueria coriacea is characterized by the long-
tubular flowers, dull leaf surfaces, and fruit with thick leathery rind. The 2° veins are often some- what impressed in thick mature leaves when dried. The larger leaf dimensions noted above in paren- theses are from Steyermark's Flora of Venezuela treatment ( 1 974) and may not occur in Costa Rica. The breeding system of this species was studied by Bawa and Beach (1983).
Posoqueria grandiflora Standl., J. Wash. Acad. Sci. 18: 166. 1928. P. maxima Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 57. 1930. Figure 15.
Small trees, treelets, or shrubs, 2-7(-17) m tall, trunk to 18 cm dbh, leafy stems 2.5-12 mm thick, glabrous or minutely puberulent with thin erect hairs 0.1-0.2 mm long; stipules 8-18 mm long, 4-1 1 mm broad, triangular, glabrescent, deciduous. Leaves well spaced along the stem, petioles 7-20 mm long, 2-4 mm thick, glabrous or mi- nutely puberulent; leaf blades ( 1 3-)l 8-34(-46) cm long, 6-1 7(-23) cm broad, elliptic to elliptic-obovate or ellip- tic-oblong, apex acuminate to short-acuminate or blunt- ly rounded, base tapering gradually to slightly rounded and obtuse or acute, drying subcoriaceous to coriaceous, pale grayish green beneath, glabrous above, with thin erect hairs 0.1-0.5 mm beneath or the hairs difficult to see and apparently glabrous, slightly rough to the touch beneath, 2° veins 4-8/side and prominent on both sur- faces, minor venation obscure on both surfaces. Inflo- rescences only 3-6 cm long (not including the corolla tubes), corymbose with ca. 10-20 flowers, peduncles 1-
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2 cm long and 3-8 mm thick, pedicels 2-12 mm long, ca. 2 mm thick, glabrous or minutely puberulent. Flow- ers glabrous, hypanthium 3-6 mm long, little differen- tiated from the pedicel, calyx tube 3-6 mm long, 3-6 mm diam., calyx lobes 5, 0.5-1.5 mm long, broadly rounded distally; corolla long-tubular with rotate lobes, white, tube ( 1 2.5-) 1 5-36 cm long 2.5-4 mm diam., lobes 5, 20-38 mm long, 7-1 3 mm broad, oblong-obovate and rounded; stamens with unequal filaments to 1 5 mm long, 0.5-1 mm thick, anthers 9-12 mm long, to 2 mm broad, linear-lanceolate. Fruits 7-12 cm long, 4-7 cm diam., ovoid to ellipsoid, with a slightly roughened bark-like (minutely lenticellate) brownish surface, pericarp 5-12 mm thick; seeds 6-10 mm long, orange to white.
Trees and treelets of evergreen lowland Carib- bean rain forest formations, from 40 to 300 m elevation. Flowering in February, April, July, and September-October; probably fruiting throughout the year. The species ranges from northeastern Costa Rica to Colombia.
Posoqueria grandiflora is recognized by its very long flowers on small terminal inflorescences, the relatively large corolla lobes, and the large leaves with obscure minor venation and pubescence (when present), which makes the lower leaf surfaces slightly rough to the touch. This species appears to be less common than its congeners in Costa Rica. The species has been called "wild coffee" in the Caribbean lowlands. An unusual collection (Wilbur 207 '11 DUKE, F) from southern Costa Rica has large leaves with conspicuous long hairs, and flowers with very long corolla tubes, but is prob- ably no more than an extreme form of this species. The Colombian material ascribed to P. maxima appears to be conspecific; Steyermark erred in making it a subspecies of P. coriacea Mart. & Gal.
Posoqueria latifolia (Rudge) Roem. & Schult., Syst. Veg. 5: 227. 1819. Solena latifolia Rudge, PI. Guian. 1: 26, t. 40. 1806. Stannia panamensis Walp. & Duchass., Linnaea 23: 755. 1850. P. panamensis (Walp. & Duchass.) Walp., Ann. Bot. Syst. 2: 797. 1852. Figure 15.
Small trees to 9(-20) m tall, trunks to 25(-40) cm dbh, wood hard and reddish, leafy stems 2-6 mm thick, gla- brous; stipules 7-1 8 mm long, 3-8 mm broad at the base, triangular to ovate-oblong, sometimes slightly (1 mm) united above the petioles, apex obtuse to acuminate, stiff, glabrous. Leaves distant along the stem, petioles (4-) 7- 20 mm long, 1 . 5-4 mm thick, glabrous; leaf blades (7-) 1 0- 20(-24) cm long, (3-)4-10(-14) cm broad, elliptic-ob- long to elliptic-ovate or ovate, apex obtuse or short- acuminate, base abruptly narrowed and obtuse or round- ed, drying stiffly chartaceous to subcoriaceous, glabrous
above and below, 2° veins 5-7/side, the minor venation visible on both surfaces. Inflorescences 2-5 cm long (not measuring the corolla tubes), corymbose with 7-1 8 flow- ers, peduncles 1-2 cm long, 2-3 mm thick, bracteoles ca. 0.5 mm long, pedicels 3-9 mm long, glabrous. Flow- ers glabrous externally, sweet aromatic, hypanthium 3- 6 mm long, calyx lobes poorly developed, 0-0.5 mm long; corolla long-tubular with usually reflexed lobes, white, tube 8-14(-16) cm long, 2-3.3 mm diam., lobes 5 (4), 12-20(-26) mm long, 4.5-5.5 mm broad, narrowly oblong, rounded; stamens 5 (4), filaments to 6 mm long, ca. 0.4 mm thick, anthers 6-7.5 mm long, ca. 1 mm broad, linear-oblong, with basal lobes ca. 1 mm long; style ca. 9 cm long. Fruits 4-6 cm diam., globose to ovoid, yellow or orange at maturity, the pericarp only 1-3 mm thick, surface becoming wrinkled; seeds 6-12 mm long, often triangular, translucent in life, arils white to yellow-orange, in a fleshy sweet pulp.
Trees and treelets found in rain forests, partly deciduous forests, and moist sites in deciduous forests, from 2 to 700(-1200) m elevation. Prob- ably flowering and fruiting throughout the year (but mostly flowering in March-October and fruit- ing in October-April). This common species oc- curs in all the lowland evergreen areas of Costa Rica. The species ranges from southern Mexico to the Amazon basin of Brazil and Bolivia.
Posoqueria latifolia is recognized by the larger stiff glabrous leaves, the small terminal inflores- cences with very long tubular flowers, and the glo- bose fruit. The barely visible minor (3° and 4°) venation appears to be a consistent way of differ- entiating the leaves of P. latifolia from those of P. coriacea. Croat (1978, p. 814) remarks that the anthers are united at anthesis along one side of the tube and burst apart when contacted. He also notes that the flowers open late in the day and do not persist on the following day. Common names re- corded in Costa Rica are boca de vieja, carica,fruta de mono, guayaba de mono, guayaba mica, man- zana de mico, picarito, and querica. The names jicarillo and querica are used in southeastern Nic- aragua. The fruit's pulp is edible and sweet.
Psychotria Linnaeus Nomen conservandum
REFERENCES— C. Hamilton, A revision of Me- soamerican Psychotria subgenus Psychotria (Ru- biaceae), part I: Introduction and species 1-16. Ann. Missouri Bot. Gard. 76: 67-1 1 1 . Part II: Spe- cies 17-47. Ann. Missouri Bot. Gard. 76: 386- 429. Part III: Species 48-6 1 and appendices. Ann. Missouri Bot. Gard. 76: 886-916. 1989. A. Mo-
220
FIELDIANA: BOTANY
lina, Revision de las Especies de Cephaelis en Mexico, Centre America y las Antilllas. Ceiba 4: 1-38. 1953. (Other references are listed under spe- cific species below, under the family description, and at the end of the text.)
Shrubs or small treelets, less often medium-sized trees or herbaceous subshrubs (rarely lianas), terrestrial or rarely epiphytic, stems often slightly thickened at the nodes, glabrous or puberulent; stipules usually united and in- terpetiolar with 1 or 2(-5) apices or lobes on each side, sometimes also united around the stem above the pet- ioles (intrapetiolar) and forming a short tube or sheath, rarely separate to the base and appearing free, deciduous or persisting, colleters often present on the stems at the adaxial base of the stipules (and drying reddish in sub- genus Psychotria). Leaves opposite and decussate (rarely 3 or 4/node), petiolate or rarely subsessile, usually acu- minate at the apex; leaf blades entire and pinnately veined, raphides (cystoliths) present and obscure or conspicuous, some species with domatia in the vein axils beneath (some African species with bacterial nodules in the leaves). Inflorescences mostly terminal and occasionally becom- ing pseudoaxillary by further growth of an axillary branch, less often consistently axillary, usually solitary at a node, usually pedunculate and paniculate with opposite branching, bracts large to small or rarely undeveloped, bracts forming an involucre in some species formerly placed in Cephaelis, flowers often borne in distal brac- teolate cymes, sessile or pedicellate. Flowers bisexual (in Central America), radially symmetrical, usually small, often distylous with long-styled (pin) and short-styled (thrum) forms within the same species, calyx tube usually short and cupulate, with 4 or 5(-6) short calyx lobes or without lobes and entire; corolla tubular to funnelform or campanulate, white to pink or yellowish, corolla tube usually short or narrow, often with tufted hairs at the throat within, glabrous or puberulent externally, corolla lobes 4-5(-6), always valvate in bud, often thickened at the tips; stamens 4-5(-6), borne from the middle or up- per part of the tube, filaments slender, anthers narrow, included or exserted, disc ring-shaped and encircling the base of the style; ovary usually 2-locular (rarely with 4- 6 locules), 1 erect ovule borne from the base of each locule or from the base of the thick septum, style long or short, with 2 (rarely 4-6) linear stigmas. Fruits fleshy drupes, red, yellow, blue, purple, black, or white when ripe, often with spongy arenchymatous tissue, usually with 2 (4-6) hard pyrenes, pyrenes plano-convex with a flattened interior (adaxial) face and a rounded exterior (abaxial) surface, often with longitudinal ridges on the exterior surface (rarely with transverse ribs or projec- tions), usually with a median longitudinal sulcus on the inner face.
Psychotria is the largest genus of Rubiaceae and one of the largest genera of Angiosperms, with an estimated 1,500-1,600 species in the tropics and subtropics of both hemispheres. Together with Piper (Piperaceae) and Miconia (Melastomaceae),
Psychotria is one of Costa Rica's three most spe- ciose woody genera, especially common in the un- derstory of evergreen forests and forest edges. Some species are difficult to separate from similar spe- cies in Coussarea, Faramea. and Palicourea (q.v.). While Cephaelis appeared to be a very distinctive genus in Central America, a great number of in- termediate species have necessitated the inclusion of its species in Psychotria. Our treatment has ben- efited greatly from Hammers work on Psychotria at La Selva (in Taylor, 1991). Likewise, Hamil- ton's recent revision (see references above) of the species in subgenus Psychotria has been especially helpful, as these species are often difficult to dif- ferentiate; we have not deviated from his species concepts in this treatment.
Psychotria is generally characterized by the sol- itary terminal inflorescences with opposite branching (in most species), the 1 - or 2-lobed in- terpetiolar stipules, entire and pinnately veined opposite leaves, smaller flowers with minute calyx lobes, short corolla tubes, valvate corolla lobes, basal solitary ovules, and fleshy fruit usually with 2 hard seed-like pyrenes. The short shrubby habit, preferences for areas of high rainfall, fruit often with longitudinal ridges, and white or yellowish flowers are additional characteristics. Those spe- cies with axillary inflorescences tend to have un- branched succulent stems or few-branched woody stems (see below). All the fruits appear to be bird- dispersed; they are fleshy and bright red, blue to purplish and black, or arenchymatous and whitish. Two species of Psychotria found in Costa Rica, P. emetica and P. ipecacuanha, are used medicinally.
Most species of Psychotria in Costa Rica are readily recognizable but the differences between closely related species can be subtle. There appear to be real problems of intcrgradation in the epi- phytic species, and the succulent-stemmed her- baceous species with axillary inflorescences pre- sent problems of species delimitation. Except for the groups just mentioned, the majority of species stand well apart and give little evidence of hy- bridization or intergradation. The difficulty in identifying a specimen to species often is due to the large number of species in the genus, and the wide range of variation found within some species. Below we provide keys to four groups within the genus; groups 2 and 3 appear to be largely mono- phyietic. The illustrations are an additional ave- nue for identification and are grouped by inflo- rescence position, subgeneric placement, and general similarity.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
221
Key to Four Species Groups of Psychotria in Costa Rica
1 a. Plants nearly always epiphytic and with subcoriaceous leaves drying grayish; ovary usually with 4 locules; fruits red and often with 4 seeds; or plants of Cocos Island Group 1
Ib. Plants not epiphytic, leaves various; ovary with 2 (rarely 5) locules; fruit usually with 2 (rarely 5) seeds; and plants not known from Cocos Island 3
2a. Inflorescences regularly axillary; main stems usually short (ca. 1 m) and succulent in a majority of species, both succulent and woody stems often with hollow sections within when dried . . . Group 2
2b. Inflorescences terminal, only occasionally axillary where lateral shoots have continued to develop; main stems usually woody and rarely with hollow sections when dried 3
3a. Leaves drying greenish, gray-green, yellow-green, yellowish brown, or brown (note that leaves treated with isopropyl alcohol may turn reddish brown and that some species have pinkish venation in life), domatia rarely present; stipules often persisting, the stipules not subtending and enclosing a ring of reddish hairs (colleters) in early stages; fruit usually becoming blue, purple, or black (red only in P. haematocarpa with very small inflorescences, or orange in later stages in P. racemosa with 5 locules and 5 -seeded fruit); subgenus Heteropsychotria Group 3
3b. Leaves drying grayish to black, pinkish brown or dark reddish brown, domatia present in a few species; stipules usually falling off as the leaves expand (caducous), and usually enclosing a short ring of reddish hairs (these often persisting just above the stipule scar); fruit always red at maturity (never with more than 2 seeds); subgenus Psychotria Group 4
Key to Group 1: Epiphytic Species of Psychotria in Costa Rica and Three Species of Cocos Island
The epiphytic species appear to represent a very natural (probably monophyletic) group. However, there is the strong likelihood that a large number of our collections belong to a single polymorphic entity, for which P. guadalupensis is the earliest name. Psychotria maxonii appears to be quite distinct, but there are a significant percentage of collections that appear to be intermediate between P. guadalupensis and P. pithecobia. See the discussion under P. guadalupensis.
la. Epiphytic plants of continental Costa Rica 2
1 b. Trees and shrubs of Cocos Island 4
2a. Leaves to 5(-7) mm wide, 10-28 mm long, narrowly lanceolate to narrowly oblong; inflores- cences to 3 cm long P. maxonii
2b. Leaves 8-40 mm wide, 20-120 mm long, more than 40 mm long if lanceolate or narrowly
oblong; inflorescences 3-14 cm long 3
3a. Leaves with 6 or more pairs of secondary veins, the veins clearly elevated and visible on the dried leaf surfaces, leaves more than 4 cm long and 2 cm broad; inflorescences with peduncles
2-7 cm long P. pithecobia
3b. Leaves with fewer than 6 pairs of major secondary veins, the veins often obscure on the surfaces
of the dried leaves, leaves 3-10 cm long and 1-5 cm broad, very variable (on different plants)
as regards size, shape, and texture; peduncles 0.5-3 (rarely to 5) cm long . . .P. guadalupensis
4a. Leaves to 25 cm long, drying subcoriaceous and dark reddish brown; stipules forming an acute
F/CMs-like cap over the shoot apex and early deciduous P. cocosensis
4b. Leaves to 1 5 cm long, drying thin-chartaceous and greenish or grayish; stipules with 2 acute lobes
on each side and persisting 5a
5a. Inflorescences less than 3 cm long, with thick (1 mm) lateral branches less than 5 mm long; stipule
lobes 2—4 mm long P. brachybotrya
5b. Inflorescences usually becoming more than 3 cm long, with slender (0.3 mm) lateral branches 5- 10 mm long; stipule lobes ca. 1 mm long Psychotria sp. A
Key to Group 2: Species of Psychotria with Axillary Inflorescences
la. Inflorescences axillary to both leaves of the node, usually with 2 or more inflorescences at distal nodes, inflorescences usually less than 5 cm long; fruits becoming blue, black, or purple 2
222 FIELDIANA: BOTANY
Ib. Inflorescences axillary to only 1 leaf at each node, usually with 1 inflorescence at distal nodes,
inflorescences often more than 5 cm long; fruits white to yellow or red .8
2a. Inflorescences subsessile and forming a dense verticillate cluster at the node 3
2b. Inflorescences short-pedunculate, not forming dense verticillate clusters around the node
5
3a. Inflorescences lacking an involucre of bracts, calyx lobes to 3 mm long (not Psychoiria
spp.) Hoffmannia spp.
3b. Inflorescences subtended by an involucre of broad bracts, calyx lobes to 2 mm long
4
4a. Bracts subtending the inflorescences pale green; stipules to 10 mm long and 5 mm
broad; widely distributed at 1 200-2800 m elevation P. aubletiana
4b. Bracts reddish; stipules 12-22 mm long and almost as broad; 700-1500 m near Orosi
and Muneco P. cartagoensis
5a. Leaves drying greenish to greenish brown, shrubs or small trees with many branches, young stems glabrous; stipules forming a short sheath; inflorescences with many branches, often
with more than 1 0 fruits P. cooperi
5b. Leaves drying grayish or dark brown, subshrubs or few-branched shrubs, young stems mi- nutely puberulent; stipules not forming a sheath; inflorescences few-branched, rarely with
more than 5 fruits 6
6a. Inflorescences usually 4/node; plants growing to 2(-5) m tall, with lateral branches; leaf blades
drying stiffly chartaceous [often obtuse at the base; peduncles to 1 5 mm long] . . . P. erecta
6b. Inflorescences 2/node; plants growing to 1 m tall, unbranched; leaf blades drying thin-char-
taceous 7
7a. Leaf blades drying brownish, often cuneate basally; peduncle to 5 mm long, inflorescence
less than 2.5 cm long P. emetica
7b. Leaf blades drying greenish, often decurrent basally; peduncle to 70 mm long, inflorescences
more than 3 cm long P. aggregata
8a. (from Ib) Stipules lobed or fimbriate distally, to 3 cm long, translucent to opaque 9
8b. Stipules entire distally, bluntly obtuse to rounded, to 1 cm long, thick and opaque 11
9a. Stipules to 20 mm broad, drying yellowish and translucent, fimbriate to bluntly lobed [in- florescences neither capitate nor hirsute; fruit becoming white] P. cartagoensis
9b. Stipules to 7 mm broad, drying dark and opaque, with stiff narrow or filiform teeth; fruit
becoming red, purple, or blue 10
lOa. Leaves pilose with hairs to 2 mm long; inflorescences to 15 cm long, much-branched and
lacking an involucre; fruit becoming purple or blue P. pilosa
lOb. Leaves glabrous or with short (0.5 mm) hairs; inflorescences to 5 cm long, capitate and
involucrate; fruit red P- ipecacuanha
1 la. Leaves with more than 18 pairs of closely parallel secondary veins, veins becoming prominently raised and the leaf corrugated in age, bluntly obtuse to rounded distally; plants rarely exceeding
40 cm in height [fruits red] - P- polyphlebia
1 Ib. Leaves with fewer than 18 pairs of secondary veins, not becoming corrugated, usually acuminate
at apex; plants usually more than 50 cm tall
1 2a. Fruits becoming orange or red
12b. Fruits becoming white or yellowish green
13a. Leaf blades drying dark above and much paler beneath, young leaves glabrous; pyrenes with
a longitudinal costa on the back; commonly collected . P. uliginosa
13b. Leaf blades drying greenish to dark brown above and only slightly paler beneath; young leaves densely hirsute beneath with hairs to 1 mm long; pyrenes lacking dorsal costae; rare
P. siggersiana
14a. Young leaves villose beneath with hairs to 1 mm long, a definite arcuate submarginal vein present 2-3 mm from the leaf edge, with 16-22 pairs of major secondary veins; fruit becoming greenish
yellow P capacifolia
14b. Young leaves minutely puberulent or glabrous, a definite submarginal vein absent, with 10-16 pairs of secondary veins (and often loop-connected in the distal part of the leaf); fruit becoming white and spongy. (Note: This is a variable group of plants that may intergrade.)
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 223
1 5a. Inflorescences with thin widely spreading distal branches, bracts subtending the major branches less than 4 mm long, flowers borne separately along the distal branches P. macrophylla
15b. Inflorescences with thick (1 mm) distal branches or the inflorescences compact and short (5 cm), bracts subtending the basal branches or basal flowers more than 4 mm long 16
1 6a. Peduncles 0-3 cm long, flowers loosely clustered to capitate; leaves usually elliptic (rarely consis- tently narrow); widespread in Costa Rica P. aggregata
1 6b. Peduncles to 1 5 cm long, flowers usually capitate on the peduncle or on 3 short branches on the
peduncle; leaves usually elliptic-lanceolate; eastern part of the Cordillera de Talamanca
P. aggregata (sensu stricto)
Key to Group 3: Species of Subgenus Heteropsychotria and Similar Species in Costa Rica (Including Species Formerly in Cephaelis)
la. Flowers borne in dense capitate or subcapitate inflorescences with closely clustered flowers, bracts usually conspicuous and enclosing the flowers; inflorescences terminal and solitary or 3 closely
grouped together (species formerly placed in Cephaelis and others) 2
Ib. Flowers in open or congested inflorescences but not capitate or subcapitate (flowers sometimes in small distal capitula or glomerules on secondary branches within branched inflorescences); bracts
small to conspicuous or absent, bracts rarely enclosing and obscuring the flowers 16
2a. Inflorescences subtended by 2(-3) large bracts forming a basal cupulate involucre, capitulum
solitary, 3-10 cm broad 3
2b. Inflorescences usually with more than 4 basal bracts, lacking a single cupulate involucre at
the apex of the peduncle; inflorescence of 1 or 3 capitula, 1-5 cm broad 6
3a. Bracts reddish (rarely yellow), inflorescences erect; very common shrubs in lowland
evergreen formations 4
3b. Bracts purple to lilac, inflorescences erect or pendant; rarely collected species 5
4a. Stems and leaves glabrous; stipules with 2 rounded lobes on each side . . P. elata 4b. Stems and leaves densely tomentulose; stipules with 2 sharp teeth on each side
P. poeppegiana
5a. Montane (800-1200 m) plants; bracts purple, inflorescences pendant; leaf blades with
ca. 1 4 pairs of 2° veins P. correae
5b. Lowland (0-500 m) plants; bracts lilac, inflorescences erect; leaf blades with ca. 7 pairs
of 2° veins P. borucana
6a. Stipules with more than 2 slender teeth on each side (more than 6 teeth per node); unbranched subshrubs less than 1 m tall; fruit red [medicinal plants rarely collected in Costa Rica] . . .
P. ipecacuanha
6b. Stipules with 2 slender teeth on each side, or with 2 rounded lobes or unlobed (with 4 or fewer stipule lobes per node); plants mostly shrubby and more than 1 m tall (except P.
guapilensis); fruit blue or purple 7
7a. Corollas more than 4 cm long; fruit becoming more than 1 2 mm long; bracts usually bluntly
obtuse and green P. chiapensis
7b. Corollas less than 3 cm long; fruit less than 1 2 mm long; bracts acute to rounded and green
to purple 8
8a. Basal bracts of the inflorescences rounded to bluntly obtuse at the apex 9
8b. Basal bracts acute to acuminate 14
9a. Bracts white to pale green or bluish, glabrous, broadly rounded distally forming a tight
cupulate involucre beneath the congested capitulum [0-600 m elevation]
P. glomerulata
9b. Bracts green to deep purple, glabrous to puberulent, bluntly obtuse distally (rounded
in P. platypoda), not forming a definite cup at the base of the inflorescence 10
1 Oa. Bracts deep purple or reddish purple, inflorescences often densely compacted and spher- ical; plants usually less than 1 m tall P. guapilensis
1 Ob. Bracts greenish to purple, inflorescences never spherical, often loosely compacted; plants usually more than 1 m tall 11
224 FIELDIANA: BOTANY
1 la. Inflorescences usually of 3 pedunculate capitula; bracts white to pink (purple in fruit); stipular tube 2-5 mm long; 2° veins 8-12/side [1000-2500 m elevation] ...P. dichroa
lib. Inflorescences usually of a single pedunculate capitulum; bracts whitish to green or purple; stipular tube ca. 1 mm long; 2° veins 1 1-22/side .12
12a. Bracts broadly ovate and rounded at the base, purple; stipule lobes short and bluntly rounded; 1200-2300 m elevation p, molinianum
1 2b. Bracts usually narrowed at the base, whitish to green or purple; stipules lobes short to long, acute; 0-800 m elevation 13
13a. Bracts elliptic-obovate to oblanceolate; flowers usually puberulent, corolla tube 9-13 mm long; commonly collected p. suerrensis
1 3b. Bracts broadly rounded; flowers glabrous, corolla tube 3-6 mm long; rare in Costa Rica
P. platypodd
14a. (from 8b) Bracts orange or reddish orange, ovate and broadly overlapping; plants confined
to the southern Pacific slope, 600-1300 m elevation P. aura nti bract ea
1 4b. Bracts green or greenish with white, blue, or purple, linear lanceolate to ovate but not broadly
overlapping; plants wide ranging 15
1 5a. Bracts narrowly ovate, often marked with blue; inflorescences ca. 3 cm long and 2 cm broad
with erect bracts; stipule lobes thin and translucent, to 6 mm long; Caribbean slope, 900-
1 600 m elevation P. hazenii
15b. Bracts often linear-lanceolate, usually marked with white or purple; inflorescences short (1
cm) with broadly spreading bracts; stipule lobes stiff and opaque, to 4 mm long; widely
ranging in moist evergreen formations, 0-800 m elevation P. hoffmannseggiana
1 6a. (from 1 b) Young stems densely and conspicuously puberulent with hairs 0.3-2 mm long; peduncles
and branches of the inflorescence usually densely puberulent 17
16b. Young stems glabrous or sparsely puberulent with inconspicuous hairs less than 0.3 mm long;
peduncles and branches of the inflorescences glabrous or puberulent 25
17a. Hairs usually becoming more than 0.8 mm long, leaf blades drying dark above and 12-30
cm long, often with more than 14 pairs of major 2° veins; flowers in dense distal clusters,
subtended by conspicuous bracts 18
17b. Hairs rarely exceeding 0.8 mm in length; leaf blades usually drying greenish (rarely dark
brownish above), usually less than 20 cm long, with less than 14 pairs of major 2° veins;
flowers in open or small distal clusters, bracts various 19
1 8a. Leaf blades with 9-18 pairs of major 2° veins; flowers subtended by densely puberulent lanceolate bracts ca. 2 mm broad; wide-ranging P. pilosa
18b. Leaf blades with (14-) 17-23 pairs of major 2° veins; flowers subtended by sparsely
puberulent ovate bracts 2-3 mm broad; southwestern Costa Rica .... P. mortoniana
19a. Ovary with 5 locules, fruits often with 5 seeds; stipules with 2 long (6-14 mm) stiff persisting
awns on each side; leaf blades with conspicuous subparallel 3° veins [9-21 (-26) cm long]
P. racemosa
19b. Ovary with 2 locules, fruits never with more than 2 seeds; stipules rarely with 2 long stiff
awns (P. umbelliformis); leaf blades rarely with conspicuous 3° veins 20
20a. Inflorescence umbelliform, flowers in 3 small glomerules on equal primary branches at the
apex of a long peduncle; leaves glabrous above [rare] P. umbelliformis
20b. Inflorescences not as above; leaves puberulent or glabrescent above .21
2 la. Leaf blades cuneate at the base and long-decurrent on the petiole, thin in texture; flowering
portion of the inflorescences often broader than long; deciduous and evergreen formations
of the Pacific slope P pubescens
21b. Leaf blades not cuneate and long-decurrent at the base (sometimes decurrent in P. steyer-
markii), mostly stiff-chartaceous when dried; flowering portion of the inflorescence rarely
broader than long; plants of evergreen formations
22a. Pubescence of young stems usually in narrow longitudinal lines; leaf blades 3-12 cm long
and 1-3 cm broad; stipules with narrow awns 2-5 mm long [plants of lower montane (800-
1 800 m) cloud forests; inflorescences few-branched and racemose] P. steyermarkii
22b. Pubescence not in narrow longitudinal lines; leaf blades 6-20 cm long, 2-5 cm broad; lines;
plants rarely collected above 800 m elevation
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 225
23a. Plants of the Osa Peninsula; inflorescences with hairs to 1.5 mm long, branches and flower clusters distant and subtended by linear bracts to 8 mm long [leaf blades 5-12 cm long and with 10-12 pairs of major 2° veins] P. acicularis
23b. Plants not known from the Osa Peninsula or from the Pacific slope below 400 m elevation; inflorescences with hairs usually less than 1 mm long, flower clusters congested or distant, subtended by linear bracts usually less than 4 mm long 24
24a. Inflorescences with crowded flowers and branches, usually less than 4 cm broad, narrowly pyramidal, erect, calyx lobes 0.3-1.5 mm long; leaf blades 7-20 cm long, with 8-14 pairs of major 2° veins; awns of the stipules to 3 mm long P. hebeclada
24b. Inflorescences usually with open branching and separate flower clusters, usually more than 3 cm broad, broadly pyramidal, often pendant; calyx lobes 0.2-0.5 mm long; leaf blades 6- 1 5 cm long and with 8-1 1 pairs of major secondary veins; awns of the stipules to 7 mm long
P. pittieri
25a. (from 16b) Inflorescences large and many-branched, usually becoming more than 12 cm long and
1 0 cm broad, broadly paniculate; leaf blades usually large (often to more than 1 8 cm long); fruit
rarely more than 4 mm diam. when dried (to 5 mm in P. solitudinum) 26
25b. Inflorescences smaller, rarely more than 12 cm long (and if so narrowly racemiform) and usually
less than 10 cm broad, paniculate to subcapitate or racemiform; leaf blades often less than 15 cm
long; fruit 3-12 mm diam. when dried 31
26a. Stipule lobes to 15 mm long and 5 mm broad at the base, conspicuous; flowers distant in small (1-2) alternate sessile groups along the slender distal branches of the inflorescence; fruits grayish blue to white P. microbotrys
26b. Stipule lobes rarely more than 5 mm long, usually less than 2 mm broad at the base; flowers distant in small sessile groups only in P. solitudinum; fruit blue to black 27
27a. Stipules at first acute distally but splitting and developing 2 acute teeth or awns separated by a broad U-shaped sinus; leaf blades often drying dark brown above; rarely collected below 400 m elevation P. berteriana
27b. Stipules at first rounded or obtuse at the apex, splitting into 2 broad lobes separated by a narrow V-shaped sinus; leaf blades drying dark greenish brown to yellowish green or grayish green; plants growing from 0 to 1 700 m elevation 28
2 8a. Flowers in small (1-2) separate groups along the (usually dichotomous) slender distal branches of the inflorescence [corolla tube 4-5 mm long, narrowed at the base and urceolate distally]; southwest Pacific slope of Costa Rica P. solitudinum
28b. Flowers in distal small cymes on multiple-branched inflorescences with opposite and cymose
branching; corolla various; plants not collected from the southwest Pacific area of Costa Rica
29
29a. Plants only known from the upper Rio Grande de Orosi and Tapanti above 1 300 m elevation; floral bracts ca. 3 mm long, often persisting with the flowers and obtuse distally; corolla tube ca. 6 mm long and tubular P. tapantiensis
29b. Plants not collected above 1200 m elevation; floral bracts to 3 mm long, early deciduous; corolla tube 2-7 mm long 30
30a. Corolla tube 2-3 mm long, funnelform or tubular; floral bracts usually rounded at the apex; leaf blades with 8-14 pairs of major 2° veins; fruit blue P. luxurians
30b. Corolla tube 6-7 mm long, narrowed at the base and expanded distally; floral bracts usually acute at the apex; leaf blades with ca. 8-10 pairs of major 2° veins; fruit becoming blue or
black P. angustiflora
3 la. (from 25b) Inflorescences 12-20 cm long and only 2-5 cm broad, racemiform or thyrse-like,
peduncles 6-12 cm long; leaf blades with a distinct marginal vein within ca. 0.3 mm of the leaf
edge and connecting the distal ends of the major secondary veins; stipules narrowly 2-lobed . . .
P. cincta
31b. Inflorescences rarely over 14 cm long, usually more than 4 cm broad when over 10 cm long, not
racemiform or narrowly thyrse-like; leaf blades lacking a distinct marginal vein along the leaf edge;
stipules various 32
32a. Inflorescences with conspicuous bracts and bracteoles to 14 mm long and 3 mm broad, the bracts
persisting with the flowers . 33
FIELDIANA: BOTANY
32b. Inflorescences without conspicuous bracts, bracts rarely more than 3 mm long or more than 1 mm
broad, bracts often deciduous before the flowers mature 37
33a. Inflorescence small and compact, less than 3 cm long but expanding in fruit; stipules with
narrow awns 2-6 mm long [0-900 m elevation] p. brachybotrya
33b. Inflorescences larger and more open, to 10 cm long; stipule lobes less than 2 mm long or
triangular (not narrowly awned) 34
34a. Stipules very shortly (1-3 mm) bilobed with a U-shaped sinus between them [lateral branches
of the inflorescences not subtended by bracts; stipules 2-5 mm long] P. officinalis
34b. Stipules bilobed distally with a short or long V-shaped sinus between the lobes 35
35a. Flowers subtended by ovate bracteoles 2-3 mm long, lateral branches of the inflorescences
subtended by adnate bracts to 8 mm long; stipules 3-7 mm long P. brachiata
35b. Flowers subtended by lanceolate to oblanceolate bracts 4-14 mm long, lateral branches of
the inflorescence usually without subtending bracts; stipules 8-20 mm long 36
36a. Flowers subtended by lanceolate bracts 4-9 mm long; inflorescences with 4 or more nodes
with lateral branches; corolla tube 3-7 mm long P. capitata
36b. Flowers subtended by oblanceolate bracts 5-14 mm long; inflorescences with 1-3 nodes with
lateral branches; corolla tube 7-10 mm long P. calochlamys
37a. (from 32b) Stipules with 1 narrow lobe (usually 2/node), the awn-like lobe acute (rarely bifid)
distally and with an inner (adaxial) tooth; 500-1600 m elevation P. valeriana
37b. Stipules with 2 lobes on each side (4/node), the lobes separate laterally and without a tooth-like
adaxial (inner) appendage, or the lobes/teeth not developed; 0-2000 m elevation 38
38a. Inflorescences less than 1.5 cm long, less than 12 mm wide; fruit becoming red at maturity ....
P. haematocarpa
38b. Inflorescences becoming more than 2 cm long and more than 2 cm broad; fruit blue to purple or
black at maturity (orange during development in P. racemosd) 39
39a. Leaf blades linear-lanceolate to narrowly elliptic-oblong, largest blades less than 3 cm broad; 1000-
2000 m elevation P. goldmanii
39b. Leaf blades rarely linear-lanceolate, larger leaves over 3 cm broad; 0-1500 m elevation 40
40a. Inflorescence branches 0.7-1.5 mm thick when dried, flowering part of the inflorescence (beyond
the peduncle) broader than long 41
40b. Inflorescence branches slender (ca. 0.4 mm thick) when dried, flowering portion of the inflorescences
usually longer than broad [fruit less than 8 mm long and 7 mm diam. when dried] 43
4 la. Corolla tubes less than 3 mm long; inflorescences consistently with 4 lateral branches at the
first node; rare in Costa Rica at 1 500 m elevation P. allenii
41b. Corolla tubes 10-15 mm long; inflorescences with 2 or 4 opposite lateral branches at the first
node; from below 1 200 m elevation in Costa Rica 42
42a. Fruit 8-12 mm long and 7-10 mm diam., without transverse projections; leaf blades stiffly
chartaceous; calyx lobes ca. 0.2 mm long P. eurycarpa
42b. Fruit 4-5 mm long and ca. 6 mm diam., with transverse projections when dried; leaf blades
thinly chartaceous; calyx lobes 0.2-1 mm long P. domingensis
43a. Leaf blades drying membranaceous or thin-chartaceous, usually cuneate at the base and decurrent
on the petiole; plants often found in deciduous forest formations .44
43b. Leaf blades usually drying chartaceous, rarely cuneate and conspicuously decurrent on the petiole;
never collected in deciduous areas
44a. Leaf blades with 3-7 pairs of major 2° veins, often obovate; corolla tube 6-10 mm long; fruit
7-10 mm long P microdon
44b. Leaf blades with 9-15 pairs of major 2° veins, elliptic to ovate; corolla tube 2.5-4 mm long;
fruit 5-6 mm long P pubescens
45a. Fruits often with 5 seeds, ovary with 5 locules; stipules with stiff sharp awns 6-14 mm long and
persisting; leaf blades with 3° veins subparallel [with 7-12 pairs of major 2° veins] P. racemosa
45b. Fruit never with more than 2 seeds, ovary with 2 locules; stipules with awns or lobes 2-8 mm
long; 38 veins not subparallel 46
46a. Stipules with lobes or teeth absent or rarely 1 mm long, corolla tube 5-8 mm long (rare in Costa
Rica) P. phanerandra
46b. Stipules with lobes or teeth 2-8 mm long; corolla tubes 0.7-5 mm long 47
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 227
47a. Stipules with narrow lobes 2-3 mm long; fruits ca. 4 mm long and 6 mm with smooth rounded surfaces (dried); leaf blades with long (1-3 cm) drip tips; inflorescences often small (to 6 cm long) P. acuminata
47b. Stipules with narrow awns 3-8 mm long; fruits ca. 3 mm long and 4 mm diam. with longitudinal ribs and transverse depressions; leaf blades with narrowed tips 1-2 cm long; inflorescences to 12 cm long P. deflexa
Key to Group 4: Species of Subgenus Psychotria and Similar Species in Costa Rica
la. Leaf blades 3-9 cm long, very rarely more than 7 cm long; small shrubs of evergreen forests ... 2 Ib. Leaf blades 5-30 cm long, the largest leaf blades usually more than 9 cm long; plants of evergreen
and deciduous habitats 6
2a. Young stems densely puberulent with short (0. 1-0.2 mm) reddish brown hairs; stipules acute at the apex and caducous; inflorescences less than 1 6 mm long [ 1 200-2200 m elevation] . .
P. parvifolia
2b. Young stems glabrous (except for a ring of reddish hairs above the stipular scar); stipules bilobed or bifurcate distally; inflorescences usually more than 20 mm long (except in P.
chagrensis) 3
3a. Inflorescences capitate and sessile, less than 1 7 mm long and subtended by large bracts to 1 cm long, corolla tube 4-8 mm long; stipules bifurcate distally with a V-shaped sinus [wet
lowlands, 0-300 m elevation] P. chagrensis
3b. Inflorescences paniculate and pedunculate usually more than 20 mm long and subtended by inconspicuous bracts, corolla tubes 2-4 mm long; stipules with 2 distal lobes separated by a
broad U-shaped sinus 4
4a. Stipule lobes glabrous or minutely (< 0.05 mm) papillate puberulent; inflorescence branches
ca. 0.4 mm thick when dried [0-500 m elevation in Nicaragua and Panama]
P. fruticetorum
4b. Stipule lobes minutely (ca. 0.1 mm) ciliolate; inflorescence branches 0.2-0.3 mm thick when
dried; plants known to occur in Costa Rica 5
5a. Leaf blades rarely more than 5 cm long; petioles 2-10 mm long (the leaves not sessile);
inflorescences not exceeding 5 cm in length [0-1800 m elevation] P. graciliflora
5b. Leaf blades (at least the larger blades) usually exceeding 1 0 cm in length, petioles 0-50 mm
long (leaves sessile in some spp.); inflorescences usually exceeding 5 cm in length 21
6a. (from Ib) Young stems conspicuously puberulent with reddish hairs 0.2-2 mm long, internodes
puberulent above the ring of reddish hairs (colleters) at the stipule scar 7
6b. Young stems glabrous or inconspicuously puberulent with minute (0.03-0. 1 mm) hairs, but a ring
of reddish hairs (colleters) often present just above the stipule scars 14
7a. Inflorescences small (1-5 cm), flowers usually congested distally; peduncles to 2 cm long . .
8
7b. Inflorescences larger (4-15 cm long), flowers separate or congested, peduncles 1.5-15 cm long
10
8a. Leaf blades with an arcuate submarginal vein; stipules with a tube 4-14 mm long and
narrow awns; fruits 5-6 mm long [0-200(-800) m elevation] P. psychotriifolia
8b. Leaf blades usually lacking an arcuate submarginal vein but the distal veins often loop- connected; stipules with a tube 0-5 mm long, usually lacking narrow awns; fruits 6-14
mm long 9
9a. Leaf blades usually elliptic-obovate or elliptic; stipules 3-1 1 mm long; fruits 6-8 mm
long; widespread at 20-800(-1200) m elevation P. nervosa
9b. Leaf blades usually oblanceolate to narrowly elliptic-oblong; stipules 8-14 mm long;
fruits unknown; Chiriqui Highlands ca. 1 200 m elevation P. boquetensis
lOa. Leaf blades with 14-33 pairs of major 2° veins, leaf blades 12-30 cm long; peduncles 4-14
cm long 11
lOb. Leaf blades with 8-13 pairs of major 2° veins, leaf blades 6-30 cm long; peduncles 1.5-8 cm long 12
228 FIELDIANA: BOTANY
1 la. Leaf blades with 14-22 pairs of major 2° veins; lacking a definite submarginal vein, 3°
veins subparallel; floral bracts ca. 3 mm long, narrowly triangular P. micrantha
1 Ib. Leaf blades with 20-34 pairs of major 2° veins, with definite submarginal veins and
subparallel 3° veins; floral bracts ca. 7 mm long and ovate P. sixaolensis
12a. Leaf blades rounded at the base and subcordate at the petiole, to 30 cm long, petioles 6-12 cm long; stipules ca. 2 cm long; rarely collected plants of the Caribbean escarpment [200-
500 m elevation] p. insignis
1 2b. Leaf blades not rounded and subcordate at the base (or if so not regularly exceeding 1 8 cm
in length), petioles usually less than 6 cm long; widespread 13
1 3a. Corolla tubes 2-2.5 mm long; fruit 4-6 mm long; peduncles 1.5-5 cm long; leaf blades usually drying reddish brown or brown, major veins not impressed on the upper surfaces; (200-)400r
1600 m elevation P. jimenezii
13b. Corolla tubes 1.5-2 mm long; fruit 5-7 mm long; peduncles 3-8 cm long; leaf blades usually drying grayish, major veins impressed on the upper leaf surfaces; 20-400 m elevation ....
P. neillii
1 4a. (from 6b) Larger leaf blades usually more than 1 0 cm broad and usually more than 20 cm long
15
14b. largest leaf blades less than 10 cm broad, rarely more than 20 cm long 21
15a. Inflorescences short (3-1 1 cm long) and compact (2-5 cm broad), with lateral branches to 1
cm long and often difficult to see; plants 0.5-2 m tall 16
1 5b. Inflorescences to 30 cm long and 4-15 cm broad, with lateral branches more than 1 cm long
and clearly visible; plants 1-10 m tall 18
16a. Leaf blades with 16-19 pairs of major 2° veins, usually more than 25 cm long, rounded
to obtuse at the apex; bracts ca. 5 mm long P. chitarriana
16b. Leaf blades with 8-14 pairs of major 2° veins, usually less than 25 cm long, short- acuminate to acute at the apex; bracts 0.2-2 mm long 17
17a. Stipules with a basal tubular sheath and 2 narrow distal lobes; leaf blades with 10-14 major 2° veins on each side, usually obovate; inflorescences to 7 cm long, compact
panicles of cymes : P. alfaroana
1 7b. Stipules not tubular at base, broadly ovate and bluntly obtuse at the apex; leaf blades with 8-1 1 pairs of major 2° veins, usually broadly elliptic; inflorescences to 3 cm long,
with short lateral branches and appearing subumbellate P. lamarinensis
18a. Stipules united over the shoot apex (as in Ficus) and tearing off as the leaves begin to grow,
narrowly conical in early stages; fruit 5-9 mm long (var. compressicaulis of)
P. panamensis
18b. Stipules united only at their base, open distally and often with the base persisting, ovate- triangular in form; fruit 4-6 mm long
19a. Leaf blades subsessile and subcordate at base, usually 3-4 times longer than wide, 2° veins
arising from the mid vein at angles of 80-100° P. rosulatifolia
19b. Leaf blades with prominent petioles, never rounded at the base, blades 2-3 times longer than
wide, 2° veins usually arising at angles of 60-70° .20
20a. Secondary veins not usually loop-connected; stipules 8-30 mm long and acute to acuminate
at the apex; inflorescences to 30 cm long .P. grandis
20b. Secondary veins loop-connected distally to form an arcuate submarginal vein; stipules 5-12
mm long and rounded to acute; inflorescences to 10 cm long . P. limonensis
2 la. (from 5b and 14b) Flowers in dense sessile verticils on a single elongated (4-9 cm) rachis or in
dense clusters in a compact often globose arrangement, inflorescences solitary and apical, spiciform
or capitate/subcapitate, primary branches short (< 1.5 cm) or absent
21b. Flowers usually in open cymes, or on open spreading branches of the inflorescences when closely clustered, never spiciform or globose-capitate, primary branches of inflorescences usually more
than 1 cm long
22a. Flowers in dense sessile verticillate clusters along the slender elongated (3-9 cm) rachis, inflorescences spiciform with a single pair of proximal opposite branches or none; leaf blades usually with domatia in the distal vein axils • • P- viridis
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 229
22b. Flowers in dense subcapitate globose of ellipsoid inflorescences with a rachis less than 4 cm
long; leaf blades lacking large domatia P. alfaroana
23a. Stipules united over the shoot apex and forming a cap with a single acute apex as in Ficus (usually not split or open distally during development), narrowly conical to cylindrical and elongate, usually becoming more than 5 mm long before splitting, caducous as the leaves begin to expand (note that new lateral branches beneath the inflorescences often are enclosed within F/cws-like stipules after other stipules have fallen); inflorescences with opposite branches; leaves lacking a submarginal
vein 24
23b. Stipules united over the shoot apex only in very early stages (if present the united cap usually splitting before reaching 5 mm in length and open distally in later development), caducous or tardily deciduous, larger F/c«s-like stipules not present; inflorescences with opposite or whorled
branches; leaves with or without a submarginal vein 27
24a. Leaf blades usually with pit domatia or tufted hairs in the vein axils beneath, drying grayish
or pinkish gray above, with 10-13 pairs of major 2° veins; ca. 600 m elevation in the Cordillera
de Tilaran P. mexiae
24b. Leaf blades lacking pit domatia or tufted hairs in the vein axils, drying grayish to dark reddish
brown, with 5-16 pairs of major 2° veins; 400-2100 m elevation 25
25a. Stipules 2-6 mm long, central 2° veins arising at angles of 60-90°, leaf blades 4-12 x 1.5-
5 cm [drying dark grayish above, with 5-7 pairs of major 2° veins; fruits 4-7 mm long] . .
P. orosiana
25b. Stipules usually becoming more than 6 mm long; central 2° veins arising at angles of 50-70°,
leaf blades 6-22 x 2-10 cm 26
26a. Leaf blades usually drying grayish above, with 6-8 pairs of major 2°, usually narrowly elliptic
and with long petioles; stipules becoming 5-45 mm long; dried fruits 7-10 mm long
P. sarapiquensis
26b. Leaf blades usually drying reddish brown to dark brown (less often dark grayish), with 6-16
pairs of major 2° veins, very variable in shape and texture; stipules becoming 8-80 mm long;
fruits 5-9 mm long P. panamensis
27a. (from 23b) Inflorescences only 2-4 cm broad, with short (< 2 cm) lateral branches (sometimes becoming larger as the fruit develop) [flowers often closely congested distally]; leaf blades with 2° veins often weakly loop-connected near the margin; plants usually found in deciduous and partly
deciduous formations 28
27b. Inflorescences usually 4-8 cm broad, the lateral branches usually more than 2 cm long; leaf blades
with or without loop-connected 2° veins; plants not found in deciduous formations 31
28a. Corolla tube 4-5 mm long; fruit 6-9 mm long; inflorescences 1-3 cm long, with opposite
branches or the flowers subsessile at the end of the peduncle [broad ovate stipules with an
obtuse apex often subtending the young inflorescence] P. quinqueradiata
28b. Corolla tube 1-3.5 mm long; fruit 4-8 mm long; inflorescences 1 .5-10 cm long, with opposite
or whorled branches 29
29a. Inflorescences with only 2 branches at each node, often umbelliform; corolla tube 1.5-2 mm
long; stipules 6-14 mm long, acute and becoming narrowly 2-lobed at the apex; leaf blades
narrowly elliptic, to 20 cm long P. tenuifolia
29b. Inflorescences usually with 4 branches at the first node (2 longer and 2 shorter), rarely
umbelliform; corolla tube 2.2-3.5 mm long; stipules 2-8 mm long and bluntly obtuse; leaf
blades usually obovate to elliptic, to 1 5 cm long 30
30a. Pit domatia rarely present in distal vein axils beneath; leaves usually drying gray to pinkish
gray; calyx not usually persisting on the fruit; deciduous formations (in Costa Rica)
P. carthagenensis
30b. Domatia or tufts of hairs often present along the midvein beneath; leaf blades usually drying
gray to greenish gray [calyx often persisting on the fruit with lobes 0.5-3 mm long]; in both
evergreen and deciduous areas P. horizontalis
3 la. (from 27b) Inflorescences usually with 4 lateral branches at the first node (2 smaller descending
and 2 larger ascending); leaf blades usually narrowly elliptic to oblanceolate and with an arcuate
submarginal vein formed by the loop-connected secondaries (compare dichotomy 28 also) . . 32
31b. Inflorescences with 2 opposite lateral branches at the first node; leaf blades various 33
230 FIELDIANA: BOTANY
32a. Corolla tube 2-2.5 mm long; leaves usually drying grayish; wet lowlands (0-800 m elevation)
P. clivorum
32b. Corolla tube 3-5 mm long; leaves usually drying reddish or grayish brown; montane (2000-
2600 m elevation) forests p. sylvivaga
33a. Leaf blades usually with large (1-3 mm) domatia in the vein axils beneath, often opening by ellipsoid slits [blades 1 1-24 cm long, usually elliptic-oblong and drying dark reddish brown; inflorescence axes winged; peduncles ca. 2 mm thick; fruit 7-9 mm long; rarely collected in Costa
Rica] P. remota
33b. Leaf blades without large webbed domatia in the vein axils, smaller domatia present or absent
34
34a. Calyx lobes well developed (0.5-1 mm long); peduncles 1.5-2.5 mm thick when dried, primary branches of the flowering inflorescences 1-1.5 mm thick when dried; leaf blades lacking domatia'.
often oblanceolate, to 22 cm long; montane (1300-2600 m elevation) forest species 35
34b. Calyx lobes minute (0.3 mm) or absent; peduncles 0.5-1 .5 mm thick when dried, primary branches of the flowering inflorescences usually less than 1 mm thick when dried; leaf blades often with domatia on the lower surfaces, not usually oblanceolate or more than 1 5 cm long (except in P.
sarapiquensis); 0-2500 m elevation 37
35a. Primary branches of the inflorescence usually diverging at more than 100° from the more distal rachis (± reflexed); leaf blades with 10-15 pairs of major 2° veins, with or without an arcuate submarginal vein [flowers on open cymes on tertiary branches of a much-branched
inflorescence, peduncles 5-9 cm long; 1 700-2000 m elevation] P. stockwellii
35b. Primary branches of the inflorescence usually diverging at 90° or less from the rachis, per- pendicular or ascending; leaf blades with 8-1 1 pairs of major 2° veins; usually with a definite
submarginal vein 36
36a. Flowers in dense glomerules at the ends of the few (ca. 5) 1° or (ca. 9) 1° and 2° branches;
peduncles 1.5-5.5 cm long; 900-1500 m elevation P. monteverdensis
36b. Flowers in open cymes or pedicellate on 2° or 3° branches of the inflorescences; peduncles
4-1 2 cm long; 1 300-2600 m elevation P. sylvivaga
37a. Stipules developing 2 small (1-3 mm) slender lobes at the apex (4 acute tips per node) and usually
united to form a short tube at the base; corolla tube usually more than 3 rtim long 38
37b. Stipules acute at the apex (without small lobes), a short tube not apparent at the base; corolla tubes
rarely more than 3 mm long (except in P. chiriquina)
38a. Leaves drying dark reddish brown, petioles 7-18 mm long; stipules 4-10 mm long; 1500-
2500 m elevation P. chiriquina
38b. Leaves drying dark to pale grayish, petioles 0-5 mm long; stipules 2-5 mm long; 400-1900
m elevation P- orosiana
39a. Fruits 7-10 mm long, obovid-oblong; stipules conical, united and early caducous; leaf blades
glabrous beneath and without domatia [600-1600 m elevation] P. sarapiquensis
39b. Fruits 4-8 mm long (unknown in P. laselvensis), subglobose; stipules free distally, not united into
a conical cap; leaf blades glabrous or minutely puberulent and often with domatia beneath 40
40a. Highland (1500-2500 m) plants of easternmost Costa Rica and Chiriqui, Panama; leaves usually
drying reddish brown; corolla tube 3-6 mm long P. chiriquina
40b. Lower (0-1000 m) elevation plants; leaves usually drying grayish; corolla tube 1-3 mm long 41 4 la. Leaf blades with domatia rarely present in distal vein axils, blades to 13(-16) cm long; stipules
triangular; La Selva area, 50-300 m elevation P laselvensis
41b. Leaf blades with domatia often present along the central and proximal part of the midvein, blades to 17 cm long; stipules ovate-triangular; widespread, 0-1000 m elevation P. marginata
Psychotria acicularis C. M. Taylor, sp. nov. Figure TYPUS-/. Utley A K. Utley 1036 (holotypus CR. ««>-
„ typi F, MO!), from region to southwest of airstrip, 20-60
:>:>- m alt., 20 July 1974, Rincon de Osa, Puntarenas, Costa
Rica.
Species Psychotriae pittieri Standley affinis. sed ab ea
bracteismajoribus(3-10mmlongis)etangustoribusdif- Small shrubs, 0.5-2 m tall, leafy stems 0
fert thick, densely hirsutulous with thin hairs 0.4-1.5 mm
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 231
long, older stems glabrescent; stipules 3-8 mm long, with a short (1-3 mm) tube and 2 slender teeth 3-7 mm long on each side, densely hirsutulous in early stages, per- sisting or deciduous. Leaves with petioles 2-9(-13) mm long, ca. 0.7 mm thick, densely hirsutulous, often some- what unequal at the same node; leaf blades 5-12 cm long, 2-4.5 cm broad, narrowly ovate-elliptic to elliptic or elliptic-oblong, apex acuminate with a tip 4-12 mm long, base usually obtuse, drying thinly chartaceous or membranaceous, dark green above, with slender hairs 0.5-1.2 mm long on the upper surface, especially along midvein, becoming glabrescent, more densely hirsutu- lous beneath with slightly shorter more persistent hairs, 2° 10-15/side. Inflorescences terminal and solitary, 5- 12 cm long, open or elongate panicles with usually 4 branches at apex of the peduncle and 2 additional pairs of opposite or alternate branches, peduncles 3-8 cm long, ca. 0.7 mm thick (dried), hirsutulous with hairs to 1.5 mm long, bracts 3-10 mm long and linear, flowers sub- sessile in condensed distal cymes, bracteoles 2-4 m long, linear. Flowers densely puberulent externally, hypanthi- um ca. 1 mm long, sericeous with ascending thin whitish hairs ca. 0.5 mm long, calyx lobes 0.5-1 mm long, lan- ceolate; corolla white, narrowly salverform, tube ca. 3 mm long and 0.5 mm diam., corolla lobes ca. 1.2 mm long; anthers ca. 1 mm long. Fruits 3-3.5 mm long, 2.5- 3 mm diam., broadly ellipsoid, bluish with thin ascend- ing hairs ca. 0.5 mm long, persisting calyx lobes ca. 0.5 mm long.
Plants of the rain forest interior on the Osa Pen- insula, at elevations of 20-80 m. Flowering in May and July; fruiting in May-August. This species is only known from near Rincon de Osa on the Osa Peninsula in southwestern Costa Rica.
Psychotria acicularis is recognized by its hir- sutulous indumentum on all parts, small stature, small leaves, open inflorescences with conspicuous linear bracts, sessile flowers borne in distal clus- ters, small blue fruit, and limited geographic range. This species, a member of subgenus Heteropsy- chotria, resembles P. pittieri vegetatively but dif- fers in its linear bracts and inflorescences with four branches at the first (basal) node. Additional spec- imens seen were Aguilar 6423 (CR), Burch 4413 (DUKE, MO), Burger & Stolze5449(cR, F, MO), Duke 16111 (MO), Kennedy 1 934 (MO), Raven 21532 (CR, F), and Utley & Utley 1174 (CR, F).
Psychotria acuminate Benth., Bot. voy. Sulph. 107. 1 845. P. cuspidata sensu Standl. et auctores, non Bredem. ex Roem. & Schult. fide Steyermark 1974. Figure 56.
Shrubs or subshrubs, l-2(-3) m tall, much-branched, leafy stems 1 .5-5 mm thick, glabrous, quadrate to terete; stipules with a short (0.5-2 mm) truncated tube with 2 narrow awns 1.5-3 mm long on each side, these decid- uous. Leaves with petioles 5-15(-18) mm long, 0.7-1.4
mm thick, minutely (0.05 mm) papillate-puberulent; leaf blades 7-15(-19) cm long, 3-8(-10) cm broad, elliptic to oblong-elliptic or ovate-elliptic, apex short- to long- acuminate or caudate-acuminate with tip 1-3 cm long, base acute to obtuse, drying thin-chartaceous and green- ish or brownish, glabrous above and below, or with tuft- ed domatia in the vein axils beneath, 2° veins 7-12/side, midveins sometimes reddish beneath. Inflorescences solitary and terminal, 2-6 cm long (to 7 cm in fruit); 2- 4(-5) cm broad, paniculate, peduncles 1-4 cm long, 0.5- 1 mm thick, minutely papillate-puberulent (or appar- ently glabrous), primary branches opposite or alternate, bracts subtending the inflorescence branches absent or small (1 mm) and adnate to branches, floral bracts ab- sent, pedicels 0-2 mm long. Flowers distylous, minutely (0.05 mm) papillate-puberulent externally, hypanthium ca. 0.5 mm long and 0.5 mm diam., calyx 0.3-0.7 mm long with weakly denned lobes; corolla white or pale yellow, tube 2-4(1-5) mm long and 0.5-1 mm diam., lobes 5, 1.5-2.5 mm long, rounded; anthers 1-1.2 mm long, linear. Fruits 4 mm long, 5-7 mm diam., 2-lobed and rounded, smooth, becoming blue, bluish gray, or purple; pyrenes 2.5 mm long and 3 mm broad, hemi- spheric with a longitudinal sulcus on the inner face.
Plants of evergreen lowland rain forest forma- tions on both the Caribbean and Pacific slopes, from near sea level to 900(-1 100) m elevation in Costa Rica. Flowering in January-August in southern Central America (usually May-June at La Selva). Fruiting is primarily in June-Decem- ber. The species ranges from Mexico and the West Indies to Venezuela and Peru.
Psychotria acuminata is characterized by its gla- brous or minutely papillate-puberulent parts, lus- trous acuminate leaves with pronounced narrow drip tips, small flowers and inflorescences, 2-awned stipules, and smooth rounded bilobed blue fruit. In some specimens, the stipules appear to be un- developed. The small stature, leaf size, and foliage drying greenish are similar to many other species, but the small yellowish papillate-puberulent inflo- rescences and smooth bilobed fruit are distinctive. Also, the veins on the undersides of the leaves are sometimes pink. The presence of domatia-like tufts of hairs in the vein axils is uncommon among our species of Psychotria subgenus Heteropsychotria. The breeding biology was studied by Bawa and Beach (1983). Psychotria valeriana of higher ele- vations is similar but has a single stipular awn with an adaxial tooth on each side of the stem. [The name P. cuspidata deserves more careful re- view; it may be that Steyermark failed to recognize a specific distinction between the Caribbean ma- terial and his Venezuelan specimens.— C.M.T.]
Psychotria aggregata Standl., Contr. U.S. Natl. Herb. 18: 128. 19 16. P. tonduziiSiandl, J. Wash.
232
FIELDIANA: BOTANY
Acad. Sci. 15: 287. 1925. Montamans pana- mensis Dwyer, Ann. Missouri Bot. Gard. 67: 286. 1980. Figure 12.
Herbacous subshrubs with succulent stems, 0.5-1.5(-2) m tall, leafy stems 1 .6-6 mm thick (dried), quadrangular, glabrous; stipules 2-9(-12) mm long, to 7 mm broad, triangular with a short (2-3 mm) cupular base and bifid conical appendages 4-7 mm long, base persisting. Leaves with petioles 3-1 0(- 1 3) cm long, 1 -2 mm thick, glabrous; leaf blades 12-30(-38) cm long, (3-)5-17 cm broad, el- liptic-oblong to elliptic-obovate or elliptic-oblanceolate, apex tapering gradually and acute or acuminate, tip 5- 1 5(-22) mm long, base obtuse to cuneate or acute and decurrent on petiole, drying chartaceous and dark to pale grayish green, glabrous above, glabrous or minutely pa- pillate-puberulent beneath, 2° veins 10-19/side. Inflo- rescences axillary, solitary, 4-1 5 cm long, 2-4 cm broad, congested to capitate or with 1-2 pairs of lateral branch- es, peduncles l-5(-8) cm long, 1-2 mm thick, glabrous or minutely and sparsely puberulent, bracts at the lateral branches 2-8 mm long, flowers subsessile in closely crowded distal glomerules on the 1° or 2° branches, brac- teoles 1-4 mm long. Flowers with hypanthium 1-2 mm long, glabrous or sparsely and minutely puberulent, calyx 0.5-2 mm long, lobes 5, 0.5-4 mm long, obtuse to linear; corolla white, funnelform glabrous or minutely puber- ulent externally, tube 2-4 mm long, ca. 1 mm diam., corolla lobes 5, ca. 1 mm long, obtuse and galeate; an- thers ca. 1.2 mm long. Fruits 5-7 mm long (dried), 4-6 mm diam. (to 10 mm in life), ovoid to subglobose, spongy, white; pyrenes ca. 6 mm long, with margin and median keel thickened dorsally.
Plants of evergreen montane rain forest and cloud forest formations, from 10 to 2300 m elevation (but most common in the 400-1 700-m range). Ap- parently flowering and fruiting throughout the year. The species ranges from northern Costa Rica into western Panama.
Psychotria aggregata is recognized by its short succulent unbranched main stems, large essen- tially glabrous leaves, axillary inflorescences with flowers in dense bracteolate clusters, and spongy white fruits. This species is quite similar to P. macrophylla but differs in the more condensed inflorescences and the pyrenes. (This and related species have recently been studied by Molly Ne- pokroeff, wis, 1 992.) The type of P. aggregata (Pit- tier 3264 us from Horqueta, Chiriqui, Panama) is a very poor specimen, but it is an early name for this variable taxon, commonly called P. tonduzii by previous authors.
Psychotria alfaroana Standl., J. Wash. Acad. Sci. 18:273. 1928. Figure 66.
Herbaceous erect subshrubs, 0.2-0.6(-1) m tall, usu- ally with a single erect unbranched stem, rhizomatous,
leafy stems 2-4 mm thick, glabrous, terete; stipules to 20 mm long, basal sheath 5-8 mm long and 4 mm broad, with 2 narrow distal lobes 3-12 mm long, glabrous or sparsely puberulent, caducous. Leaves with petioles (2-)6- 25(-30) mm long. 1.5-2 mm thick, sulcate above, gla- brous; leaf blades 1 1-22(-30) cm long, 4-1 1 cm broad, obovate to elliptic-obovate or elliptic-oblanceolate. apex short-acuminate or subacutc. base gradually narrowed and cuneate, decurrent on petiole, drying thin- to stiffly chartaceous, grayish or grayish green, glabrous above. with minute (0. 1-0.3 mm) thin hairs along major veins beneath, 2° veins 10-14/side, weakly connected by an arcuate submarginal vein. Inflorescences terminal, sol- itary, 3-8 cm long, 2-4 cm broad, compact globose or elongate dense panicles of cymes, peduncle 1-4 cm long, 1.2-2 mm thick, glabrous, bracts 0.5-1 mm long, ped- icels 1-2 mm long. Flowers glabrous externally, hypan- thium ca. 1.5 mm long, calyx tube ca. 1 mm long, cu- pulate, calyx lobes 5, 0.5-2 mm long, narrowly oblong; corolla white, tube 2.5-4 mm long, ca. 2 mm diam.. lobes 5, 1 .5-3 mm long, ca. 1 mm broad at base; stamens 5, anthers 0.8-1.3 mm long. Fruits 8-12 mm long, 4-6 mm diam., ellipsoid, red at maturity, becoming black when dried, persisting calyx to 3 mm long; pyrenes ca. 7 mm long, usually with 5 dorsal ribs.
Plants of wet rain forest formations of the Ca- ribbean slope and continental divide, 20- 900(-1100) m elevation. Flowering in January- September; fruiting in January-February and June- August. The species ranges from the Cordillera de Guanacaste in Costa Rica to Bocas del Toro Prov- ince in Panama.
Psychotria alfaroana is recognized by its very short stature, usually obovate leaves, compact subglobose inflorescences, and larger fruit. The bright red fruit and leaves drying grayish are char- acteristics of subg. Psychotria. Standley and Vale- ric made a number of collections of this species near Tilaran, including the holotype (41579 us). A collection from along Rio Corobici (Opler 138 F) is provisionally placed here; this is the only collection from the Pacific lowlands. Several col- lections with exceptionally large (35 x 15 cm) leaves attenuate almost to the base and then abruptly subauriculate are tentatively placed here: Gdmez-Laurito 11570, Hammelet al. 16895, and Herrera & Chacdn 2355 (all at CR).
Psychotria allenii Standl., Ann. Missouri Bot. Gard. 27: 342. 1940.
Small trees or shrubs, (2-)4-10 m tall, leafy stems 1- 4 mm thick, glabrous or minutely and sparsely puber- ulent; stipules united around the stem for 0.5-3 mm, with 2 acute lobes 1-5 mm long, separated by a V- or U-shaped sinus, persisting. Leaves with petioles 6-18 mm long, 0.8-1.3 mm thick, glabrous; leaf blades 5-16 cm long, 3-7(-8) cm broad, ovate-elliptic or ovate-ob-
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
233
long to narrowly elliptic-oblong, apex acuminate with tip ca. 5 mm long, base obtuse to slightly rounded, drying dark olive green or grayish green and chartaceous, gla- brous above, glabrous or minutely puberulent beneath, 2° veins 5-7/side, distal veins strongly arcuate-ascend- ing. Inflorescences 8-10 cm long, 5-7 cm broad, pyra- midal paniculate with distal branches progressively shorter, peduncles 3.5-6.5 cm long, 0.9-2.3 mm thick, glabrous or puberulent, the first node with 4 lateral branches 2-3.5 cm long subtended by linear bracts 1-3 mm long, flowers mostly sessile and crowded in distal cymes. Flowers with hypanthium 0.4-0.8 mm long, gla- brous or sparsely papillate-puberulent, calyx ca. 0.7 mm long with glabrous lobes ca. 0.3 mm long; corolla white, tube 0.7-2 mm long and 1 mm diam., lobes 5, ca. 1.7 mm long; stamens 5. Fruits 6-1 5 mm long and ca. 6 mm diam. when dried, bright blue; pyrenes smooth, subglo- bose.
Plants of moist evergreen lower montane for- ests, 50-900(-1500) m elevation. Flowering and fruiting in February-August in Panama. The spe- cies is known only from Monteverde in Costa Rica (Koptur 74 MO with flowers and early fruits in October). The species ranges to central Panama.
Psychotria allenii is recognized by its distinctive pyramidal inflorescences with relatively thick pe- duncles, four stout lateral branches at the first few nodes, and small sessile flowers. This species is quite similar to P. officinalis, but that species has floral bracts and larger flowers and lacks the robust lateral branches of the inflorescence. The Costa Rican collection has narrower elliptic-oblong leaves than Panamanian material, and it was a 2-m shrub rather than a tree.
Psychotria angustiflora K. Krause, Bot. Jahrb. 54, Beibl. 119: 43. 1916. P. mima Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 386. 3 1 Oct. 1 940 (holotype: Skutch 4589 F from Costa Rica), not P. mima Standl., loc. cit. 22: 204. 10 Sept. 1 940 (holotype: L. B. Smith 1878 F from Brazil). Figure 57.
Shrubs or small trees to 5 m tall, leafy stems 1.5-5 mm thick, glabrous, terete; stipules 3-8 mm long, 2-6 mm broad, triangular to ovate and rounded at the nar- rowed tip, with a short (1-3 mm) tube, obtuse or bilobed with a small ( 1 mm) sinus, glabrous, persisting. Leaves with petioles 15-35(-60) mm long, 1.2-3 mm thick, gla- brous, with 2 adaxial ridges; leaf blades (9-) 1 5-24(-30) cm long, (4-)7-14(-16) cm broad, oblong to ovate-ob- long, broadly elliptic or broadly elliptic-obovate, apex short-acuminate with a tip 5-10 mm long, base broadly obtuse (rarely acute), drying thin-chartaceous and green- ish gray or greenish brown above, glabrous above and below, 2° veins 8-1 I/side, central 2° veins ca. 2 cm dis- tant in larger leaves. Inflorescences terminal, solitary (3-) 17-45 cm long, (8-) 15-25 cm broad, pyramidal pa-
niculate with distant opposite lateral branches, peduncles 6-15 cm long, 1.5-2 mm thick, glabrous or minutely puberulent, bracts to 7 mm long and 1.5 mm broad, flowers subsessile in distal cymes or along slender distal branches, subtended by bracteoles ca. 1 mm long. Flow- ers minutely papillate-puberulent externally, hypanthi- um ca. 0.7 mm long, calyx ca. 0.4 mm long, calyx lobes ca. 0.2 mm long, acute; corolla pale yellow or greenish yellow (rarely white), tubular-obovoid or funnelform, tube 5-7.5 mm long, 0.9-1 .3 mm diam. in the lower half and 2-3 mm distally, lobes 5, 1-2.5 mm long, triangular and acute, erect. Fruits 4-5 mm long and 3-5 mm diam. (to 8 mm in life), subglobose with ca. 6 longitudinal ribs, becoming blue or black; pyrenes 1.5-3.5 mm long.
Plants of evergreen rain forest formations, from near sea level to 900 m elevation on the Caribbean slope. Flowering in January-August; fruiting in June and October-November. The species ranges from near La Selva, Heredia, to Brazil.
Psychotria angustiflora is recognized by its large open inflorescences with small distal bracteoles, narrowly obovoid yellowish corolla, larger leaves, and ovate stipules that are obtuse or minutely two- lobed at apex. The stipules may become two-part- ed as the expanding stem breaks them apart. The flower buds are quite narrow before the corolla is fully expanded. The isotype of P. angustiflora (Tonduz 12996 us) fits in well with the material placed here. This species should be compared to P. berteriana (smaller flowers, different stipules, narrower leaves) and P. luxurians (smaller flowers, larger bracteoles).
Psychotria aubletiana Steyerm., Mem. New York Bot. Card. 23: 694. 1972. Cephaelis axillaris Sw., Prodr. 45. 1788, not P. axillaris Willd. P. aubletiana var. centro-americana Steyerm., loc. cit. 698. 1972.
Herbs or subshrubs, 0.5-1 .5(-4) m tall, erect and few- branched, stems slightly succulent, leafy stems 1-5 mm thick, glabrous or rarely densely puberulent with soft yellowish hairs ca. 0.2 mm long, quadrangular, drying dark; stipules 5-10 mm long, to 6 mm broad, basal sheath 3-5 mm long, ovate with a short (2-4 mm) sinus and 2 irregular rounded lobes per side. Leaves with petioles 6- 20(-40) mm long, 0.7-1.8 mm thick, glabrous or mi- nutely puberulent; leaf blades 4-15(-18) cm long, 1- 4(-6.5) cm broad, elliptic-oblong, narrowly oblong-ob- ovate to narrowly elliptic-obovate, oblong-lanceolate or narrowly elliptic, apex acuminate (rarely acute) with tip 4-1 2 mm long, acute at the base (obtuse in larger leaves), drying stiffly chartaceous, dark greenish above, much paler beneath, glabrous above, glabrous beneath or with short (0.1-0.2 mm) hairs on major veins, 2° veins 6-87 side (with lesser parallel intermediate 2° veins). Inflo- rescences axillary and sessile, 1.3-2 cm diam., densely compact, subglobose, with distal 2-lobed stipules resem-
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bling bracts, bracts obovate, purple or green, 2-7 mm long, flower 5-1 5/head, sessile. Flowers with 5 calyx lobes 1-2 mm long; corolla white, glabrous, tube ca. 10 mm long, lobes 5, 3-3.5 mm long, triangular, stamens 5, anthers 1-2 mm long. Fruits 5-7 mm diam., ellipsoid, bright blue; pyrenes 3-5 mm long and 2-4 mm broad.
Understory plants of evergreen montane rain forest formations, from (600-)1 200 to 2300(-2800) m elevation in Costa Rica. Probably flowering and fruiting throughout the year. The species ranges from Guatemala through Central America to Co- lombia, Venezuela, and the lesser Antilles.
Psychotria aubletiana is recognized by its small stature, sessile rounded inflorescences often encir- cling distal nodes and subtended by an involucre of broad bracts, short white corolla tubes, and bright blue fruit. This species is not easily confused with other species of Psychotria.
Psychotria aurantibractea C. M. Taylor, nom. nov. Cephaelis pittieri K. Krause, Bot. Jahrb. Syst. 54, Beibl. 1 19: 45. 1916, not Psychotria pittieri Stand i. Figure 18.
Shrubs, 1-3 m tall, leafy stems 1.3-4.5 mm thick, glabrous, often narrowed below the node when dried; stipules united for only 0.5-2 mm around stem, with 2 narrowly triangular or linear teeth 2-8 mm long, sepa- rated by 2-3 mm, persisting. Leaves with petioles 5-40 mm long, 0.8-1.5 mm thick, glabrous; leaf blades 8- 21 (-28) cm long, 2-6 cm broad, narrowly oblong to nar- rowly elliptic-oblong or lanceolate, apex gradually ta- pering and acute or acuminate, tip to 1 5 mm long, base cuneate to acute, drying membranaceous or thin-char- taceous and dark brown, glabrous above, glabrous or sparsely and minutely (0. 1 mm) puberulent on the veins beneath, 2° veins 10-16/side. Inflorescences terminal, 1 or 3, 2.5-6 cm long, 4-6 cm broad (to 8 cm when tri- partite), usually with 3 capitate or cymose clusters of flowers on short (1-4 mm) primary branches, peduncles 5-25 mm long, 1.3-2.2 mm thick, minutely puberulent, bracts 8-15 mm long, 4-9 mm broad, ovate to lanceo- late, acuminate, orange (rarely red), sparsely puberulent and ciliolate along margins, flowers sessile. Flowers en- closed by bracts and bracteoles, hypanthium ca. 1 .5 mm long, densely pubescent with straight ascending hairs, calyx lobes 5, ca. 1.3 mm long, dentate with narrrow lobes; corolla yellow or yellow-rose, tube ca. 5 mm long, lobes ca. 2 mm long. Fruits 5-7 mm long, 2.5-5 mm diam., ovoid-ellipsoid, with prominent longitudinal cos- tae and short thin hairs, color unknown, persisting calyx 1 .3-2 mm long, conical; pyrenes ridged.
Plants of continually wet or seasonally dry ev- ergreen forest formations, from 200 to 1200 m elevation. Flowering in May-July; fruiting in Au- gust-September and January. This species is en- demic to southern Costa Rica, from San Isidro del
General to San Vito and the Osa Peninsula, and on the adjacent Caribbean slope (at 83°W).
Psychotria aurantibractea is recognized by its terminal inflorescences composed of large ovate, orange or reddish bracts, narrow leaves, and re- stricted geographical range. The colorful bracteate inflorescences made this and other species for- merly placed in Cephaelis so distinctive in Central America.
Psychotria berteriana DC., Prodr. 4: 515. 1830. Figure 58.
Shrubs or small trees, 1.5-6(-8) m tall, leafy stems (1.5-)3-5 mm thick, glabrous or minutely puberulent with hairs 0.1-0.2 mm long; stipules at first triangular and acute with a small distal cleft, becoming 2-lobed or 2-awned (on each side), basal sheath 0.5-2 mm long with acute lobes 1-3.5 mm long. Leaves with petioles 10- 30(-52) mm long, 0.6-1.8 mm thick, glabrous or mi- nutely puberulent; leaf blades 8-20(-26) cm long, 3- 9(-ll) cm broad, narrowly ovate-elliptic to elliptic or elliptic-oblong, apex acuminate with 5-15 mm long, base acute and cuneate to obtuse (rarely rounded and sub- truncate in large leaves), drying membranaceous to thin- chartaceous, dark green or brown, glabrous or with min- ute (0.1-0.2 mm) hairs along the veins above, glabrous or with thin whitish hairs ca. 0.2 mm long beneath, 2° veins 7-15/side. Inflorescences terminal and solitary. 5- 16(-22) cm long, 5-15 cm broad, oblong but becoming broadly pyramidal paniculate with opposite or alternate branches, yellowish in life, peduncles 5-10 cm long. 1 .3- 2.2 mm thick, sparsely puberulent with thin hairs ca. 0.2 mm long, bracts 3-8 mm long, ca. 0.6 mm broad, flowers sessile in distal cymes, bracteoles 1-2 mm long. Flowers minutely puberulent externally (rarely glabrous), hypan- thium ca. 0.5 mm long, calyx lobes ca. 0.2 mm long; corolla tubular-salverform, yellow to white, tube 1.5-4 mm long, 1-2 mm diam., glabrous proximally, lobes 5. 1-1.5 mm long; anthers 1-1.4 mm long. Fruits 3—4 mm long (5 mm including the persisting calyx), 3-4 mm diam.. subglobose, fleshy, lustrous black; pyrenes with 3-5 dor- sal ridges.
Plants of evergreen rain forest formations, from near sea level to 1 700 m elevation (to 1 200 m on the Pacific slope of southern Costa Rica). Flow- ering throughout the year; probably fruiting throughout the year. The species ranges from Mex- ico to South America.
Psychotria berteriana is characterized by its large, many-branched, pyramidal inflorescences, thin leaves often with long slender petioles and drying dark, two-lobed stipules, small flowers, and small rounded fruit. Compare P. luxurians and P. an- gustiflora (with longer thinner corolla tubes). A broader interpretation of P. berteriana might re- quire the inclusion of P. luxurians. The greenish
BURGER & TAYLOR: FLORA COSTARICENSIS. RUB1ACEAE
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yellow inflorescences resemble those found in Pal- icourea tilarensis.
Psychotria boquetensis Dwyer, Ann. Missouri Bot. Gard. 67: 349. 1980.
Shrubs, 0.5-2 m tall, leafy stems 1.2-3.5 mm thick, tomentulous with reddish hairs 0.4-1 mm long; stipules 8-14 mm long, 2-6 mm broad, oblanceolate to ovate, obtuse to acuminate at apex (not bilobed in the type as described by Hamilton, 1989), caducous. Leaves with petioles 1-6 mm long, 0.7-1.2 mm broad, with curved thin reddish hairs ca. 0.3 mm long; leaf blades 4-16 cm long, 1-4 cm broad, narrowly elliptic-oblong to oblan- ceolate, apex acuminate with tip 4-15 mm long, base gradually narrowed and acute or cuneate, decurrent on petiole, drying thinly chartaceous and dark reddish brown, puberulent along the primary and secondary veins above and below, with hairs 0.2-0.7 mm long along the margin, 2° veins 10-13/side, with minute tufted domatia in the axils beneath. Inflorescences terminal or pseudoaxillary, several at a node, 2-4 cm long, small panicles, peduncles ca. 20 mm long and 0.6 mm thick, bracts represented by dense whitish hairs 0.5-1 mm long, flowers sessile or subsessile in compact cymes. Flowers glabrous on the distal surfaces, hypanthium ca. 1 mm long and puber- ulent, calyx lobes 1-1.5 mm long, narrowly triangular, corolla white, urceolate, tube 1.5-2.5 mm long and 1 mm diam., lobes 5, 1-1.5 mm long; stamens 5, anthers 0.5-0.7 mm long. Fruits unknown.
Plants of the Boquete region of the Chiriqui Highlands at about 1 200 m elevation. Flowering material was collected in May-June. The species is only known from western Panama.
Psychotria boquetensis is recognized by the nar- row leaves with conspicuous reddish hairs, mem- branaceous broad caducous stipules, small inflo- rescences, and narrow calyx lobes to 1.5 m long. The species is closely related to P. nervosa but differs in the large calyx lobes, narrower leaves, and smaller inflorescences. This distinctive species is a member of subgenus Psychotria.
Psychotria borucana (A. Molina) C. M. Taylor & W. Burger, comb, nov., Cat. fl. Peru (in press). Cephaelis affinis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 1 84. 1 930, not P. affinis Baker. C. borucana A. Molina, Ceiba 4: 31. 1951. Fig- ure 17.
Shrubs or woody herbs, 1.5-4 m tall, leafy stems 2.5- 8 mm thick, glabrous and drying dark, quadrangular or flattened; stipules 9-20 mm long and 5-10(-15) mm broad, basal sheath 1-2 mm long, ovate-triangular, ob- tuse, with parallel venation, glabrous or minutely brown- ish puberulent. Leaves with petioles 12-50 mm long,
1 .7-3(-5) mm thick, glabrous (as in the type) or minutely papillate-puberulent with erect brownish hairs 0. 1 mm long; leaf blades 1 5-28(-32) cm long, 7- 14(- 16) cm broad, obovate to elliptic obovate (as in the type) to broadly elliptic, apex short-acuminate (acute) with tip 5-15 mm long, base gradually narrowed and cuneate to obtuse, drying stiffly chartaceous, dark grayish above and mark- edly paler beneath, glabrous above, glabrous beneath (as in the type) or with minute hairs 0.1-0.2 mm long throughout, 2° veins 6-10/side. Inflorescences terminal, capitulae solitary, 2-4 cm broad, peduncle 5-20 cm long, 2.5-4 mm diam., glabrous or minutely puberulent, basal bracts mostly 2, 4-15 mm long, broadly ovate to reni- form, interior bracteoles 4-8 mm long, spatulate, pale purple, lavender, or green marked with purple, flowers enclosed by bracts. Flowers with hypanthium ca. 2 mm long, calyx 1-3 mm long and 3 mm broad; corolla fun- nelform, white, glabrous, 20-22 mm long, tube 14-19 mm long and 2-4 mm diam., lobes 5(-6), 2.5-3 mm long 1.5- 2 mm broad at the base, acute; stamens 5(-6), an- thers subsessile, ca. 2.8 mm long; style ca. 16 mm long. Fruits ca. 8 mm long and 8 mm diam., rounded-oblong, becoming blue or purple, 1 0-ribbed when dried; pyrenes with 4-6 ribs.
Plants of evergreen forest formations of the Pa- cific slope, from around 50 to 500 m elevation in southern and south-central Costa Rica. Flowering in May-July; fruiting in late August and Novem- ber. The species is also known from Peru.
Psychotria borucana is recognized by its large leaves, long-pedunculate purple heads, two broad inflorescence bracts with reticulate venation, long corolla tube with short lobes, unusual stipules, and restricted range (in Costa Rica). The leaves are quite variable in shape. Compare P. correae with pendant heads and more secondary veins.
Psychotria brachiata Sw., Prodr. 45. 1788. Figure 58.
Shrubs or small trees, (0.5-)1.5-3(-5) m tall, leafy branchlets 1.2-5 mm thick, glabrous, quadrangular be- coming terete; stipules 3-7 mm long, 2-5 mm broad, rounded and bilobed with a sinus 0.5-2 mm deep and 2 rounded lobes, glabrous and persisting. Leaves with petioles (1-) 1.5-3. 5 cm long, 1-1.5 mm thick, glabrous; leaf blades 9-17(-21) cm long, 3-7(-ll) cm broad, el- liptic to elliptic-oblong or elliptic-obovate, apex usually short-acuminate with tip 5-10(-l 5) mm long, base acute to attenuate (obtuse), drying thin-chartaceous or char- taceous and greenish or brownish, glabrous above, gla- brous beneath except for a line of short (0.5 mm) hairs along sides of midvein, 2° veins 7-10/side. Inflorescences terminal and solitary or 3 (4, 5), 10-20 cm long, 3-10 cm broad, open pyramidal panicles with opposite sep- arated (6-30 mm) lateral branches mostly arising at 90° angles, peduncles 1.5-6 cm long, with thin hairs ca. 0.4 mm long (mostly in 2 longitudinal rows), often with 3 2° branches bearing the distal flower clusters, bracts 3- 8 mm long and 1.5-3 mm broad, lanceolate, floral bracts
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2-3 mm long, broadly ovate, ciliolate along the margins, pedicels 0-2 mm long, flowers borne in capitulae 5-10 mm broad. Flowers with hypanthium 1-1.5 mm long, calyx 1-1.5 mm long, yellowish, calyx truncate or slightly dentate; corolla yellow or white (often with bluish tips and yellow throat), funnelform, tube 3-5 mm long and 0.7-2 mm diam., usually glabrous externally on the low- er half, lobes 5, 1-1.5 mm long. Fruits 4-5 mm long (including calyx) and 3-6 mm diam., obovoid to ellip- soid but becoming subglobose and spongy in life (to 10 mm diam.), bright blue to dark blue or purple, with prominent longitudinal ribs, persisting calyx ca. 1 mm long; pyrenes 3-4 mm long, with 3-5 ribs.
Plants of secondary vegetation in evergreen tropical and premontane wet forest formations on the Caribbean slope and in the General Valley, from 5 to 800 m elevation in Costa Rica. This species appears to flower and fruit throughout the year in Central America. At La Selva flowering is primarily in May-July and fruiting is mostly in July-September. The species ranges from the West Indies and Mexico through Central America to Peru.
Psychotria brachiata is recognized by its open inflorescences with well separated short opposite branches bearing closely clustered yellowish flow- ers in distal groups subtended by prominent per- sisting bracts. The small stature, nearly glabrous leaves usually drying greenish, spongy bright blue fruit with longitudinal ribs, and persisting stipules are additional features. Immature inflorescences of P. brachiata are racemose with the congested flowers in unexpanded capitula at the ends of the opposite lateral branches.
Steyermark (1972, pp. 585-586) distinguishes P. caerulea Ruiz & Pavon from P. brachiata by its larger corollas (11-12 mm long), external co- rolla surface that is densely puberulent through- out, more prominent calyx lobes, larger clusters of flowers, and larger floral bracts forming a more definite involucre. Specimens that fit Steyermark's definition of P. caerulea appear to occur in Bocas del Toro, Panama (D. Simpson in herb. 1977), and it is possible that they also occur in nearby Costa Rica. The question of whether or not these two species intergrade and are conspecific might make a worthwhile study.
Psychotria brachybotrya Muell.-Arg. in Mart., Fl. Bras. 6(5): 327. 1881. P. iquitosensis Standl., Publ. Field Columb. Mus., Bot. Ser. 8: 195. 1930. Figure 56.
Shrubs or subshrubs, l-2(-3) m tall, leafy branchlets 0.8-4 mm thick, glabrous; stipules separate or with a
short (0.5-1 mm) sheath, lobes 2/side, 2-7 mm long, 0.5-1 mm broad at base, linear to narrowly triangular, usually separated by a U-shaped sinus, glabrous, per- sisting. Leaves with petioles 3-10 mm long (poorly dif- ferentiated from the decurrent leaf base), glabrous; leaf blades 9-16(-18) cm long, 3-7(-8) cm broad, ovate-el- liptic to ovate or elliptic, apex usually tapering gradually and short-acuminate or acute, tip 5-10 mm long, taper- ing gradually or abruptly to the attenuate base (acute to broadly obtuse above the narrowed base), drying thinly chartaceous and greenish, glabrous above and below, 2° veins 4-8/side and arcuate ascending, distal 3° veins of- ten subparallel. Inflorescences terminal and solitary, 1 .2- 3 cm long but enlarging in fruit, 1-2 cm wide, narrowly paniculate with short opposite lateral branches or some- times subcapitate, peduncle 3-18 mm long, ca. 1 mm thick, with thin whitish hairs to 0.5 mm long, 2° branches 0-5 mm long, cymes congested, usually subtended by 3 involucrate narrowly ovate to lanceolate bracts, median bracts 3-6 mm long and 2-3 mm broad, with 2 shorter lateral bracts, all 3 united at base, flowers sessile. Flowers with hypanthium ca. 0.5 mm long, puberulent, calyx lobes 5, 0.2-0.3 mm long; corolla white, salverform, tube 1.5-4 mm long, ca. 1 mm diam. distally, glabrous or puberulent, lobes 5, 0.8-1.5 mm long. Fruits ca. 4 mm long and 4 mm diam., globose to oblong, becoming pur- ple-black at maturity (blue-black on Cocos Island), with prominent longitudinal ribs (transverse ridges some- times visible when dried), calyx minute (0.3 mm high) or obscure; pyrenes 3-4 mm long, with 4-5 prominent ridges.
Plants of the lowland rain forest formations on both the Caribbean and Pacific slopes, from near sea level to 400(-900) m elevation and usually found in shaded, poorly drained sites. Flowering in July-August; fruiting in August-October and December. In addition to the mainland plants, a distinctive population of this species is found on Cocos Island (see below). The species ranges from Costa Rica and Panama to Brazil and Bolivia.
Psychotria brachybotrya is distinguished by its well-separated stipule lobes, glabrous leaves dry- ing green, small compact inflorescences, flowers subtended by an involucre of bracts, small white flowers, and purplish black fruit. Inflorescences change in shape as they grow and expand. Collec- tions from Cocos Island placed under this name differ from the mainland material in having small- er leaves with blades (4-)6-9 cm long and 1.5-3.5 cm broad; the stipules, flowers, and fruit are very similar. Compare this species to P. hoffmannseg- giana, P. platypoda, and P. officinalis. Hammel (in Taylor, 1991) noted that P. brachybotrya is usually a larger plant than P. officinalis and the latter has a linear ventral sulcus on the pyrene.
Psychotria ca loch lam ys Standl., Publ. Field Co- lumb. Mus., Bot. Ser. 8: 199. 1930.
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Small shrubs or treelets, 1.5-4.5 m tall, leafy stems 2.5-4.5 mm thick, glabrous or sparsely and minutely puberulent; stipules 1 3-28 mm long and 5-9 mm broad at the base, ovate-triangular with 2 attenuate-acute lobes 6-10 mm long, greenish to wine red or violet, with fine parallel venation, chartaceous. Leaves with petioles 3- 16(-25) mm long, 0.9-2 mm thick, glabrous; leaf blades 8-23 cm long, 2-8.5 cm broad, elliptic-ovate to oblong- lanceolate or lanceolate, apex gradually tapering and acu- minate, tip 5-10 mm long, base acute to obtuse, drying chartaceous and greenish, glabrous above and below, 2° veins 1 1-17/side, often with lesser 2° veins between the major. Inflorescences solitary and terminal, 12-14 cm long and 6-7 cm broad, hemispheric panicles with 2- 3(-4) branches at the first and second nodes, peduncles 3-8 cm long and 1.5-2 mm thick, white, minutely (0.1 mm) puberulent, bracts subtending the first whorl of branches 1 .5-2 cm long, 3-4 mm wide, narrowly elliptic, resembling the stipules in texture, distal bracteoles 5-14 mm long, 2-4 mm broad, lanceolate, persisting, white to violet, enclosing the sessile flowers. Flowers ca. 15 mm long, hypanthium 0.6-1 mm long, calyx 0.6-0.8 mm long, lobes 5, triangular; corolla white to purple or pale violet, yellow in throat, tubular-salverform, tube 7-10 mm long, lobes 5, 2-3 mm long; anthers ca. 1.5 mm long and curved. Fruits ca. 6 mm long and 4 mm diam., oblong, dark purple or black; pyrenes with 4-5 ribs.
In Costa Rica this disjunct species is only known from the hills near Golfito, Puntarenas, at about 1 00 m elevation, flowering in December-January. The species also occurs in Choco, Colombia, and Amazonian Peru.
Psychotria calochlamys is distinguished by its large deciduous stipules, long and persisting white or lilac bracts and bracteoles, and large flowers. The unusual form and size of the stipules is similar in P. capitata, a close relative. The inflorescence with large white (or purplish) bracts and flowers is unlike any other Costa Rican species of Psy- chotria and reminiscent of some species of Pali- courea. The disjunct occurrence in Costa Rica is similar to that of some other species in the Golfo Dulce region. This species may prove to be con- specific with P. stipulosa Muell. Arg. of Amazo- nian South America, but that species (interpreted in a broad sense) includes variation not seen in our material.
The leaves drying grayish and the bright red fruit are characteristics of the subgenus Psychotria. This species has a disjunct distribution and is not known from Costa Rica. It is found in southern Mexico and Guatemala and again in Colon and Code provinces in Panama.
Psychotria camponutans (Dwyer & Hayden) Ham- mel, Selbyana 12: 139. 1991. Cephaelis cam- ponutans Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 35. 1968.
Semisucculent herbs or subshrubs 0.5-1.2 m tall, rhi- zomatous, stems unbranched, leafy stems 2—4 mm thick, quadrangular or terete, glabrous, drying black and con- tracted below the nodes; stipules 6-8(-15) mm long, 5- 6(-10) mm broad, broadly ovate with a serrated margin and 2 long lobes, glabrous and thin, distal parts decid- uous but with a persisting truncated sheath 1-1.5 mm long. Leaves with petioles 18-55 mm long, 1.2-2 mm thick, glabrous, drying black; leaf blades (1 1-)18-25 cm long, (4-)7-9 cm broad, narrowly elliptic-oblong to ob- long or oblong-obovate, apex acuminate with tip 6-13 mm long, acute at base and slightly decurrent on petiole, drying thinly chartaceous, dark or grayish above, gla- brous above and below, 2° veins 8-15/side but these difficult to see. Inflorescences axillary and I/node, 1-2 cm long, to 2 cm diam., capitate, peduncle 4-10 mm long, with involucrate bracts 8-15 mm long, 5 mm broad, ovate and acuminate, glabrous and green, flowers sessile and congested, each enclosed by 2 lanceolate bracteoles 6-8 mm long. Flowers glabrous, calyx lobes 5-6, ca. 1 mm long; corolla white, funnelform, tube ca. 3 mm long, corolla lobes 1-1.5 mm long. Fruits 7-8 mm long, 4-5 mm diam., ellipsoid to obovoid, crowned by a persisting calyx, deep maroon red or pale rose red, bracts red to purple; pyrenes 6-7 mm long, ribbed.
Plants of the wet evergreen forests of the Carib- bean slope, from 300 to 900 m elevation. Flow- ering in April and July; immature fruit were col- lected in September. Costa Rica to central Panama.
Psychotria camponutans is distinguished by its short semisucculent erect stems with small axillary capitate inflorescences, reddish fruit, and restric- tion to the Caribbean slope. Compare P. uliginosa and P. cartagoensis.
Psychotria calophylla Standl., Contr. U.S. Natl. Herb. 18: 129. 1916.
Small trees, 3-18 m tall, with larger (13-26 x 5-12 cm) elliptic to obovate leaves usually reddish tomentulose beneath, with an arcuate submarginal vein; the inflorescences paniculate with usually 3 ranks of branching, the flowers in distal glomer- ules, and larger (10-14 x 6-9 mm) ellipsoid fruit.
Psychotria capacifolia Dwyer, Ann. Missouri. Bot. Gard. 67: 353. 1980. Figure 54.
Herbs or subshrubs, 0.4-1 (-2?) m tall, usually suc- culent and unbranched, leafy stems 3-12 mm thick, gla- brous or with hairs to 1 mm long at nodes, terete, drying dark; stipules 3-6 mm long, to 8 mm broad, broadly obtuse to truncate and entire, with a conical caducous lateral appendage to 3 mm long, densely hirtellous. Leaves densely hirtellous in early stages but usually glabrescent
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in age, petioles 3-10 cm long, 2-4 mm thick, with hairs ca. 1 mm long in early stages; leaf blades 20—40 cm long, 9-1 6(-20) cm broad, oblong to elliptic-oblong or elliptic- obovate, apex abruptly narrowed and short-acuminate, base cuneate to obtuse, drying thinly chartaceous and grayish green, usually glabrous above, glabrescent or with thin hairs to 1 .4 mm long beneath, 2° veins 1 7-22/side, connected by a distinct slightly arcuate submarginal vein 2-3 mm from the margin. Inflorescences axillary, I/node, (8-) 15-3 5 cm long, 5-12 cm wide, open panicles with opposite branching, peduncles 7-22 cm long, 1.5-2.2 thick, with thin yellowish to reddish brown hairs 0.2- 0.7 mm long, bracts 1-2 mm long, flowers sessile on distal dichotomous branches or distal clusters, bracteoles ca. 0.5 mm long. Flowers puberulent externally, hypan- thium ca. 0.7 mm long, calyx to 1 mm long, calyx lobes 5, 0.3-0.5 mm long, triangular; corolla white or pale yellow, funnelform, tube 2.5-4 mm long, 0.5-1 mm diam. basally and 2 mm distally, lobes 5, ca. 1 mm long. Fruits 5-6 mm long, 3-4 mm broad, ovoid, yellow-green or whitish, with ca. 10 longitudinal ribs; pyrenes ca. 5 mm long.
Plants of evergreen rain forest formations on the Caribbean slope, ranging from 5 to 1300 m ele- vation. Flowering in January-September; fruiting in March-November. The species ranges from southern Nicaragua to western Panama.
Psychotria capacifolia is recognized by its short succulent stems, pubescence on younger parts, large leaves, axillary often large puberulent inflores- cences, well-developed (only slightly arcuate) sub- marginal vein, and the yellowish green fruit. The corolla interior is said to have a "ball" of hairs within (Dwyer, 1980, p. 354). This species closely resembles some material placed under P. macro- phylla, P. siggersiana, and P. aggregate, but the differences used in the keys appear to reflect a real discontinuity of gene flow.
Psychotria capitata Ruiz & Pav., Fl. Peruv. 2: 59, pi. 206. 1799.
Shrubs or small treelets l-3(-4) m tall, leafy stems 2- 4 mm thick, glabrous; stipules 8-18 mm long, 4-7 mm broad, ovate-lanceolate, acute at apex, with 2 triangular lobes 1-10 mm long, glabrous, deciduous. Leaves with petioles 4-20 mm long, 1-2 mm thick, glabrous, often drying yellowish green; leaf blades 6-16(-22) cm long, 2.5-6(-8) cm broad, elliptic-oblong to elliptic, ovate- elliptic or elliptic-lanceolate, apex short-acuminate with tip 5-12 mm long, base acute or obtuse and slightly decurrent on petiole, drying stiffly chartaceous, greenish, glabrous above and below (except for short hairs along the sides of the midvein beneath), 2° veins 9-14/side, arising at almost 90° from midvein. Inflorescences ter- minal and solitary, 5-1 3 cm long, 3-6 cm broad, an open racemose or congested panicle, white at anthesis, pe- duncles (l-)2.5-7 cm long, ca. 1.5 mm thick, glabrous (rarely puberulent), branches opposite or ternate (alter-
nate) and usually without subtending bracts, flowers ses- sile in distal cymose groupings of 2-4 subtended by white lanceolate bracteoles 4-9 mm long and 1-2 mm broad. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx ca. 0.5 mm long, calyx lobes 4-5, ca. 0.3 mm long, triangular; corolla white or cream (yellow in throat), tube 3-7 mm long, 1 mm diam. near base, lobes 4-5, 1.5-4 mm long; stamens 4 or 5 anthers 1-2 mm long. Fruits 5-7 mm long, ca. 5 mm diam., subglobose, dark blue- black to purple-black or black; pyrenes ribbed.
Plants of lowland evergreen forest formations, 0-200 m elevation. Flowering and fruiting throughout the year (mostly in December-July). The species ranges from Belize (Croat, 1978), the Caribbean lowlands of Nicaragua and Panama to Peru and Brazil; it is not known from Costa Rica.
Psychotria capitata is recognized by its white flowers and inflorescences, conspicuous bracts on the distal parts of the inflorescences, leaves usually glabrous and drying greenish, and the large bilobed stipules. This species is similar to P. calochlamys, which has larger bracts and stipules and less prom- inent secondary venation. Compare also Psycho- tria officinalis and Palicourea tilaranensis. This species appears to be less variable in Central America than in South America (cf. Steyermark, 1974).
Psychotria cartagoensis Nepokroeff, nom. nov. Cephaelis latistipula Standl., J. Wash. Acad. Sci.18: 281. 1928, not Psychotria latistipula Benth. Figure 13.
Herbaceous subshrubs, 0.3-1 m tall, stems erect and usually unbranched, leafy stems 2-6 mm thick, glabrous, quadrangular or terete; stipules (5-) 1 2-22 mm long, 1 2- 18 mm broad, ovate in general outline, bilobed with a narrow sinus 2-7 mm deep, glabrous, stiffand persisting. Leaves well separated along the stem, petioles 2-6(-7) cm long, 1.2-3.7 mm thick, glabrous, drying dark; leaf blades 13-27 cm long, 5-9 cm broad, elliptic-oblong, oblong-lanceolate, elliptic-obovate, or ovate-elliptic, apex abruptly narrowed or rounded and shortly acute or short- acuminate, base acute to cuneate, drying stiffly charta- ceous, dark gray-green above, much paler beneath, 2° veins 9-13/side, arising at 60-80° and arcuate distally. Infloresences axillary and sessile or subsessile, broadly capitate, 1-2 cm long and 2—4 cm broad, becoming dark magenta or purplish, bracts 7-8 mm long and 2-4 mm broad, innermost bracteoles lanceolate, flowers enclosed by many bracteoles. Flowers glabrous externally, hypan- thium ca. 2 mm long, turbinate, calyx cup 0.3-1 mm long, calyx lobes 4-5(-6), 1-4 mm long, lanceolate; co- rolla 4-5 mm long, funnelform, white, lobes 5, 1.2-3 mm long, acute; anthers 2.5 mm long. Fruits ca. 6 mm long and 4 mm broad (not including the 2 mm long persisting calyx), reddish purple.
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239
Understory plants of evergreen wet forests of the Caribbean slope, from 800 to 1 500 m eleva- tion. Flowering in March (Standley 39695 us the type of C. latistipuld) and May-June; fruiting in June and November. The species has only been collected in the valley of Rio Sarapiqui (Heredia Province) and near Orosi and Mufieco (Cartago) in central Costa Rica.
Psychotria cartagoensis is recognized by its short unbranched habit, broad bilobed stipules, axillary and subsessile heads, and absence of pubescence. Among our species, Hoffmannia congesta is most likely to be confused with this species, but that species lacks the involucrate bracts and has many- seeded fruits. This species and its allies have been studied by Molly Nepokroeff (wis, 1992); P. \vil- buriana Dwyer and P. dukei Dwyer of Panama are closely related.
Psychotria carthagenensis Jacq., Enum. PL Carib. 16. 1760. Figure 65.
Shrubs, subshrubs, or small treelets, 0.3-3(-4) m tall, leafy branches 1.5-4 mm thick, glabrous and terete, be- coming gray; stipules 3-8 mm long, 1.5-5 mm broad, ovate to oblong or slightly obovate, apex rounded or bluntly acute, glabrous, drying dark reddish brown, ca- ducous. Leaves with petioles 3-10 mm long, ca. 1 mm thick, with decurrent lamina base; leaf blades (4-)6- 13(-15) cm long, (1.7-)2-5(-7) cm broad, oblanceolate to narrowly elliptic-obovate or narrowly elliptic, apex obtuse to very shortly acuminate (tip ca. 4 mm long), base gradually narrowed and cuneate or acute, decurrent on petiole, drying chartaceous, gray or reddish gray, gla- brous on both surfaces, 2° veins 6-8/side, domatia rarely present in distal vein axils. Inflorescences terminal and solitary, 3-8(-10) cm long, 2.5-4 cm broad, a compact panicle with usually 3 nodes of opposite branches di- minishing in size toward apex (also verticillate or um- bellate), peduncle to 4.5(-6) cm long, 0.6-1 .3 mm thick, glabrous, bracts subtending the lateral branches 0.5-1.5 mm long, subulate to broadly ovate, flowers in distal cymes, pedicels 0-2 mm long, bracteoles to 0.5 mm long. Flowers glabrous or minutely (0.05 mm) papillate-pu- berulent externally, hypanthium ca. 1 mm long, calyx ca. 0.5 mm long, subentire or with 5 short triangular lobes; corolla white, funnelform-salverfbrm, tube 2.5-3 mm long, ca. 1 mm diam. near base, lobes 5, 1.2-2 mm long; anthers 1-1.5 mm long. Fruits ca. 5 mm long and 4 mm diam., oblong or ellipsoid, becoming red or or- ange, with 10 vertical ribs, glabrous; pyrenes with 2 sulci on the inner face and 5 ribs on the exterior surface.
Plants of shaded sites in seasonally dry decid- uous forest formations of Guanacaste and north- ernmost Alajuela Province, from 5 to 200(-600) m elevation (rarely to 1400 m elsewhere). Flow- ering in November-August (primarily February-
July); fruiting throughout the year. The species ranges from Mexico and the West Indies to Bolivia and Argentina.
Psychotria carthagenensis is recognized by its deciduous forest habitat, smaller oblanceolate or obovate leaves, lack of conspicuous pubescence, small flowers and inflorescences drying reddish, and inflorescences often with nodes bearing two larger and two smaller lateral branches. The ten- dency of the leaves to dry gray or reddish and the red or orange fruit are characteristics of members of subgenus Psychotria. The correct spelling of this species is carthagenensis (not carthaginensis}. This species is similar to P. tenuifolia of similar habitats but with bilobed stipules.
Psychotria chagrensis Standl., J. Wash. Acad. Sci. 15: 105. 1925. Figure 60.
Shrubs, 1-3 m tall, many-branched and often flat- topped, leafy branchlets 1-3 mm thick, glabrous, reddish brown or dark; stipules 3-10 mm long, 1 -3 mm broad near base (broader when subtending the inflorescence), narrowly tubular to narrowly ovate and acute at apex with 1 or 2 slender awns 1-3 mm long, glabrous and drying dark reddish brown, caducous. Leaves usually closely clustered distally, petioles 1-12(-17) mm long, ca. 0.7 mm thick, glabrous; leaf blades 2.5-8(-10) cm long, 0.9-3(-3.5) cm broad, obovate to oblanceo- late, elliptic-obovate or elliptic, apex acuminate or cau- date-acuminate to bluntly obtuse, tip ca. 5 mm long, base cuneate to acute and decurrent on petiole, drying stiffly chartaceous, gray or reddish brown, glabrous above and below, 2° veins 6-9/side (but sometimes difficult to see), joined by an arcuate submarginal vein near margin. Inflorescences terminal (rarely pseudoaxillary), 6-16 mm long, to 7 mm broad, sessile, subtended by broad stipule- like dark reddish brown glabrous bracts to 1 cm long, fasciculate or capitate, with few sessile flowers, bracteoles 4-5 mm long. Flowers distylous, glabrous externally, hy- panthium ca. 1 mm long, calyx 2-4 mm long, lobes 1- 2 mm long, narrowly triangular, corolla funnelform, white, tube 4—9 mm long and 1-1.3 m diam., lobes 5, 2-3 mm long; stamens 5, anthers ca. 1 m long. Fruits becoming 8 mm long (not including persisting calyx), 3-6 mm diam., oblong-ellipsoid to ovoid, persisting calyx 2-3 mm long, becoming red or purple pyrenes ca. 6 mm long, weakly ribbed.
Understory plants often found in low wet de- pressions in evergreen rain forest formations, from near sea level to 700 m elevation on the Caribbean slope and Osa Peninsula. Probably flowering and fruiting throughout the year (flowering mostly in March-August). At La Selva flowering is most common in May-June and fruiting in November- December. The species ranges from southern Nic- aragua to Colombia and Peru, but with disjunct
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populations in Veracruz, Mexico, and Izabal, Guatemala.
Psychotria chagrensis is recognized by its short (often flat-topped) habit, very small obovate leaves, distinctive stipules, small sessile fasciculate or cap- itate inflorescences, and unusual calyx lobes. The leaves drying grayish or reddish and the red fruit are characteristic of subgenus Psychotria. The slender secondary veins often have point inden- tations on the lower surface after the leaves have been dried. The very small leaves are unusual among the woody plants of the lowland rain forest floor; compare P. graciliflora, P. parvifolia, and Randia loniceroides.
from near sea level to 700 m elevation. Flowering in May-September and January; fruiting in July- March. The species ranges from southern Mexico and Belize to central Panama.
Psychotria chiapensis has the longest corolla tubes among Central American Psychotria species; it is pollinated by long-tongued sphingid moths (cf. Bawa & Beach, 1983). The terminal subcapi- tate inflorescences (often in groups of three) with broad bracts and the relatlively large four-edged fruit are also distinctive. The paucity of puberu- lence, and leaves usually broadest at or above the middle are additional characters that make this species stand apart. It is called cocobolito in Pan- ama.
Psychotria chiapensis Standl., Contr. U.S. Natl. Herb. 23: 1390. 1926. Cephaelis tetragona J. D. Smith, Bot. Gaz. 61: 376. 1916, not Psychotria tetragona Seem., 1867. Figure 18.
Psychotria chiriquiensis (Standl.) C. M. Taylor, comb. nov. Cephaelis chiriquiensis Standl., Ann. Missouri Bot. Gard. 28: 469. 1941. Figure 17.
Shrubs, treelets, or small trees, 2.5-8(-10) m tall, much- branched with rounded crown, leafy stems 2-6(-8) mm thick, terete and often conspicuously contracted for 5- 10 mm below the node after drying, glabrous or with short (0.2 mm) hairs in new growth, center with white spongy pith; stipules 2-5 mm long, 2— 4(-8) mm broad, broadly triangular, rounded with a small apical sinus, with 2 triangular acute lobes to 2 mm long. Leaves with petioles 7-35 mm long, 0.8-2.3 mm thick, glabrous (sometimes puberulent near inflorescences); leaf blades 9-20(-23) cm long, 3-8(-9) cm broad, elliptic-obovate to elliptic-oblong or narrowly oblong-obovate, apex short- acuminate or acute with tip 5-10 mm long, base usually gradually narrowed and cuneate base, slightly decurrent on petiole, drying thin-chartaceous and grayish green to dark brown above, distinctly paler beneath, glabrous above and below except for short (0.2 mm hairs) along sides of midvein beneath, 2° veins 9-12/side. Inflores- cences terminal, 1 (or 3 when basal lateral branches are axillary to distal leaves), 4-12 cm long, 5-8 cm broad, subcapitate or corymbiform with 3-7 dense bracteate flower clusters, peduncles l-8(-15) cm long, 1.3-3 mm thick, minutely puberulent with hairs 0.2 mm long, bracts 6-10 mm long, 4-8 mm broad, ovate to ovate-elliptic, obtuse to rounded, minutely ciliolate along margins, with 6-many flowers closely clustered and sessile. Flowers distylous, calyx resembling the bracts in texture and col- or, 3-5 mm long, calyx lobes 5, ca. 1.5 mm long, tri- angular; corolla salverform, white, glabrous externally in ours, tube 20-45 mm long, 1-3 mm diam. for most of its length, lobes 5, 9-15 mm long and 3 mm broad; anthers ca. 4 mm long. Fruits 12-16 mm long (not in- cluding the calyx) and 9-1 3 mm diam., ellipsoid to ovoid, purple-black, strongly 4-angled, persisting calyx 2-3 mm long; pyrenes 10-13 mm long, 7 mm broad and 3 mm thick, bony, with 3 dorsal ribs and concave areas be- tween.
Plants of evergreen lowland rain forest forma- tions on both the Caribbean and Pacific slopes,
Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla- brous; stipules 3-5 mm long, with a short (1 mm) tube and 4 rounded ovate lobes 2-4 mm long at each node, these separate on the internode but overlapping above petioles, persisting. Leaves well separated along stems, petioles 15-40(-57) mm long, 0.5-1.5 mm thick, gla- brous; leaf blades 8-16(-20) cm long, 3-7 (-9) cm broad, elliptic to elliptic-oblong, apex acuminate with tip 5-10 mm long, base obtuse to acute, drying stiffly chartaceous, dark green to yellowish brown or grayish brown, glabrous above and beneath, 2° veins 1 1-22/side, loop-connected distally to form an arcuate submarginal vein. Inflores- cences terminal (pseudoaxillary), solitary or 3, 3-12 cm long, 3-8 cm broad, capitulum often with 3 short (2-5 mm) primary branches (especially in fruit), involucral bracts 12-20 mm long, 20-50 mm broad, broadly ovate, purple to reddish violet, peduncle l-6(-8) cm long, 2- 3.5 mm thick, glabrous, flowers subsessile within many imbricate bracteoles 8- 12 x 5-1 Omm, elliptic to oblong. Flowers glabrous externally, hypanthium ca. 1 mm long, 0.5 mm diam., turbinate, calyx tube ca. 1 mm long, lobes ca. 0.5 mm long; corolla tubular-funnelform, purplish, tube 9-12 mm long and 1.7 mm diam., lobes 5-6, 1.5- 2 mm long; anthers to 2 mm long. Fruits elliptic, blue, enclosed within the inflorescence; pyrenes with 4-5 ridg-
Plants of lower montane rain forest formations, from 1 200 to 1 800 m elevation. Flowering in Jan- uary-September; fruiting in October-December. This species ranges from Volcan Tenorio to south- ward along the Caribbean slope and continental divide to the Chiriqui Highlands.
Psychotria chiriquiensis is recognized by its in- volucrate capitate inflorescences with broad pur- ple bracts, long-petiolate leaves with many prom- inent secondary veins, lack of pubescence, and rounded stipule lobes usually overlapping above
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the petiole. This species is named in honor of An- tonio Molina R., who monographed the Mexican and Central American species ofCephaelis in 1953 and has made many important collections in Cen- tral America. This species is very similar to P. data, with larger red bracts, heads not three- branched, and white corollas. It is also similar to P. dichroa.
Psychatria chiriquina Standl., Contr. U.S. Natl. Herb. 18: 129. 1916. Figure 61.
Shrubs or small trees, 2-6 m tall, leafy stems 1-4 mm thick, glabrous, dark reddish brown; stipules 4-8(-10) mm long, 3-5 mm broad, ovate to elliptic, apex acute, ciliolate to erose or shortly bilobed, united around stem at base, glabrous or puberulent, caducous. Leaves with petioles 7-18(-35) mm long (sometimes variable on the same stem), 0.4-1.3 mm broad, glabrous; leaf blade 5- 15 cm long, 2-6 cm broad, narrowly elliptic, elliptic- oblanceolate to oblanceolate or oblong-obovate, apex gradually tapering and acute or acuminate, tip ca. 10 mm long, gradually narrowed to an acute base and de- current on petiole, drying stiffly chartaceous, dark red- dish brown above, glabrous above and below except for hairs along sides of midvein, occasional pit domatia in vein axils beneath, 2° veins 6-12/side. Inflorescences terminal, usually solitary or 3, 3-10(-l 7) cm long, a few- branched panicle, often trichotomous, peduncles 1.5-5 cm long, 0.5-2 mm thick, essentially glabrous, pedicels 0-3 mm long. Flowers glabrous externally, calyx ca. 1 mm long, truncate or with 5 small (0.6 mm) lobes; corolla white, salverform or funnelform, tube 3-6 mm long, 1.5- 2 mm diam., glabrous or puberulent externally, lobes 5, 1.5-2 mm long; stamens 5, anthers ca. 1 mm long. Fruits 5-8 mm long, 4-6 mm diam., oblong to subglobose, becoming red, persisting calyx 0.5-1 mm long; pyrenes with 4—5 rounded ribs.
Psychotria chitariana Dwyer & C. Hamilton, Phy- tologia64: 221. 1988.
Subshrubs, 30-100 cm tall, leafy stems 4—9 mm thick, glabrous, drying dark; stipules 15-18 mm long, ovate, with 2 awns ca. 3 mm long, caducous. Leaves with pet- ioles 0-27 mm long, ca. 3 mm thick, glabrous, with lateral margins and flattened above; leaf blades 25-40 cm long, 10-17 cm broad, elliptic-obovate to obovate- oblong or oblanceolate, apex obtuse to acute, base grad- ually narrowed and cuneate or expanded and auriculate at the petiole, drying thinly chartaceous, grayish brown to reddish brown, glabrous above and below, 2° veins 12-17/side, loop-connected 2-3 mm from the margin. Inflorescences terminal and solitary, dense panicles of congested cymes, globose or hemispheric, 3-1 1 cm long, 3-5 cm broad, peduncles l-2.5(-^.5?) cm long, bracts ca. 5-6 x 1-3 mm, glabrous, flowers closely congested, pedicels 3-5 mm long, bracteoles to 0.5 mm long. Flow- ers glabrous externally, calyx ca. 1 mm long, cupulate, lobes 5, short; corolla funnelform, greenish white, tube 4. 5-5 mm long, 1-1.5 mm diam., lobes 1.5-2. 3 mm long, triangular or oblong; anthers ca. 1.5 mm long. Fruits not seen.
Known only from the Caribbean slope of the Cordillera de Talamanca in Costa Rica, from 200 to 800 m elevation. Flowering in April-May; im- mature fruits were collected in September. The species ranges from the valley of Rio Chitaria east of Turrialba (Liesner et al. 15400 CR, MO holotype) to the valley of Rio Estrella.
Psychotria chitariana is distinguished by its short stature, large obovate leaves with rounded and obtuse apex, and unusual inflorescence. The leaves drying dark grayish (or reddish brown) is a char- acteristic of subgenus Psychotria. Compare P. al- faroana with more open inflorescences and acute leaf bases.
Plants of evergreen montane cloud forest for- mations, from (900-)1500 to 2500 m elevation. Flowering in January-May (July-August in Nic- aragua); fruiting in August-April (November-May in Nicaragua). The species is known from northern and central Nicaragua and from the easternmost part of the Cordillera de Talamanca in Costa Rica and the Chiriqui Highlands in Panama.
Psychotria chiriquina is recognized by its higher elevation habitats, stipules often with two small distal lobes, narrow leaves tapering gradually to apex and base, pit domatia, small flowers, and poorly developed calyx lobes. The red fruit and leaves/branchlets drying dark reddish brown are characters of subgenus Psychotria. Compare P. sarapiquensis, with a shorter calyx, and P. sylvi- vaga. This species is also similar to P. panamensis and with very different stipules.
Psychotria cincta Standl., Publ. Field Columb. Mus., Hot. Ser. 7: 90. 1930. Figure 58.
Shrubs, l-2.5(-4) m tall, leafy branches 1.5-4 mm thick, glabrous and drying greenish or dark; stipules forming a small basal (1-3 mm) sheath, with 2 awns (4/ node), awns 6-14 mm long, ca. 0.5 mm broad, glabrous, persisting. Leaves with petioles 8-20 mm long, 1-1.5 mm thick, glabrous; leaf blades 12. 5-21 (-28) cm long, 3. 5-7 (-9) cm broad, elliptic-oblong to oblong or ovate- oblong, apex tapering gradually and acuminate, tip 8- 18 mm long, base acute to obtuse, drying stiffly char- taceous, green or grayish green, glabrous above and be- low, 2° veins 9-1 1 (- 1 4)/side, leaf margin with a distinctly thickened vein along the edge. Inflorescences terminal and solitary, usually pendant, 12-20 cm long, 2-4 cm broad, racemiform thyrsoid panicles with short (1-2 cm) usually alternate lateral cymose branches, peduncles 6- 14 cm long, 0.6-1.2 mm thick, sparsely and minutely puberulent with short (0.1-0.2 mm) thin hairs, bracts
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linear, adnate to lateral branches or absent, bracteole 0- 0.5 mm long, pedicels 0-2 mm long. Flowers minutely puberulent externally, hypanthium 0.5 mm long, calyx 0.2-0.4 mm long, truncate or with minute lobes; corolla funnelform or salverform, white or greenish white, tube 2-3 mm long, ca. 0.5 mm diam., lobes 5, 1-1.5 mm long. Fruits 4-6 mm long, subglobose, obscurely ribbed, gla- brous, red becoming black; pyrenes shallowly ribbed.
Plants of evergreen lowland Caribbean rain for- est formations, from near sea level to 500 m ele- vation. Flowering in February-July; fruiting in August. The species ranges from Nicaragua to Co- lombia.
Psychotria cincta is recognized by its persisting stipules with long narrow awns, the often oblong leaves usually drying yellowish green, the long nar- row pendant inflorescences with unusually short lateral branches, and the small puberulent flowers. In addition, the thickened leaf margins are nota- ble. This distinctive species is presently known only from La Selva and nearby areas in Costa Rica. Compare P. deflexa, which lacks the thickened leaf margins and has pyrenes with transverse ribbing.
Psychotria clivorum Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 87. 1944, based on P. limonensis var. angustifolia Standl., loc. cit. 17: 282. 1937. Figure 61.
Shrubs or small trees, 1.5-6 m tall, leafy stems 2-4 mm thick, glabrous; stipules 10-16 mm long, 2-5 mm broad (broader beneath the inflorescences), with a sheath 1-4 mm long, ovate to elliptic, usually bilobed with 2 acute lobes and a short (1-3 mm) sinus at apex, glabrous, dark reddish brown, caducous. Leaves variable in shape (on different plants), petioles 6-35 mm long, 1-1.7 mm thick, glabrous; leaf blades 10-22 cm long, 3.5-8 cm broad, narrowly elliptic-oblong to elliptic or oblanceo- late, apex acuminate, tip 7-15 mm long, base acute to cuneate and often long-decurrent on petiole, drying membranaceous to thinly chartaceous, grayish or gray tinted with red, glabrous above and below (midvein rare- ly sparsely puberulent beneath), 2° veins 8-12/side, loop connected 1-3 mm from margin, small pit domatia usu- ally present. Inflorescences terminal and solitary, 6-10 cm long, 3-7 cm broad, open panicles, usually with 4 lateral branches from 2 proximal nodes, peduncles 1-5 cm long and 2.2 mm thick, minutely puberulent or gla- brous, reddish brown, bracts to 2(-3) mm long, pedicels 0-2 mm long. Flowers glabrous externally, hypanthium ca. 0.6 mm long, conical, calyx tube ca. 0.5 mm long with minute (0.2 mm) broadly triangular lobes; corolla funnelform, white, tube 2-2.5 mm long, 1.2-1.5 mm diam., lobes 1-2 mm long; anthers ca. 1 mm long. Fruits 5-6 mm long, 4-5 mm diam., ellipsoid or obovoid, lon- gitudinally ribbed, red; pyrenes with 3-5 longitudinal ridges.
Plants of evergreen forest formations of the Ca- ribbean slope; from near sea level to 800 m ele- vation (to 1400 m in Guatemala). Most flowering collections have been made in March-May. A lit- tle-collected species ranging from Veracruz, Mex- ico, to central Panama.
Psychotria clivorum is recognized by its larger narrow leaves gradually narrowed to the base with numerous pit domatia and distinct submarginal vein, stipules tubular at base and sometimes bi- lobed distally, inflorescences with whorled lateral branches at first two nodes (usually of two unequal pairs), small flowers, and reddish fruit. The ten- dency of the leaves to dry grayish and the stipules to dry reddish is characteristic of subgenus Psy- chotria. Specimens of this species may be difficult to separate from material of P. orosiana and P. sarapiquensis.
Psychotria cocosensis C. Hamilton, Phytologia 64: 222. 1988. Figure 62.
Shrubs or small trees to 5 m tall, leafy stems ca. 2-7 mm thick, glabrous; stipules (7-) 1 2-35 mm long, ( 1 .5-)2- 5 mm broad, lanceolate, glabrous, drying dark reddish brown, caducous. Leaves with petioles 5-16(-23) mm long, 1-2.5 mm thick, glabrous; leaf blades (8-) 12-1 7 cm long, (3-)5-7 cm broad, narrowly elliptic to elliptic- oblong or broadly elliptic-obovate, apex acuminate or caudate-acuminate with tip 4-12 mm long, base acute to attenuate and decurrent on petiole, drying subcoria- ceous, reddish brown to grayish brown, glabrous above and beneath, 2° veins 9-1 2/side, small pit domatia often present in the vein axils beneath. Inflorescences terminal or pseudoaxillary, solitary or 3-parted, 1.3-3 cm long, few-branched panicles with distal 3-flowered cymules, peduncles 0-7 mm long, ca. 1 mm thick, glabrous, bracts 0.5-1.5 mm long, triangular, glabrous, pedicels 2-3 mm long. Flowers puberulent externally, hypanthium ca. 0.5 mm long, glabrous, calyx tube ca. 1 mm long and 2 mm broad, cupulate, slightly denatate; corolla funnelform, white, tube ca. 3 mm long and 2 mm diam., lobes 5, ca. 4 mm long, 2 mm wide; stamens 5, anthers ca. 2 mm long. Fruits ca. 6 mm long, 5-6 mm diam., ellipsoid to globose, orange then red and finally purple, drying dark reddish brown, persisting calyx 0.8-1.5 m long.
Plants known only from Cocos Island, from near sea level to 50 m elevation. We have seen four collections: Foster 4132 F, Holdridge 5169 us, Pit- tier 12375 us (the holotype), and Pittier 16279 GH, us. Flowering in April-June; fruiting in January, April, and June.
Psychotria cocosensis is similar to P. panamen- sis but differs in its subcoriaceous leaves and larger fruits with persisting cupulate calyx. The very small domatia, short inflorescences, and flowers with long
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corolla lobes are distinctive. The red fruit and ten- dency of the leaves to dry reddish brown are char- acteristics of subgenus Psychotria. The Holdridge collection is unusual in having broadly elliptic- obovate leaves that have dried grayish and sig- nificantly paler beneath.
Psychotria cooperi Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 296. 1929. Figure 13.
Small trees or shrubs, 3-8 m tall, to 10 cm dbh, leafy stems 1 .4-5 thick, glabrous, terete or slightly tetragonal; stipules 5-9(-l 1) mm long, to 8 mm broad, with a broad tube, truncate to rounded or bilobed with a small ( 1 mm) sinus at apex, glabrous, persisting or deciduous. Leaves opposite or 3 at a node, petioles 7-25 mm long, 0.7-1.7 mm thick, glabrous; leaf blades 8-21 cm long, 3.5-7 cm broad, elliptic-oblong to elliptic or elliptic-obovate, apex acuminate with tip 3-10 mm long, base gradually nar- rowed and acute or cuneate base, slightly decurrent on petiole, drying chartaceous, dark yellowish green to dark brown above, distinctly paler beneath, glabrous except for short (0.2-0.3 mm) stiff hairs along the sides of the midvein above and below (sometimes with thin hairs along the secondary veins beneath), 2° veins 7-1 I/side, arcuate-ascending near the margin. Inflorescences ter- minal or pseudoaxillary and becoming verticilate, sessile or subsessile, 1-3.5 cm long, to 3.5 cm broad, a compact globose irregularly branched panicle, peduncle 1-7 mm long, bracts to 10 mm long, bracteoles ca. 1 mm long and digitate-glandular, pedicels to 6 mm long in fruit. Flowers glabrous externally, distylous, hypanthium ca. 0.7 mm long, calyx lobes 5, ca. 1 mm long; corolla fun- nelform, white, tube 4-5 mm long, 1.3 mm diam., lobes reflexed at anthesis, ca. 2 mm long. Fruits 4-5(-10?) mm long, 2.5-3.5 mm diam., obovoid or turbinate and trun- cated distally, dark blue or purple, persisting calyx ca. 1 mm long; pyrenes 3-4 mm long, with 4-5 longitudinal ribs.
Plants of lowland rain forest formations from 20 to 300(-600) m elevation, often found in light gaps and stream sides. Flowering and fruiting throughout the year with most collections from January-March. The species ranges from north- eastern Costa Rica to Colombia.
Psychotria cooperi is recognized by its small con- gested axillary inflorescences (sometimes appear- ing verticillate), the broad short-tubular stipules, leaves often drying yellowish green, small white flowers, and blue or purple obovoid fruit. The flowers are mostly sessile in terminal inflores- cences but become axillary as the stems continue to grow. These small treelets are called cocobolito in Bocas del Toro, Panama. Fruiting material may resemble Palicourea copensis.
Psychotria correae (Dwyer & Hay den) C. M. Tay- lor, comb. nov. Cephaelis correae Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 36. 1 968. Figure 17.
Shrubs or small trees, 2-3.5(-5) m tall, leafy stems 3- 6 mm thick, glabrous, quadrangular; stipules with a very short (1-3 mm) sheath with 2 broadly rounded often overlapping lobes ca. 4 mm long and 4 mm broad sep- arated by a U-shaped sinus, glabrous. Leaves with pet- ioles 1 5-80 mm long (opposing petioles often unequal), 1.2-2.5 mm thick, glabrous; leaf blades 1 1-28 cm long, 5-13 cm broad, broadly elliptic-oblong, ovate-oblong, apex abruptly narrowed and short-acuminate with tip ca. 4 mm long, base broadly obtuse, drying chartaceous, green, glabrous above and below, 2° veins 13-17(-20)/ side. Inflorescences solitary and terminal or pseudoax- illary, capitulae 1-2 cm long, 2-3 cm diam., involucre 5-10 cm wide, peduncles (3-)5-12(-22) cm long, ca. 2 mm thick, glabrous, erect and becoming pendant, the 2 large involucral bracts 2.5-5 cm long and 4-5 cm broad, ovate to reniform, rose red to magenta or purple, gla- brous, bracteoles 8-15 mm long, obovate. Flowers gla- brous externally, calyx lobes 5, ca. 1 mm long; corolla narrowly funnelform, white, tube to 10 mm long, cylin- drical, lobes 5, ca. 3 mm long; anthers ca. 3 mm long. Fruits 10-15 mm long, ca. 6 mm diam., obovoid-oblong, blue; pyrenes with 4-5 slightly elevated ridges
Plants of evergreen cloud forest formations along the central Cordilleras at 900-1100 m elevation. Flowering in March-August; fruiting in October. This species is only known from the Cordilleras de Guanacaste and Tilaran and the Province of Code in Panama.
Psychotria correae is recognized by the larger leaves with many secondary veins, the unusual stipules, the long-pedunculate inflorescences be- coming pendant in later stages, and the large pur- plish bracts subtending the capitulum. Specimens of this species can be confused with P. elata, which has erect capitulae with smaller involucral bracts and shorter peduncles, and to P. chiriquiensis, which lacks involucral bracts. This attractive spe- cies was named in honor of our respected Pana- manian colleague Mireya Correa.
Psychotria deflexa DC., Prodr. 4: 510. 1830. P. patens auct. non Sw. fide Steyermark 1972. Fig- ure 56.
Shrubs, (0.5-)l-3 m tall, leafy stems 0.5-5 mm thick, glabrous or sparsely puberulent in early stages; stipules united around stem for 0.5-2 mm and with 2 subulate or linear awns 3-8 long on each side, glabrous, persisting. Leaves with petioles 3-9(-12) mm long, 0.5-1.3 mm thick, glabrous; leaf blades 7-15(-18) cm long, (1.5-)2-
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5(-7) cm broad, narrowly ovate-elliptic to narrowly el- liptic, elliptic-lanceolate or rarely lanceolate, apex ta- pering gradually and acute or acuminate, tip 10-20 mm long, base broadly obtuse to acute, drying chartaceous, greenish or brown, glabrous above and below, 2° veins 4-8/side. Inflorescences terminal (rarely pseudoaxillary) and solitary, 4-1 1 cm long and 2-4 cm broad, narrowly pyramidal or thyrsiform panicles with slender central rachis and short (5-14 mm) opposite or subopposite lateral branches (expanding somewhat in fruit), pedun- cles 2.5-5 cm long, 0.4-1.1 mm thick, glabrous or mi- nutely puberulent, lateral branches without subtending bracts, pedicels 1-2 mm long, bracteoles ca. 1 mm long. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx ca. 0.5 mm long, dentate; corolla white, funnel- form, tube 1.6-3 mm long, 0.3-1 mm diam. (in Costa Rica), lobes 4-5, ca. 1 mm long; stamens 4, anthers ca. 1 mm long. Fruits 2-3 mm long and 3-4 mm diam. (8- 10 mm when aerenchymatous), oblate and somewhat bilobed, longitudinally ridged, becoming purple, violet, or blue (pulpy white within); pyrenes with 3-4 ridges and transverse ribs.
sheath, with 2 short (1-2 mm) obtuse or rounded lobes separated by a U-shaped sinus, glabrous. Leaves with petioles 4-25 mm long, 0.7-1 .2 thick, glabrous; leaf blades 4-1 1(-13) cm long, 1.8-4.3 cm broad, narrowly elliptic to lanceolate, apex tapering gradually and acuminate, tip 5-10 mm long, base acute and slightly decurrent on pet- iole, drying subcoriaceous, dark brownish green above, glabrous above and below, 2° veins (6-)8-12/side. Inflo- rescences 1-3 and terminal, 3-10 cm long, 3-8 cm wide, usually subtended by a pair of smaller (15 mm) leaves, capitula ca. 1 5-25 mm long and equally broad (enlarging in fruit), subtended by 2 involucrate bracts 12-20 mm long and 8-15 mm broad, white becoming pinkish (dark purple in fruit), peduncles 3-9 cm long and 1.5-3.5 mm thick, glabrous, flowers sessile within bracteoles. Flowers glabrous externally, hypanthium ca. 1.5 mm long, calyx ca. 3 mm long, lobes ca. 1.5 mm long and triangular; corolla funnelform, white to pink, tube 5-10 mm long, ca. 2 mm diam., corolla lobes 2-3 mm long, bluntly acute. Fruits 8-9 mm long (including the 1 mm high calyx), 4 mm diam., narrowly obovoid, blue-black, dry- ing dark reddish brown; pyrenes usually smooth.
Understory plants of wet evergreen or partly deciduous forest formations from ca. 500 to 1200 m elevation on the Caribbean slope of the cordil- leras and near sea level on the Osa Peninsula. Flowering in June-August; fruiting in July-March. This species ranges from Mexico to Peru and Bo- livia.
Psychotria deflexa is recognized by its persisting narrowly awned stipules, narrow leaves, general lack of pubescence (only the slender narrow inflo- rescence is minutely puberulent), small flowers, and purple or blue fruit with transverse ribs. Flow- ers in Costa Rican material appear to be much smaller than those described by Steyermark (1974, p. 1284). This species may resemble P. cincta and P. microbotrys. There are collections (Kernan & Phillips 661 & 789 CR) from Corcovado National Park that have unusually thin pedicels and inflo- rescences. However, these aberrant samples are bridged by other collections (Liesner 2842 & 3198 CR, also from the Osa Peninsula) that appear to be intermediate with the more normal inflores- cences. These populations are worthy of further study.
Psychotria dichroa (Standl.) C. M. Taylor, comb, nov. Evea dichroa Standl., Contr. U.S. Natl. Herb. 18: 124. 1916. Cephaelis dichroa (Standl.) Standl. Publ. Field Columb. Mus., Bot. Ser. 4: 296. 1929.
Small shrubs, 0.6-2 m tall, leafy stems 1.3-4 mm thick, glabrous; stipules 2.5-6 mm long, with a short (2-3 mm)
Plants of evergreen montane forest formations, 1 200-2600 m elevation. Flowering January-Sep- tember. This species is only known from the Chi- riqui Highlands of western Panama and the Ca- ribbean slope of the Cordillera de Talamanca.
Psychotria dichroa is recognized by the leaves with prominent 2° veins, one to three terminal small involucrate heads, white flowers, and high- land habitat. In Costa Rica, it is rarely collected and appears to be restricted to the understory of high-elevation (2000-2500 m) Quercus forests. This species is similar to P. chiriquiensis with red- dish purple capitula.
Psychotria domingensisJacq.,Enum. PI. 16. 1760. Psychotria pavetta Sw., Prodr. 45. 1788. Pali- courea domingensis (Jacq.) DC., Prodr. 4: 529. 1 830. Palicourea pavetta (Sw.) DC., loc. cit. 525. 1830. Psychotria mombachensis Standl., Publ. Field Columb. Mus., Bot. Ser. 8: 188. 1930.
Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla- brous, often with a shrunken area below the node after drying; stipules 2-5 mm long, with 2 triangular lobes separate almost to base by a broad (1-2 mm) U-shaped sinus, acute or awned, glabrous, deciduous or persisting. Leaves with petioles 5-20 mm long, 0.7-1.7 mm thick, glabrous, often drying yellowish; leaf blades 8-19 cm long, 3-7 cm broad, elliptic-oblong to narrowly elliptic- obovate, apex acuminate or gradually tapering and acute, base acute to obtuse and slightly decurrent on petiole, drying membranaceous to thin-chartaceous, yellowish green to greenish brown or brown, glabrous above and below, 2° veins 8-1 I/side. Inflorescences solitary or 3, terminal or pseudoaxillary, 4-9 cm long, 3-6 cm broad,
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a rounded open panicle with opposite branching, pe- duncle 3-20 mm long, ca. 1.5 mm thick and yellowish when dried, glabrous or minutely papillate-puberulent, bracts absent or 1-6 mm long and linear, flowers in distal cymules of 3, pedicels 0-3 mm long, ca. 1 mm thick, bracteoles minute or absent. Flowers glabrous or mi- nutely puberulent externally, hypanthium ca. 1.2 mm long, calyx tube ca. 0.5 mm long, lobes 0.2-1 mm long, narrowly acute; corolla white, tubular-salverform, tube ca. 12 mm long, 0.8-1.7 mm diam., lobes ca. 4-5 mm long and 2 mm broad. Fruits 4-5 mm long, ca. 6 mm diam., subglobose to ellipsoid, purple-black, with 4-5 longitudinal costae and smaller transverse ribs when dried; pyrenes angled.
Plants of evergreen or seasonally deciduous for- mations, from 20 to 900 m elevation. Flowering in April and June (the Nicaraguan type of P. mom- bachensis (Maxon et al. 7818 F) was flowering in July); a Costa Rican collection from near Tilaran (Standley & Valeria 45222 us) was fruiting in Jan- uary. The species is frequent in the West Indies and Guatemala-southern Mexico but is known from only a few collections in southern Central America.
Psychotria domingensis is recognized by the in- florescences with thick branches and relatively few flowers, larger fruit with longitudinal and trans- verse ridges, thin glabrous leaves that often dry yellowish green, two-lobed stipules, and dried stems often conspicuously contracted beneath the node. The few collections from Costa Rica come from both evergreen and deciduous areas. This species is rather similar to P. eurycarpa of wet forests, but that species has stiffer leaves lustrous above, poor- ly developed calyx lobes, and larger fruit without transverse ribs. It is also similar to P. microdon and some species of Coussarea (see fig. 47).
Psychotria dukei Dwyer, Ann. Missouri Bot. Gard. 67: 371. 1980.
Shrubs or herbaceous subshrubs, 0.5-1. 5(-2) m tall, leafy stems succulent in life, 1.8-7 mm thick (dried), glabrous; stipules 7-20 mm long, 8-1 8 mm broad, ovate- triangular, bilobed or fimbriate, yellowish and membra- naceous, lobes 1-4 mm long, deciduous. Leaves with petioles 1-5 cm long, 1-2.3 mm broad, glabrous; leaf blades (1 1-) 13-27 cm long, 4-1 1 cm broad, elliptic-ob- long to oblong or elliptic-ovate, apex short-acuminate with tip 5-10 mm long, base obtuse to cuneate and de- current on petiole, drying chartaceous, dark green or greenish brown, glabrous above, minutely (0. 1 mm) pu- berulent along the major veins beneath, 2° veins 10-18/ side. Inflorescences axillary, solitary at each node, 2.5- 9 cm long, pyramidal, often with the flowers crowded in 3 capitula on 3 primary branches 3-20 mm long (rarely with further secondary branches), peduncles 8-35(-60)
mm long, ca. 1.2 mm thick, minutely puberulent or gla- brous, bracts 2-10 mm long, triangular to lanceolate, bracteoles 2-5 mm long, often forming a small involucre beneath the sessile flowers. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx tube ca. 0.5 mm long, entire; corolla tubular-funnelform, white, tube 3-4 mm long, ca. 0.5 mm diam. basally, lobes ca. 1.5 mm long. Fruits 5-6 mm long, 3-4 mm diam., oblong, white; py- renes longitudinally ridged.
Plants of lower montane rain forest formations from (800-)1 100 to 1800 m elevation. Flowering in March and June-August; fruiting in March- November. The species ranges from the Cordillera de Tilaran to Choco, Colombia.
Psychotria dukei is recognized by its usually un- branched succulent stems, axillary inflorescences with flowers in capitate clusters, white fruit, and unusual stipules. This species is easily confused with P. aggregata or P. macrophylla, but the large thin stipules and inflorescence bracts are distinc- tive. This species is rarely collected in Costa Rica. Compare P. dukei with P. wilburiana Dwyer of Panama.
Psychotria elata (Sw.) Hammel, Selbyana 1 2: 1 39. 1991. Cephaelis elata Sw., Prodr. 45. 1788. C. punicea Vahl, Eclog. Amer. 1:19. 1796. C. cos- taricensis Schlechtend., Linnaea 28: 546. 1856. Evea elata (Sw.) Standl., Contr. U.S. Natl. Herb. 18: 123. 1916. C. elata forma lutea Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1279. 1938. Figure 17.
Shrubs, subshrubs, or small trees, 0.5-5(-8) m tall, leafy stems 1.3-5 mm thick, glabrous, terete or qua- drangular; stipules united around stem for 1-2 mm, with 2 ovate lobes on each side, 2-5 mm long and ca. 3 mm broad, obtuse to broadly rounded and separated by a U- or V-shaped sinus, glabrous, persisting. Leaves with pet- ioles 4-22(-30) mm long, 1-2 mm wide, glabrous; leaf blades 6-25 cm long, 2.5-7(-8.5) cm broad, oblong to elliptic-oblong, oblong-obovate, or oblanceolate, apex acute to short-acuminate, tip 4-10 mm long, base cu- neate to acute or obtuse, drying thinly to stiffly charta- ceous, green, glabrous above and below, 2° veins (9-)l 3- 20(-23)/side. Inflorescences terminal and solitary (rarely 2-3), capitate and globose-hemispherical, capitula 1.5- 4 cm long, involucre 2-7 cm broad, peduncles 2-13 cm long, 1.5-3 mm thick, erect or becoming pendant, gla- brous, the 2 involucrate basal bracts 15-55 mm long, 1 5-45 mm broad, deep red to orange-red (yellow), ovate to reniform, rounded to acuminate, glabrous, bracteoles 5-10 mm long, flowers sessile. Flowers glabrous exter- nally (rarely puberulent), calyx ca. 1 mm long, dentate; corolla white (rarely pink or yellow), narrowly funnel- form, tube ca. 16 mm long, 1.5-2.5 mm diam., lobes 5, 2.5-4 mm long, narrowly triangular; stamens 5, anthers 2-3 mm long. Fruits 5-10 mm long, 2-5 mm diam.,
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ovoid, blue becoming black; pyrenes 4-7 mm long and 3-4 mm broad, 5-ribbed.
Common shrubs of evergreen forest edges and forest interiors, from 30 to 1700 m elevation. Flowering and fruiting throughout the year (pri- marily flowering in January-August). Fruiting at La Selva in July-September. The species ranges from Mexico and the West Indies, through Central America to Colombia.
Psychotria elata is recognized by its glabrous parts, usually narrowly oblong leaves with many secondary veins, solitary long-pedunculate heads, large brilliant red (rarely yellow) bracts subtending the capitulum, and white corollas. The yellow- bracted form appears to be most common in the General Valley and Golfo Dulce area. This is one of Central America's most distinctive species of Rubiaceae, common in light gaps and on the edges of forests. The flowers are visited by butterflies and hummingbirds (Freeman & Stiles, 1990); breeding biology was studied by Bawa and Beach (1983). This is one of the species that made Ce- phaelis so distinctive a genus in Central America. Compare P. poeppigiana with conspicuous pubes- cence, P. chiriquiensis with purple capitula lacking the large involucral bracts, P. correae with longer peduncles and larger bracts, and P. dichroa with several smaller heads.
Psychotria emetica L.f, Suppl. PI. 144. 1781. Fig- ure 13.
Small herbs and subshrubs, 0.2-1 m tall, rhizomatous, stems usually unbranched, terete, leafy internodes 1-4 mm thick, appressed-puberulent with stiff crooked brownish hairs 0.2-0.4 mm long; stipules 2-4 mm long, 1-2 mm broad at base, narrowly triangular and ap- pressed-puberulent, drying dark, usually caducous. Leaves with petioles 3-12(-20) mm long, 1-2 mm broad, ap- pressed-puberulent with stiff brownish hairs; leaf blades (7-)8.5-13(-17) cm long, 2-5(-6.5) cm broad, elliptic to narrowly elliptic-oblong, elliptic-obovate or oblanceo- late, apex acute or short-acuminate, tip ca. 5 mm long, base cuneate to acute and slightly decurrent on petiole, drying thinly chartaceous, dark grayish green or dark grayish brown, glabrous above, sparsely appressed-pu- berulent with short (0. 1-0.3 mm) hairs beneath, 2° veins 5-9/side. Inflorescences axillary, usually solitary in each axil (2/node), 1-2 cm long, condensed cymose or sub- capitate, with 3-10 flowers, peduncles 3-12 mm long, ca. 0.7 mm thick, puberulent, bracts 1-2 mm long, tri- angular-acute, pedicels 0-5 mm long. Flowers with hy- panthium ca. 1 mm long, conical and appressed-puber- ulent, calyx lobes 1-1.5 mm long, narrowly triangular; corolla white, funnelform, glabrous externally, tube 2-4 mm long, 1-1.5 mm diam. at mouth, lobes 5, 1.5-2 mm long, acute. Fruits 8-10 mm long and 4-6 mm diam.,
oblong or ellipsoid, persisting calyx 1-1.5 mm long, be- coming blue; pyrenes ca. 5 mm long, smooth.
Plants of shaded understory in lowland ever- green rain forest formations, from 10 to 300 m elevation. In Costa Rica and Panama flowering in February-August; fruiting mostly in June-No- vember. This species ranges from Guatemala to Bolivia.
Psychotria emetica is recognized by its short, usually unbranched stems, small condensed axil- lary inflorescences, and bright blue fruit. These plants are occasionally cultivated and have been called raicilla and raicilla macho in Costa Rica, Nicaragua, and Panama. This species differs from the closely similar P. erecta by its smaller stature, puberulent stems, and floral details. The roots con- tain the drug ipecac but it is of inferior quality to that found in P. ipecacuanha.
Psychotria erecta (Aubl.) Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 24. 1943. Ronabea erecta Aubl., Hist. pi. Guiane 1: 156. 1775. Figure 13.
Shrubs or treelets, (0.5-)l-3(-8) m tall, leafy stems 1 .5-7 mm thick, sparsely appressed-puberulent with thin ascending hairs ca. 0.3 mm long, becoming glabrescent and drying dark; stipules with a persistent short (0.5 mm) base and single slender awn 2-6 mm long, ca. 0.5 mm wide, triangular to subulate, appressed-puberulent. Leaves with petioles 8-18(-35) mm long, 0.7-2(-2.5) mm thick, appressed-puberulent with hairs 0.2-0.3 mm long; leaf blades 8-20 cm long, 3-9 cm broad, elliptic-oblong to oblong or slightly obovate, apex abruptly narrowed and short-acuminate, tip 4-1 1 mm long, base obtuse to acute, drying chartaceous to subcoriaceous, dark olive green or brownish green, glabrous above, glabrous or sparsely pu- berulent beneath with thin appressed hairs, 2° veins 5- 8/side. Inflorescences 1-3 in each axil (2-6/node), 1-3 cm long, 5-10 mm broad, subcapitate condensed cymose with 3-7 closely crowded flowers, peduncles 0-8(-18) mm long, ca. 0.7 mm thick, appressed-puberulent with ascending yellowish hairs, flowers subsessile or sessile, bracteoles to 1 mm long. Flowers with hypanthium 1-2 mm long, sparsely appressed puberulent near the base, calyx ca. 1 mm long, glabrous, lobes ca. 0.2 mm high; corolla white, tubular-salverform, glabrous, tube 3^ mm long, 1 mm diam., glabrous externally, lobes 5(-6), 1.5- 3 mm long. Fruits 8-10 mm long, 5-8 mm diam., ellip- soid or oblong, glabrous, becoming blue-black; pyrenes 5-7 mm long, smooth.
Plants of evergreen rain forest formations, from near sea level to 700(-1000) m elevation on both the Caribbean and Pacific coasts. At La Selva this species is usually found on ridges near light gaps. Flowering in May-October; probably fruiting
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
247
throughout the year. This species ranges from Mexico and the West Indies to Bolivia.
Psychotria erecta is recognized by its axillary small subcapitate inflorescences with few small flowers, blue fruit, and oblong leaves often drying dark. This species is similar to P. emetica but grows to a larger size with broader and thicker leaves and glabrescent stems. Both species have axillary, not pseudoaxillary, inflorescences. Compare Appunia guatemalensis.
Psychotria eurycarpa Standl., J. Wash. Acad. Sci. 18: 275. 1928. Figure 47.
Shrubs or small trees, 1.5-5(-7) m tall, leafy stems 1- 3 mm thick, glabrous or rarely minutely papillate-pu- berulent; stipules 1-3 mm high and 3-4 mm broad, trun- cate to broadly rounded with a small (0.2 mm deep) sinus, persisting or deciduous. Leaves with petioles 12- 28(-35) mm long, 0.7-1.5 mm thick, glabrous and often drying yellowish; leaf blades 6-14(-21) cm long, 2.7- 7(-l 1) cm broad, ovate-elliptic to elliptic or broadly el- liptic, apex short-acuminate with a narrowed tip 3-10 mm long, base obtuse to acute, drying stiffly chartaceous, yellowish green, glabrous and lustrous above, glabrous beneath, 2° veins 4-7/side. Inflorescences terminal and solitary, 5-1 1 cm long, 5-9 cm broad, broadly pyramidal panicles with opposite branching or umbellate, the first node often with 4 lateral branches, peduncles 1 5-55 mm long, 1-2 mm thick, yellowish, glabrous, bracts ca. 1 mm long, flowers sessile or subsessile in glomerules of 2-5. Flowers nocturnal, glabrous or very minutely papillate- puberulent externally, hypanthium ca. 1 mm long and 0.6 mm diam., obconic, calyx tube 0.3-0.7 mm high with minute (0.2 mm) lobes; corolla narrowly funnel- form, white, tube 6-1 5 mm long and 0.8 mm diam. (1.7 mm distally), lobes 5 (6, 7), 3-8 mm long, 0.5-1 mm broad, narrowly oblong. Fruits 8-20 mm long (to 25 mm when succulent), 7-1 5 mm diam., oblong to subglobose, blue-black or dark purple (often drying pale yellowish brown), longitudinal ridges present but not prominent when dried; pyrenes 12-17 mm long, obscurely 3-5- angled.
Plants of lowland rain forest formations and lower montane cloud forest formations, from 20 to 900(-1100) m elevation. Flowering in Febru- ary-May; fruiting in October-April and June. This species is only known from the Caribbean coastal plain, Cordillera de Guanacaste, Cordillera de Ti- laran, and the Caribbean slopes of the Central Highlands as far east as Rio Reventazon, and in western Panama.
Psychotria eurycarpa is recognized by the usu- ally glabrous parts often drying yellowish green, the short stipules (truncated or with minute apical sinus), few-branched inflorescences with thick pe- duncles and thick lateral branches and inconspic- uous bracts, longer corolla tubes, and the large
fruit. Breeding biology was studied by Bawa and Beach ( 1 983). A collection (Taylor 3546) from Las Cruces (San Vito) is tentatively placed here; it is the highest elevation ( 1 1 00 m) and easternmost collection in Costa Rica. Folsom 9188 DUKE, from La Selva with corolla tubes to 18 mm long and corolla lobes to 1 2 mm long, is provisionally placed here. Individuals of this species may be difficult to distinguish from P. domingensis (with bilobed stipules and flowers in distal triads) and Coussarea psychotrioides (q.v.).
Psychotria fruticetorum Standl., J. Arnold Arbor. 11:42. 1930. Figure 60.
Shrubs, subshrubs, or small treelets, 0.3-3(-5) m tall, leafy stems 0.7-3.5 mm thick, glabrous; stipules 2—4 mm long, 1-3.5 mm broad at base, triangular to ovate, trun- cate to acute, with 2 awns 1-1.5 mm long, glabrous or minutely (0.05 mm) papillate, drying reddish brown, caducous. Leaves with petioles 0-6(-8) mm long, 0.7- 1.2 mm thick, glabrous; leaf blades 3-8.5(-l 1?) cm long, 1— 4(-5?) cm broad, elliptic to obovate or oblanceolate, apex acute to subacuminate, base cuneate to acute, dry- ing stiffly chartaceous to subcoriaceous, grayish to dark reddish brown, glabrous above and below, 2° veins 4— 7/side, often with minute pit domatia or tufts of hairs in the axils of 2° veins beneath. Inflorescences terminal and solitary, 3-4.5 cm long, 3-4 cm wide, rounded pan- icles with opposite or whorled branches, peduncles 5- 20 mm long, 0.5-0.8 mm thick, glabrous, bracts ca. 1 mm long, flowers subsessile in distal cymes, pedicels 0.5- 1 mm long. Flowers glabrous externally, hypanthium ca. 0.8 mm long, obconic, calyx lobes ca. 0.5 mm long; corolla funnelform to salverform, white, tube ca. 2 mm long, 1-1.2 mm diam., lobes 5, ca. 1.5 mm long and 0.7 mm broad at the base; stamens 5, anthers ca. 0.8 mm long. Fruits 4-5 mm long, 3-4 mm diam., ellipsoid with ca. 10 longitudinal ridges, bright red; pyrenes ridged.
Plants of wet evergreen lowland forest forma- tions (and in open pine savannas in Nicaragua), from near sea level to 200 m elevation (to over 1 300 m in Mexico and Guatemala). Flowering in southern Central America in March-May and Sep- tember; fruiting in July-December. The species ranges from Mexico, mostly along the Caribbean, to Panama but is disjunct from Nicaragua to cen- tral Panama.
Psychotria fruticetorum is recognized by the small, often cuneate and subsessile leaves, cadu- cous stipules with two distal awns, general lack of pubescence, small domatia, small inflorescences, and small flowers and fruit. The tendency of the leaves to dry grayish or reddish and the bright red berries are characteristics of subgenus Psychotria. Presently, this species is not known from Costa Rica, though it grows near Bluefields, Nicaragua,
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and in central Panama. It may be restricted to limestone soils.
Psychotria glomerulata (J. D. Smith) Steyerm., Mem. New York Bot. Gard. 23: 670. 1972. Ce- phaelis glomerulata J. D. Smith, Bot. Gaz. 16: 12, pi. 1. 1891. Figure 18.
Small shrubs or treelets, 0.6-2(-4?) m tall, much- branched, leafy stems 1-5 mm thick, quadrangular, gla- brous; stipules with a short (0.5-2 mm) sheath, truncated and entire at apex, glabrous, persisting. Leaves with pet- ioles 3-1 1(-15) mm long, 0.8-1.3 mm thick, glabrous, yellowish green; leaf blades 5-14(-17) cm long, 1.5- 5(-6.5) cm broad, narrowly oblong to narrowly elliptic- oblong, oblong or lanceolate, apex acuminate with tip 4-1 2 mm long, base obtuse to acute or cuneate and slightly decurrent on petiole, drying chartaceous to stiffly char- taceous, glabrous above and below, 2° veins 9-13/side and loop-connected near the the margin. Inflorescences terminal and solitary, capitula 1.2-2.5 cm long, 1.5-3.5 cm broad, oblate to subglobose, subtended by 4 decus- sate involucrate bracts 10-16 mm long and 1 2 mm broad, oblong to broadly obovate and rounded distally, greenish yellow or whitish and sometimes edged with blue or purple, glabrous, peduncle to 5(-15) mm long, flowers tightly enclosed within the bracts and spatulate bracte- oles. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx lobes 5, 0.5-1 .5 mm long; corolla funnelform, white, tube 10-15 mm long, 1-2 mm diam.. lobes 5, 1.5-3 mm long; stamens 5, anthers 2-3 mm long. Fruits 6-13 mm long and 5-10 mm diam., ellipsoid, bright blue; pyrenes 4-6 mm long and 3-4 mm diam., appar- ently smooth.
Understory shrubs of evergreen rain forest for- mations, from near sea level to 600 m elevation along the Caribbean slope and coastal plain. Prob- ably flowering and fruiting throughout the year but flowering mostly In March-April and August-Oc- tober. This species ranges from southern Mexico and Belize along the Caribbean lowlands to Pan- ama.
Psychotria glomerulata is recognized by its short stature, narrow leaves, lack of pubescence, solitary sessile terminal heads with four (to eight) white or yellow-green involucrate bracts, and lowland Ca- ribbean habitat. Our figure is based on the original illustration. This species is similar to P. guapilensis with long stipule lobes; P. apoda Steyerm. of South America may be synonymous.
Shrubs or small treelets, 1.5-3(-5) m tall, leafy stems 0.6-4 mm thick, glabrous, flattened or quadrangular at first, soon terete and with thickened nodes; stipules 1-3 mm long, 1.5-2.5 mm broad, with 2 acute lobes and distal sinus 0.3-2 mm deep, glabrous, deciduous or per- sisting. Leaves with petioles 3-22 mm long, 0.3-1 mm thick, glabrous; leaf blades (2-)3-10(-13) cm long, (0.8-)l-2.5(-3) cm broad, narrowly elliptic-oblong to lanceolate or linear-lanceolate, apex long-acuminate with tip 7-15 mm long, base gradually narrowed or abruptly acute and slightly decurrent on petiole, drying charta- ceous, dark green or brown above, glabrous above and below, 2° vein 8-1 3/side, often with parallel intermediate 2° veins. Inflorescences terminal and solitary (pseu- doaxillary), 4-14 cm long, to 1 1 cm broad, open pyra- midal panicles with distant slender opposite or subop- posite branches, peduncles 15-52 mm long, 0.7-1.2 mm thick and glabrous, bracts 3-10 mm long and linear to triangular (or leaf-like), distal branching cymose (often minutely puberulent), flowers usually sessile in triads. Flowers glabrous on the exterior, hypanthium ca. 1 mm long, obconic, calyx tube ca. 1 mm long, subentire or with 5 acute lobes to 0.5 mm long; corolla funnelform, greenish white, tube 4-6 mm long, 0.6-1 mm diam., lobes 5, 1-2 mm long. Fruits 3-5 mm long, 2.5-4 mm diam., ellipsoid-oblong, blue, purple, or white, persisting calyx 0.3-0.7 mm long; pyrenes 3-5-angled or -ridged.
Plants of evergreen montane forest formations, from 1000 to 2000 m elevation. Flowering in March, April, and October; fruiting in January and March-August. The species ranges from the Cor- dillera de Guanacaste eastward along the high- lands to Darien, Panama.
Psychotria goldmanii is recognized by its higher- elevation habitat, smaller narrow long-acuminate glabrous leaves, calyx with minute lobes, longer corolla tube, and small blue fruit. The leaves often dry with the lower surfaces markedly paler in color and obscure 3° venation. The stipules with two small lobes are also distinctive. Few other species of Psychotria in southern Central America develop such narrow linear-lanceolate leaves (an extreme form of which is represented by the type of P. torresiana, Standley 39769 us). Compare P. stey- ermarkii with more numerous 2° veins and P. val- eriana with one stipule appendage. Smaller-leaved specimens of this species may be difficult to dis- tinguish from Palicourea montivaga, but that spe- cies has shorter leaves, very slender petioles, a tendency to develop a short stipular tube, inflo- rescences that dry yellowish, and corollas that are swollen at the base.
Psychotria goldmanii Standl., Contr. U.S. Natl. Herb. 18: 130. 1916. P. torresiana Standl., J. Wash. Acad. Sci. 15: 288. 1925. P. eugenifolia Dwyer, Ann. Missouri Bot. Gard. 67: 375. 1980. Figure 55.
Psychotria graciliflora Benth. in Oerst, Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 35. 1853. P. vallensis Dwyer, Ann. Mis- souri Bot. Gard. 67: 438. 1980 (fide Hamilton 1989). Figure 60.
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249
Shrubs, 0.6-2(-3) mm tall, much-branched and often flat-topped, leafy stems 0.7-2(-3) mm thick, glabrous (except for reddish colleters at the node), usually with 2 (opposite) longitudinal ridges between nodes; stipules with a narrowly triangular to ovate base 1-2 mm long and 2 awns 1-2 mm long, with ascending hairs 0.1-0.2 mm long, reddish brown, deciduous. Leaves mostly clus- tered near the ends of branchlets, petioles 2-10 mm long, 0.5-1 mm broad, glabrous; leaf blades 1 .5-6(-8) cm long, 0.7-3 cm broad, elliptic to ovate-elliptic, apex bluntly obtuse to acute or subacuminate, base cuneate to acute and decurrent on petiole, drying chartaceous, dark gray or reddish gray above, glabrous above and below, 2° veins 3-6/side, pit domatia rarely present. Inflorescences terminal and solitary, 2-6 cm long, 2-5 cm broad, py- ramidal rounded panicles with 1-2 pairs of opposite branches or umbellate, peduncles l-2(-3.5) cm long, ca. 0.5 mm thick, glabrous and drying dark, bracts and brac- teoles to 1 mm long, flowers often sessile in distal cymes or triads. Flowers glabrous externally, hypanthium ca. 0.7 mm long, obconic, calyx tube ca. 0.5 mm long, lobes 0.2-1 mm high; corolla white, salverform, tube 2.5-3.5 mm long, 0.5-0.8 mm diam., lobes 5, 1-2 mm long; stamens 5, anthers ca. 0.9 mm long. Fruits 4-6 mm long, 3-5 mm diam., globose to ellipsoid, with ca. 10 longi- tudinal ridges, bright red or orange; pyrenes ca. 4 mm diam.
Understory plants of wet evergreen forest for- mations on both the Caribbean and Pacific slopes, from near sea level to 1700 elevation (to 2500 m elsewhere, and not found below 1000 m on the Pacific slope of Costa Rica). Flowering in Febru- ary-August (mostly in March-May); fruiting throughout the year. The species ranges from southern Mexico to Colombia.
Psychotria graciliflora is recognized by its very small leaves, usually horizontal glabrous branches, caducous stipules with two slender pubescent awns, small open inflorescences with slender peduncles, and small flowers with minute calyx lobes. The bright red fruit and tendency for leaves to dry grayish or reddish are characteristics of subgenus Psychotria. Costa Rican material of this species has smaller leaves with fewer secondary veins than material from Honduras and Nicaragua (Hamil- ton, 1989, cited under the genus). This species resembles P. carthagenensis, P. chagrensis, and P. parvifolia (see fig. 60).
Psychotria grandis Sw., Prodr. 43. 1788. Figure 64.
Shrubs or small trees, (1.5-)4-8(-10) m tall, trunks to 20 cm diam., leafy stems 4-10 mm thick, terete, glabrous or minutely papillate-puberulent in early stages; stipules 8-30 mm long, 5-20 mm broad, broadly ovate-trian- gular with the margins often revolute, acute to acuminate
and often minutely (1 mm) 2-lobed, usually glabrous, often inflated at the base, deciduous. Leaves with poorly denned petioles 0.5-3.5 cm long, 1.3-3 mm broad, gla- brous; leaf blades ( 1 1-) 1 8-40 cm long, (3-)6-l 6 cm broad, obovate to elliptic-obovate or oblanceolate, apex obtuse to short-acuminate with tip ca. 5 mm long, gradually narrowed to the cuneate base and long-decurrent on pet- iole, drying chartaceous to subcoriaceous, grayish green to pinkish gray or dark grayish above, glabrous above and below, 2° veins 12-16/side. Inflorescences terminal or pseudoaxillary and solitary, 12-25(-30) cm long, 10- 18 cm broad, open pyramidal panicules with opposite branching or with 4 branches (2 larger and 2 smaller) at lower nodes, peduncles 11-18 cm long, 2-4 mm thick, glabrous or minutely (0.05 mm) papillate-puberulent, flowers 2-5 in distal cymules, bracts ca. 2 mm long and triangular, pedicels 1-3 mm long, bracteoles ca. 0.5 mm long. Flowers mostly glabrous externally, hypanthium ca. 0.8 mm long, obconic, calyx tube ca. 0.6 mm long, lobes 5, 0-0.3 mm high, triangular; corolla white, fun- nelform, tube 2-4 mm long, 1.5-2 mm diam., lobes 5, 1.5-2 mm long and 1.1 mm broad at the base; stamens 5, anthers 0.8 mm long. Fruits 5-7 mm long and 5 mm diam., broadly ellipsoid or subglobose, orange or red, surfaces smooth and rounded; pyrenes ca. 5 mm long, with 4-5 rounded dorsal ribs.
Trees of wet evergreen forest formations on both the Caribbean and Pacific slopes of Costa Rica, from near the seashore to 800 m elevation. Flow- ering in February-July; fruiting mostly in July- February. The species ranges from southern Mex- ico and the Greater Antilles to northern South America.
Psychotria grandis is recognized by its often larger (tree) habit, large glabrous-obovate leaf blades de- current on the petioles, large ovate-acuminate stipules with reflexed lateral margins, large inflo- rescences, and small flowers. The red fruit and vegetative parts drying grayish or reddish are char- acteristics of the subgenus Psychotria. Guatemalan material is often puberulent and with larger inflo- rescences; Nicaraguan collections have larger fruit. This species is closely related to P. costavenia Gri- seb., which ranges from Mexico and Cuba into southern Nicaragua and is smaller in its parts.
Psychotria guadalupensis (DC.) Howard, J. Ar- nold Arbor. 47: 139.1 966, sensu lato. Loranthus guadalupensis DC., Prodr. 4: 294. 1830. Vis- coides pendulum Jacq., Select. Stirp. Amer. 73, pi. 51, f. 1. 1763. P. parasitica Sw., Prodr. 44. 1 788, nomen illeg. P. pendula (Jacq.) Urb., Symb. Ant. 1: 445. 1900, non P. pendula Hooker f., 1880. P. peperomiae Standl., Contr. U.S. Natl. Herb. 18: 132. 1916. P. orchidearumSiandl.J. Wash. Acad. Sci. 18: 276. 1928. Figure 60.
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Small epiphytic shrubs, stems 0.2-1 m long, leafy stems 1-4 mm thick, glabrous, often slightly succulent; stipules with a short (0.5-2.5 mm) usually membranaceous trans- lucent sheath, breaking apart and becoming thickened and whitish at the base. Leaves with petioles 1-6 mm long, 0.7-1.2 mm thick, glabrous; leaf blades 12-40 mm long, 7-22 mm broad, ovate or ovate-elliptic to obovate, apex bluntly obtuse to rounded, base obtuse to acute and slightly decurrent on petiole, drying stiffly chartaceous to subcoriaceous, gray to dark gray above, paler beneath, glabrous above and below, with 2-5 obscure 2° veins per side arising at angles of 20-40° (veins rarely prominent). Inflorescences terminal and solitary, 1.5-5 cm long, equally broad, few-flowered cymes to pyramidal panicles with opposite or 3 -partite branching, peduncles 3-30 mm long, 0.4-1 . 1 mm thick, glabrous and reddish in life but often drying black, bracts 2-6 mm long, longer bracts linear, flowers in distal cymes or triads, pedicels 0.5-5 mm long. Flowers glabrous externally, hypanthium 1- 1.2 mm long, 0.5-0.7 mm diam., calyx tube ca. 0.3 mm long, calyx lobes 0.4-1 mm long, triangular and acute, often reddish; corolla white or rose, salverform, tube (4-)6-8 mm long, ca. 1 mm diam., lobes 4, 1.5-2 mm long. Fruits 3-6 m long, 3-5 mm diam., globose to ob- late, usually with 4 pyrenes, red or purple and finally black.
Epiphytes of lower montane cloud forest for- mations of the Caribbean slope and continental divide, from (200-)1000 to 2300 m elevation. Flowering in January-August; fruiting in Febru- ary-September and December. The species ranges from Mexico and the West Indies to the Guianas.
Psychotria guadalupensis is recognized by its epiphytic habit, unusual stipular tube, thick leaves often with obscure venation, variable inflores- cences with slender peduncles, longer-tubed flow- ers, and four-locular ovary. The leaves drying grayish and red fruit are characters shared with subgenus Psychotria. The differentiation between this species and material placed under P. pithe- cobia may be artificial, though it does separate a large portion of the specimens effectively. Costa Rican material placed here has longer corolla tubes, and leaf secondary veins are less numerous and less prominent when dried as in P. pithecobia, and they arise from the midvein at a smaller angle. But there do appear to be a few intermediates, and paucity of collections makes it difficult to tell whether or not the longer corolla tubes of P. guad- alupensis are a consistent difference. (See Ho- ward's discussion of variation in this species, J. Arnold Arbor. 47: 139-142. 1966.)
The preceding description refers to Costa Rican material (excluding specimens assigned to P. pi- thecobia) and differs from that given by Steyer- mark (1974). For example, corolla tubes are ca. 3 mm long in Venezuela (and in P. pithecobia), much
shorter than usually seen in this material. It seems probable tbat the length of the corolla tube varies as greatly (between different plants) as do so many other characters of inflorescences and leaves in this complex.
The type of P. orchidearum Standl. (Standley & Valeria 50863 us) has rather small (2 cm long) leaves and small inflorescences, but the leaves are not as narrow as those in P. maxonii. The type of P. peperomiae Standl. (Pittier 3235 us from Pan- ama) has somewhat larger leaves and is interme- diate between the larger-leaved forms of this spe- cies and the type of P. orchidearum. The fact that all these plants share the same habitats and alti- tudinal range suggests that they may be part of a single polymorphic complex.
Psychotria guapilensis (Standl.) Hammel, Selby- ana 12: 139. 1991. Cephaelis discolor Polak., Linnaea 41: 572. 1877, not/*, discolor (Griseb.) Rolfe. C. tonduzii Krause, Bot. Jahrb. Syst. 54, Beibl. 1 19: 45. 1916, not P. tonduzii Standl. C. nana Standl., J. Wash. Acad. Sci. 17: 171. 1917, not P. nana K. Krause. Evea guapilensis Standl., J. Wash. Acad. Sci. 15: 104. 1925. C. nicara- guensis Standl., Trop. Woods 16: 46. 1928. C. guapilensis (Standl.) Standl., Publ. Field Co- lumb. Mus., Bot. Ser. 4: 295. 1929. Figure 7.
Herbs or subshrubs, 25-90 cm tall, with only 1 or 2 unbranched terete stems 2-4 mm thick, glabrous; stip- ules with basal portion 2-4 mm long, with 2 narrow triangular lobes 3-8 mm long, glabrous, often deciduous. Leaves with petioles 2-9.5 cm long, 0.8-2 mm thick, glabrous; leaf blades 9-24 cm long 3.5-10 cm wide, el- liptic-oblong to oblong or elliptic-obovate, apex abruptly narrowed and acuminate, tip 6-1 5 mm long, base obtuse to acute and slightly decurrent on petiole, drying thinly chartaceous, dark green or brown, glabrous above and below, 2° veins 10-14/side. Inflorescences terminal and solitary, ca. 5-8 cm long, the dense capitula 2-3 cm long and 2.5-5 cm broad, globose to oblate, peduncles 4-40 mm long, ca. 1.5 mm thick, glabrous, with many ovate- involucrate bracts 6-10 x 6-10 mm, covering the outer surface and becoming dark reddish purple in color, brac- teoles spatulate and with purple hairs, flowers sessile. Flowers glabrous externally, hypanthium ca. 1.5 mm long, calyx short-cupulate, lobes 4-5(-6), 0.8-1.5 mm long, triangular, corolla pale pink to purple, funnelform, tube ca. 2.5 mm long, lobes 5(-6), 1.5-2 mm long. Fruits ca. 10 mm long, pyriform to ellipsoid, bright blue; py- renes 5-6 mm long and 4 mm broad, smooth to slightly angled on the dorsal surface.
Understory plants in primary rain forest for- mations of the Caribbean slope in Costa Rica,
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from 30 to 900 m elevation. Probably flowering and fruiting throughout the year but most collec- tions have been made in July-September. The spe- cies ranges from southeastern Nicaragua to the Choco region of Colombia.
Psychotria guapilensis is recognized by its small herbaceous habit, long and slender petioles, dense rounded short-pedunculate terminal heads cov- ered by many purplish to maroon bracts, and bright blue fruit. This species resembles P. glomerulata.
Psychotria haematocarpa Standl., J. Wash. Acad. Sci. 18: 274. 1928. Figure 56.
Shrubs, 1-2.5 m tall, leafy stems 0.8-2.5 mm thick, glabrous and terete, with 2 opposite longitudinal ridges in early stages; stipules with a short (ca. 1 mm) truncate sheath with 2 filiform awns per side 2-5 mm long, early deciduous or the base of the stipule becoming thickened. Leaves with petioles 2-8(-14) mm long, ca. 1 mm thick, glabrous, drying yellowish; leaf blades 6-15 cm long, 2- 5.5 cm broad, elliptic to oblong-elliptic, apex acuminate with tip 5-13 mm long, base attenuate or acute and decurrent on petiole, drying membranaceous or thin- chartaceous, green or grayish green, glabrous above and below, 2° veins 6- 1 0/side and loop-connected near (1.5- 3 mm) the margin. Inflorescences terminal (pseudoax- illary), solitary, rounded to subcapitate 4-10 mm long, to 10 mm broad, condensed cymes with 5-9 flowers, peduncles 2-5 mm long, ca. 0.5 mm thick, glabrous (or minutely puberulent), bracts 0.5-2 mm long, green, gla- brous, pedicels 0-2 mm long. Flowers minutely papil- late-puberulent externally, hypanthium ca. 1 mm long, calyx 1-2 mm long, thin, lobes 5, 0.8-1.3 mm long, triangular; corolla short-salverform, pale greenish or white, tube 2-2.5 mm long, ca. 0.5 mm diam. at base, lobes 0.5-1 mm long, acute. Fruits 5-6 mm long, 4-5 mm diam., elliptic-oblong to globose, orange and finally bright red (dark purple?) at maturity; pyrenes with 4—5 rounded ridges.
Plants of evergreen forests on the Caribbean slope from 50 to 1000 m elevation and on the Pacific slope at 500-1000 m. Flowering in July-August near La Selva; fruiting in October-February. This species ranges from Nicaragua to Colombia.
Psychotria haematocarpa is recognized by its small stipules with deciduous filiform awns, small- er green dried leaves with loop-connected second- ary veins, very small condensed inflorescences, and red fruit. The calyx resembles the corolla in size and texture. This is one of only a few species of subgenus Heteropsychotria with red fruit, rather than blue or black. These are the smallest inflo- rescences of any of our species of Psychotria (or any of our shrubby species of Rubiaceae), and this small size may explain the paucity of collections, as the flowers and fruit are very difficult to see
among the leaves. Compare P. brachybotrya, with larger bracts and purple fruits, and smaller spec- imens of P. hoffmannseggiana, with larger bracts.
Psychotria hazenii Standl., Publ. Field Columb. Mus., Dot. Ser. 7: 96. 1930. P. ramonensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1360. l938.CephaelischlorochlamysSlandl, loc. cit. 1278. 1938. Figure 55.
Small shrubs, 1-2 m tall, leafy stems 0.9-3 mm thick, glabrous, drying yellowish green; stipules with a tubular sheath 3-7 mm long, with 2 narrow lobes 4-1 1 mm long and 0.5-1.5 mm broad, linear to narrowly lanceolate, glabrous. Leaves with petioles 6-20(-40) mm long, 0.3- 0.8(-1.3) mm broad, glabrous; leaf blades (3.5-)6-l 1(-17) cm long, (l-)1.8-3.5(-4) cm broad, narrowly elliptic to elliptic-oblong, apex tapering gradually or abruptly and acuminate, tip 5-12 mm long, base acute to obtuse and slightly decurrent on petiole, drying chartaceous, dark green or brownish, glabrous above and below (or sparsely and minutely puberulent along the midvein beneath), 2° veins 11-17/side. Inflorescences terminal and solitary (or 3), 2-4 cm long, 1 .5-2.5 cm broad, loosely condensed bracteate heads, peduncles 10-25 mm long (to 30 mm in fruit), ca. 1 mm thick and glabrous, bracts subtending the inflorescence branches 1-2 cm long, 4-7 mm broad, ovate and acute or acuminate, glabrous externally (pu- berulent within), chartaceous, green with bluish tips, flowers sessile within enclosing bracts. Flowers with hy- panthium ca. 1.8 mm long and 1.4 mm diam., obovoid- oblong, white, glabrous, calyx tube ca. 0.7 mm long, lobes 5, 1-2 mm long, ca. 1.2 mm broad at the base, triangular, bluish distally; corolla not seen. Fruits 4-8 mm long, oblong, becoming bright blue with a spongy texture, persisting calyx 1-2 mm long.
Plants of lower montane rain forest formations along the Caribbean slope and continental divide, from 900 to 1 600 m elevation. Probably flowering in May-July; fruiting in August-November. In Costa Rica this species is only known from the Caribbean slopes of the Cordillera de Tilaran and central Volcanic Highlands; it is also found in Co- lombia and Ecuador.
Psychotria hazenii is recognized by the tubular stipules with two long narrow awns, the slender petioles, the condensed inflorescences with large broadly imbricate bracts, and bright blue fruit. Our inability to find material with corollas sug- gests that flowering is nocturnal. This species re- sembles P. steyermarkii with smaller bracts and P. pittieri; compare also Palicourea skotakii.
Psychotria hebeclada DC., Prodr. 4: 513. 1830. Probably including P. molliramis (K. Schum. & Krarse.) Steyerm., Mem. New York Bot. Gard.
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23: 529. 1972. Palicourea molliramisK. Schum. & Krause, Bot. Jahrb. Syst., 40: 331 1910. Fig- ure 55.
Shrubs or subshrubs, 0.7-2(-3) m tall, leafy stems 1- 4 mm thick, terete, sparsely to densely puberulent or hispidulous with curved whitish hairs 0.2-0.8 mm long, often contracted beneath the node when dried; stipules with a short truncate tube 0-2 mm long, with 2 distal linear lobes or spines 1-3 mm long, puberulent, persist- ing or breaking apart. Leaves with petioles 4-15(-30) mm long, 0.8-1.2 mm thick, puberulent; leaf blades 7- 20 cm long, 3-8 cm broad, elliptic to ovate-elliptic or elliptic-obovate, apex acuminate with tip 4-14 mm long, base cuneate to acute (obtuse), drying membranaceous to thin-chartaceous, dark green to brownish green above, much paler beneath, sparsely puberulent or subglabrous above, puberulent beneath (especially along the midvein) with thin straight hairs 0. 1-0.4 mm long, 2° veins (6-)8- 14/side. Inflorescences terminal and solitary (or 3), (2.5-)3-6(-12) cm long, 1.5-4(-8) cm broad, narrowly pyramidal or subcylindrical open panicles of closely spaced flowers (in early stages), expanding in fruit, pri- mary branches opposite or alternate, peduncles to 45(-60) mm long, 1-1.8 mm thick, with curved or crooked thin hairs 0.2-0.6 mm long, bracts 2-4 mm long, linear (not apparent beneath the proximal branches), pedicels 1- 2(-5) mm long, often purplish, bracteoles 1-2 mm long. Flowers distylous, puberulent externally, hypanthium ca. 0.7 mm long and 0.5 mm diam., obconic, calyx lobes 5, 0.3-1.5 mm long, ca. 0.3 mm broad, ligulate; corolla white or tinged with pink or purple near the mouth, salverform, tube 2-4 mm long, 1-2 mm diam. and often constricted in middle, lobes 5, 1.5-2 mm long; anthers 1.2-1.7 mm long. Fruits 4-5 long, 3-5 mm diam., sub- globose, with 8 or 1 0 sharply denned longitudinal ridges, glabrous, purple or blue-black; pyrenes 2.5-4.5 mm long, ridged.
Plants of evergreen forest formations on the Ca- ribbean slope (20-600 m elevation) and on the Pacific slope of southern Costa Rica from 40 to 700(-1000?) m. Flowering in March-August in Costa Rica and Panama; fruiting in May-January. The species ranges from southern Mexico to Ec- uador and Venezuela.
Psychotria hebeclada is recognized by its small stature, distinctive pubescence on most parts, two- awned stipules, relatively compact pyramidal to subcylindrical conical inflorescences (in early stages), flowers with well-developed calyx lobes, and small fruit finally becoming black. These plants are often found along rivers, streams, and forest edges. This species was called P. pubescens in the Flora of Barro Colorado Island, but P. pubescens is found in deciduous forests and has smaller calyx lobes. It appears that P. hebeclada is synonymous with P. molliramis as used by Steyermark (1974). Poor material of this species may resemble P. ra- cemosa. Liesner 14126 CR with glabrous inflores-
cences, large calyx lobes, and minutely papillate- puberulent corolla tube is tentatively placed here; it may be a closely related South American species.
Psychotria hoffmannseggiana (Willd. ex Roem. & Schult.) Muell. Arg. in Mart., Fl. Bras. 6(5): 356. 1881. P. furcata DC., Prodr. 4: 512. 1830. P. involucrata sensu Standley and others, in part, not Sw. (cf. Steyermark, 1974). Figure 56.
Shrubs, 0.7-2(-4) m tall, leafy stems 0.7^4.5 mm thick, minutely (0.1-0.2 mm) glabrous to papillate-puberulent (conspicuously pubescent with thin straight hairs in a few collections); stipules 2-4 mm long, basal sheath 0.3- 2 mm long, with 2 awns 1-3 mm long (4/node) separated by a U-shaped sinus, glabrous (pubescent), usually per- sisting. Leaves with petioles 2-8 mm long, ca. 0.5 mm thick, glabrous or with hairs 0. 1-0.2(-0.5) mm long; leaf blades 5-14(-17) cm long, 1.5-6(-8) cm broad, ovate- elliptic, narrowly ovate, elliptic or elliptic-oblong (lan- ceolate), apex tapering gradually and acuminate with tip 3-14 mm long, base cuneate to acute and often decurrent on petiole, drying thinly chartaceous and usually green- ish, glabrous above or puberulent only on midvein, gla- brous or puberulent beneath with thin hairs 0. 1-0.3 mm long (more rarely with hairs ca. 0.5 mm long), 2° veins 5-9/side. Inflorescences terminal and solitary, 6-25 mm long, involucre up to 6 cm broad, capitate or condensed- cymose with (3-)7-30 congested flowers, peduncles 3- 20 mm long, 0.7-1 .8 mm thick, pubescent or glabrescent, bracts 3-22(-35) mm long, 1-6 mm broad, linear-lan- ceolate, conspicuous and persisting, green to purple, flowers sessile. Flowers glabrous or puberulent exter- nally, hypanthium 1-2 mm long, calyx lobes 0.2-0.8 mm long, acute to rounded; corolla white, funnelform, tube 2-6 mm long, 0.8-2.5 mm diam., lobes 1.3-2 mm long; stamens 4 or 5, anthers ca. 1 mm long. Fruits 3-6 mm long, 4-5 mm diam., subglobose to slightly oblong, with 8 longitudinal ridges, becoming purple to dark maroon or black, glabrous or puberulent; pyrenes ridged.
Plants of wet evergreen lowland rain forest for- mations, from near sea level to 700 m elevation, both on the Caribbean slope and in the General Valley and Golfo Dulce area. Flowering primarily in May-August; fruiting in July-March. The spe- cies ranges from Veracruz, Mexico, into Venezuela and Brazilian Amazonia.
Psychotria hoffmannseggiana is recognized by its persisting two-awned stipules, small capitate inflorescences with conspicuous narrow spreading or recurved persisting bracts, small flowers, and purple fruit. The subcapitate inflorescences can expand after anthesis and become slightly branched. The leaves can vary greatly in size in different plants and, while most of our plants have little pubescence, a few are densely pubescent. Plants of southwestern Costa Rica often have con-
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spicuously larger bracts than specimens from other areas. This species is very similar to P. officinalis, which has larger branched inflorescences. The ma- terial placed here was called P. involucrata Sw. by Standley (1938 and in herb.), but Steyermark (1972, p. 603) showed that the Swartz name is a synonym of P. officinalis. Some authors may prefer to retain P. furcata as a distinct species, but the South American material of P. hoffmannseggiana in- cludes a wide range of variation, some of which is similar to that seen in the type of P. furcata.
cences, and the small flowers often with large calyx lobes. The leaves are often undulate along the edge in life and may have an arcuate submarginal vein. The red fruits and tendency for the leaves to dry grayish are characteristics of subgenus Psychotria. There seems to be great variation in flowers and inflorescence, with thicker puberulent peduncles being correlated with more congested flowers. The calyx lobes also seem to vary greatly in their de- velopment. Compare P. quinqueradiata.
Psychotria horizontalis Sw., Prodr. 44. 1788. P. longicollis Benth. in Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 33. 1853. Figure 65.
Small shrubs to little treelets, 1-3 m tall, leafy stems 1—4 mm thick, glabrous (rarely minutely puberulent), usually becoming pale grayish; stipules 2-7 mm long, 1- 4 mm broad at base, triangular to narrowly ovate, obtuse to acute, usually glabrous, drying reddish brown, cadu- cous. Leaves with petioles l-7(-ll) mm long, 0.5-1.3 mm thick, glabrous or minutely puberulent; leaf blades 3— 9(-l 3) cm long, 1 .5-4.5(-6) cm broad, elliptic to ovate- elliptic, obovate, lanceolate or oblanceolate, apex acute to short-acuminate, base obtuse or slightly auriculate, drying thin-chartaceous or chartaceous, dark gray to pinkish gray or brown, glabrous above and below (rarely with thin hairs ca. 0.2 mm long on the veins beneath), 2° veins 5-9/side, occasionally with tufted hairs or pit domatia in vein axils beneath. Inflorescences terminal and solitary, 2-6 cm long (to 13 cm in fruit), l.5-4(-7) cm broad, rounded open panicles with opposite or whorled branching (often globose-umbellate), peduncles l-3(-7) cm long, 0.5-1.3 mm thick, glabrous or puber- ulent, with 2-5 flowers separate or closely clustered in distal cymes, pedicels 0-2 mm long, bracteoles 0.2-1 mm long. Flowers glabrous externally, hypanthium 0.7- 1 mm long, obconic, calyx tube ca. 0.6 mm long, lobes 5, 0.5-3 mm long, linear to ligulate; corolla funnelform, white, tube 2.3-3.5 mm long and 0.7-1.4 mm diam., lobes 5, ca. 1.5 mm long; anthers 0.7-1 .2 mm long. Fruits 4-8 mm long, 3-6 mm diam., ellipsoid to ovoid, orange becoming bright or dark red; pyrenes ca. 4 x 3 mm, with 3-5 rounded ridges.
Plants of both evergreen and seasonally decid- uous forest formations, most often collected from the seasonally dry Pacific slope, 20-1600 m ele- vation. Flowering most often in May-August; fruiting throughout the year (June-December in Costa Rica). This species ranges from Mexico to Ecuador and Brazil; it is also found in Cuba and Hispaniola.
Psychotria horizontalis is recognized by its smaller often subsessile leaves often with domatia, unlobed stipules, often umbelliform inflores-
Psychotria insignis Standl., Contr. U.S. Natl. Herb. 18: 130. 1916. Figure 66.
Small treelets or shrubs, 2.5-6 m tall, leafy stems 3- 8 mm thick, distinctly flattened in early stages, densely dark reddish brown pubescent with hairs 0.1-0.4 mm long; stipules 12-27 mm long, 4-10 mm broad, ovate- lanceolate with an acute bifid tip, awns 2-4 mm long, reddish pubescent with longer hairs along the midrib and margins, usually caducous. Leaves with petioles 3-5.5 cm long, 1.5-3 mm thick, densely pubescent with dark reddish brown hairs 0.1-0.5 mm long; leaf blades 14- 30 cm long, 7-18 cm broad, broadly elliptic to broadly elliptic-ovate or elliptic-obovate, apex abruptly nar- rowed and acuminate with tip 7-18 mm long, rounded at the subcordate based, basal lobes often unequal, form- ing a sinus 0-10 mm deep, drying chartaceous, dark grayish brown or dark reddish brown, glabrous above (except the midvein), densely pubescent beneath, 2° veins 14-18/side and loop-connected in distal part of the blade. Inflorescences solitary and terminal, 5-9 cm long, 4-6 cm broad, short pyramidal panicles with 4 lateral branches (2 short and 2 longer) at the first node and short dense branches at the closely congested second and third nodes, peduncles 1 .8-4 cm long, 1 .4-2 mm thick densely pilose, bracts of the first node 6-8 mm long, flowers sessile in dense glomerules of 3-10, bracteoles 1-3 mm long. Flow- ers reddish puberulent externally, hypanthium 0.5-1 mm long, obconic, calyx tube 0.5 mm long, lobes 5, 0.5-1 mm long, triangular; corolla white, tube 2-3 mm long, with prominent hairs 0.3-0.4 mm long, lobes 5, ca. 1 mm long. Fruits 5-6 mm long, 3-4 mm diam., ellipsoid with longitudinal sulci (dried), puberulent, calyx per- sisting.
Plants of the wet evergreen Caribbean slope, from 20 to 400 m elevation. Flowering in Septem- ber-October; fruiting in September. We have seen only three collections: Herrera 2248 (CR) and Gra- yum et al. 8929 (CR, MO) from central Costa Rica and the type (Pittier 4410 us holotype) from San Bias, Panama.
Psychotria insignis is recognized by the densely pubescent parts, large long-petiolate leaves often subcordate at the base, and unusual compact in- florescence. The upper surface of the leaf blade is sometimes vaginate at the juncture with the pet-
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iole. This species is a member of subgenus Psy- chotria. Our material differs from the type in that the 2° veins do not arise at 90° angles from the midvein and the submarginal is not as well de- veloped. This name is used provisionally for the Costa Rican collections.
Psychotria ipecacuanha (Brotero) Stokes, Bot. mat. med. 1: 365. 1812. Callicocca ipecacuanha Bro- tero, Memoria sobre a Ipecacuanha do Brasil 27. 1 801, Trans. Linn. Soc. London 6: 137. 1802. Cephaelis ipecacuanha (Brotero) A. Rich., Bull. Fac. Med. 4: 92. 1818. Figure 13.
Herbaceous subshrubs, 25-50 cm tall, usually with a single erect unbranched stem, rhizomatous, leafy stems 1.5-4 mm thick, glabrous, terete; stipules with a short (2 mm) sheath, truncate with 4-8 setae per side 3-6 mm long, persisting. Leaves crowded distally, petioles 3-8 mm long, 1-2 mm thick, glabrous; leaf blades 7-17 cm long, 4-9 cm broad, obovate to oblong or elliptic-ob- ovate, apex acute or short-acuminate, base cuneate to rounded-obtuse, drying membranaceous to thin-char- taceous, glabrous or minutely puberulent above and be- low, 2° veins 5-7/side. Inflorescences terminal or axil- lary, solitary, capitulae to 2 cm long and 1-3 cm broad, subglobose, peduncles 1-4 cm long, deflexed, involucrate bracts 5-10 mm long, ovate, acute, flowers sessile. Flow- ers distylous, glabrous externally, hypanthium ca. 2 mm long, ellipsoid, calyx teeth 5, ca. 0.5 mm long; corolla funnelform, white, tube 3-4 mm long, cylindrical, lobes 5, 1.5-2.5 mm long; stamens 5, anthers ca. 1.6 mm long. Fruits ca. 10 mm long, becoming red then black; pyrenes 6-7 mm long, ridged.
Plants of the lowland Caribbean rain forest for- mations (0-600 m elevation), and probably the result of introduction for cultivation in Central America. The species ranges from southeastern Nicaragua to the Amazon basin of Brazil.
Psychotria ipecacuanha is recognized by its short unbranched rhizomatous habit, unusual fimbriate stipules, leaves with broad blades and short peti- oles, small involucrate heads, and red to black fruit. The roots and rhizomes are the source of the alkaloid emetin and are used in folk medicine or cultivated for pharmaceutical use as ipecac. The species is referred to as "ipecac" and raicilla or ipecacuana.
Psychotria jimenezii Standl., J. Wash. Acad. Sci. 1 5: 288. 1 925. P. wendlandiana Oerst. ex Standl., loc. cit. 18: 9. 1928. Figure 63.
Shrubs and small treelets, 2-5(-7) m tall, to 15 cm dbh, leafy stems 1.2-4.5 mm thick, pubescent with as-
cending reddish hairs ca. 0.3 mm long (rarely glabrous), terete; stipules 5-12 mm long, 2-5 mm broad at the base, triangular to ovate, acute to acuminate (or shortly bifid), puberulent, margins ciliolate, drying reddish brown. Leaves with petioles variable among plants, 0-20(-35) mm long, 0.8-1.5 mm thick, hirsutulous to papillate- puberulent; leaf blades 6-16 cm long, 1.7-5 cm broad, oblanceolate to narrowly obovate or narrowly elliptic- oblong, apex acute to acuminate with tip 3-10 mm long, base gradually narrowed and cuneate or rounded-auric- ulate, drying membranaceous to thin-chartaceous, dark brown or almost black above, glabrous above, with short (0. 1-0.2 mm) reddish hairs on the veins beneath, 2° veins 8-1 2/side. Inflorescences terminal and solitary, 4-10 cm long, 3-5 cm broad, open pyramidal panicles, peduncles 1 5-50 mm long, ca. 1 mm thick, yellowish green in life, densely reddish brown hirsutulous when dried, first node with 4 branches (2 long and 2 short), bracts 2-6 mm long, triangular, flowers subsessile in crowded distal clus- ters of 3-8, bracteoles 0.5-1 mm long. Flowers usually glabrous externally, hypanthium ca. 0.7 mm long, ob- conic, calyx ca. 0.5 mm long, calyx lobes obscure; corolla tubular, cream white to yellowish or greenish yellow, tube 1.5-2.5 mm long and 1.4 mm diam., lobes 5, ca. 1.2 mm long, 0.9 mm broad at base; stamens 5, anthers 0.7 mm long. Fruits 4-6 mm long, 4-5 mm diam., sub- globose, bright red; pyrenes with 4-5 rounded longitu- dinal ridges.
Understory plants of evergreen lower montane cloud forest formations along the Caribbean slope and continental divide, from (200-)400 to 1600 m elevation. Flowering in January-June; fruiting in February- April, August-October, and Decem- ber. The species is found in the Cordilleras de Guanacaste and Tilaran and on the Caribbean side of the Meseta Central in central and north-central Costa Rica.
Psychotria jimenezii is recognized by its reddish pubescence (sometimes glabrous), usually oblan- ceolate leaves, triangular stipules acute to acu- minate, inflorescences with four branches at the first node and second node (two short basal and two longer ascending branches), and small gla- brous flowers. The tendency of the leaves to dry grayish and the fruit becoming red are character- istics of the subgenus Psychotria. The length of petioles and shape of the leaf (especially the base) can differ greatly in different collections of this species. The narrowed lamina base is subauricu- late in the type of P. wendlandii (Wendland 781 us). Compare P. horizontalis, P. orosiana, and P. laselvensis.
Psychotria lamarinensis C. Hamilton, Phytologia 64: 227. 1988. Figure 66.
Shrubs, ca. 2 m tall, leafy stems 1.7-4.5 mm thick, glabrous, grayish, terete; stipules 8-12(-20) mm long, 6-
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1 2 mm broad, broadly ovate, apex obtuse, reddish brown, glabrous, caducous. Leaves with petioles 2-8 mm long, 1.4-2.7 mm thick, glabrous, slightly sulcate above; leaf blades 1 3-22 cm long, 5-13 cm broad, elliptic to broadly elliptic or elliptic-obovate, apex acute or short-acumi- nate with tip ca. 5 mm long, base obtuse to cuneate or slightly truncated (rounded), drying chartaceous, grayish or reddish brown, 2° veins 8-1 I/side, minute pit domatia sometimes present in the vein axils. Inflorescences ter- minal or pseudoaxillary, 2-5, 1.5-3 cm long, to 2 cm broad, condensed globose panicles of cymes with short lateral branches (or appearing umbellate), peduncles 4- 20 mm long, 0.8-1.2 mm thick, glabrous, dark red, usu- ally with 4 subequal branches at first node, bracts 0.2- 2 mm long, triangular, bracteoles not apparent, pedicels 0.5-1 .5 mm long. Flowers glabrous externally, calyx tube ca. 0.5 mm long, cupulate, truncate or with very short teeth; corolla tubular, white, tube 2.5-3 mm long and 1 mm diam., lobes 5, 1-2 mm long and 1 mm broad, with a linear (1.5 mm) extension from near the apex; stamens 5, anthers ca. 0.7 mm long. Fruits 7-8 mm long, 4-5 mm diam., oblong-ellipsoid; pyrenes smooth.
Plants of deeply shaded sites in evergreen rain forest formations on the Caribbean slope, at 20- 500 m elevation. Flowering in March-May; fruit- ing in November. This endemic species ranges from Canalete (Alajuela) to Cahuita (Limon).
Psychotria lamarinensis is recognized by its larg- er broad glabrous short-petiolate leaves (cuneate basally but slightly rounded at the petiole), small short-branched globose panicles, and broadly ovate stipules. The leaves drying grayish or reddish brown are characteristic of species in subgenus Psy- chotria. The extensions on the corolla lobes are separate in bud. This species is closely related to P. quinqueradiata, but P. lamarinensis has shorter corolla tubes and larger leaves with the veins di- verging at angles of 70-80°.
Psychotria laselvensis C. Hamilton, Phytologia 64: 228. 1988. Figure 61.
Shrubs, 1.2-4 m tall, leafy stems 1.3-3 mm thick, erect, glabrous, bark smooth; stipules 2-4(-6) mm long, 1-3 mm broad, triangular to ovate, glabrous or puber- ulent on the midrib externally, caducous. Leaves sub- sessile or with petioles 1-7 mm long, 0.4-0.8 mm thick, glabrous; leaf blades 5-13(-16) cm long, 1.5-4(-6) cm broad, elliptic to narrowly elliptic-oblong or elliptic-lan- ceolate, apex acute or short-acuminate with tip to 1 cm long, base acute to cuneate and often slightly auriculate at the petiole, drying thin-chartaceous, dark grayish above, glabrous above and below, 2° veins 7-10/side, pit dom- atia often present in distal vein axils beneath. Inflores- cences terminal and solitary, 6-1 5 cm long, 4-7 cm broad, open pyramidal, panicles with thin opposite branches, peduncles 4-9 cm long, 0.4-0.7 mm thick, glabrous, with 2 or 4 lateral branches at the first node, bracts 0.5-1 .5(-3?) mm long, flowers often in distal pairs or triads, pedicels
ca. 0.5 mm long, bracteoles ca. 0.5 mm long. Flowers distylous, glabrous externally, hypanthium ca. 0.7 mm long, obconic, calyx lobes poorly developed (to 0.2 mm long); corolla narrowly funnelform, white or pale yellow, tube 1.5-3 mm long, 1-2 mm diam., lobes 5, 1.5-2 mm long; anthers 1-1.2 mm long. Fruits 4-5 mm diam., globose, red; pyrenes with 4-5 rounded ridges.
Plants of lowland Caribbean rain forest under- story, at 50-300 m elevation on the Caribbean slope. Flowering in February-June (mostly May); fruiting in July. The species is only known from the La Selva research station and nearby areas, Heredia, and northern Limon provinces.
Psychotria laselvensis is recognized by its small- er elliptic to oblanceolate leaves, larger inflores- cences with very thin open spreading branches, and pedicellate flowers. The veins are often flat- tened and expanded at the vein axils and form pit domatia distally. The leaves drying grayish and the red fruit are characteristics of subgenus Psy- chotria. This species resembles P. marginata with more acute leaf bases and is closely related to P. graciliflora and P. orosiana.
Psychotria limonensis K. Krause, Bot. Jahrb. 54, Beibl. 119: 43. 1916. Figure 64.
Shrubs, subshrubs, or small trees, 0.6-2(-6) m tall, leafy stems 2-6 mm thick, glabrous or sparsely puber- ulent, drying reddish to pale gray or dark; stipules 5-12 mm long, 3-6 mm broad, triangular to ovate, rounded to acute, entire or shortly bifid, glabrous, drying reddish brown, caducous. Leaves with petioles 1.2-6 cm long, 0.7-3 mm thick, glabrous; leaf blades 1 1-22(-28) cm long, 5-12(-15) cm broad, ovate-elliptic to elliptic-ob- long or broadly elliptic, apex acute to short- or long- acuminate with tip 5-20 mm long, base obtuse to acute (occasionally slightly decurrent on petiole), drying thin- to stiffly chartaceous, gray to dark reddish brown, gla- brous above and below, 2° veins (7-)10-19/side and distally loop-connected to form a submarginal vein. In- florescences terminal (pseudoaxillary), solitary of 3, 3- 9 cm long, 5-11 cm broad, broad panicles with opposite or whorled branches, peduncles 1-4 cm long, 1.2-2.5 mm thick, glabrous and reddish brown, bracts 0.5-1.2 mm long, triangular, flowers in congested cymes of 3-6, pedicels 0-1.5 mm long. Flowers glabrous externally, hypanthium ca. 0.5 mm long, obconic, calyx tube ca. 0.4 m long, lobes 0.2-0.5 mm high, obtuse; corolla funnel- form to rotate, white, tube 2-2.6 mm long and 1-1.7 mm diam., lobes (4-)5(-6), 1.2-1.5 mm long, triangular; stamens (4-)5(-6), anthers 0.8-1 mm long. Fruits 4-5 mm long, 3-4 mm diam., ellipsoid, bright red; pyrenes with ca. 5 longitudinal ridges.
Plants of evergreen lowland rain forest forma- tions on both the Caribbean and Pacific slopes, from near sea level to 200 m elevation (to 700 or
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rarely 1 700 m elsewhere). Probably flowering and fruiting throughout the year (fruiting primarily in June-January in Costa Rica). The species ranges from southern Mexico and Belize to Panama and Colombia.
Psychotria limonensis is recognized by its large leaves with many secondary veins and arcuate submarginal vein, usual lack of pubescence, un- lobed stipules, small flowers with four to six co- rolla lobes, and small fruit. The red fruit and ten- dency for leaves to dry gray or pinkish are characteristics of subgenus Psychotria. The sec- ondary veins arise at almost 90° angles from the midvein. This species can be mistaken for spec- imens of P. panamensis var. compressicaulis with larger leaves and calyptrate stipules. It is also sim- ilar to P. mexiae with longer corollas and acute buds.
Psychotria longipedunculoides C. M. Taylor, nom. nov. P. longipedunculata Dwyer, Ann. Missouri Bot. Card. 67: 389. 1980, non P. longepedun- culata (Gardn). Muell. Arg. in Mart.
Herbaceous subshrubs, 0.5-1.2 m tall, usually un- branched, leafy stems 1.7-6 mm thick, semisucculent, glabrous or minutely papillate-puberulent; stipules 6-8 mm long, 6-8 mm broad, broadly ovate, margin irregular or roughly 2-lobed, thin and translucent, deciduous or persisting. Leaves with petioles 14-33 mm long, 0.8-1.5 mm thick (dried), glabrous; leaf blades 12-18 cm long, 5-7(-9) cm broad, elliptic to elliptic-oblong or elliptic- obovate, apex acute or acuminate with tip 5-10 mm long, gradually narrowed to the acute base and slightly decurrent on petiole, drying membranaceous to thin- chartaceous, pale olive gray above, 2° veins 12-15/side, an arcuate submarginal vein present. Inflorescences sol- itary and axillary from distal nodes, 12-16 cm long, 1.5- 3 cm broad, congested pyramidal panicles to subcapitate, flowering portion 1.2-3 cm long, peduncles 10-14 cm long, 0.9-1.4 mm thick, glabrous, bracts l-6(-12) mm long, linear-subulate, flowers sessile or subsessile. Flow- ers glabrous externally, hypanthium ca. 0.5 mm long, ellipsoid, calyx lobes 5, ca. 1.5 mm long; corolla to 2.5 m long, funnelform, greenish white, lobes 5, stamens 5; anthers ca. 0.8 mm long. Fruits 4-6.5 m long, ca. 3 mm diam., oblong-obovoid; pyrenes with 4-5 rounded ridg- es.
Recently collected on the Caribbean slope of the Talamanca mountains at ca. 700 m elevation near the border with Panama (Herrera 3255 CR, MO, usj). Flowering and fruiting in July. This species is only known from eastern Costa Rica, western Panama, and Choco, Colombia.
Psychotria longipedunculoides is recognized by the single-stemmed semisucculent habit, glabrous
parts, solitary axillary inflorescences with small flower clusters on a long peduncle, and membra- naceous ovate stipules. Similar in habit to P. mac- rophylla but differing by linear bracts, congested flowers, and stipules.
Psychotria luxurians Rusby, Mem. Torrey Bot. Club 6: 50. 1896. P. berteriana subsp. luxurians (Rusby) Steyerm., Mem. New York Bot. Gard. 22: 534. 1972. Figure 57.
Small trees or shrubs, 2-8(-l 2) m tall, leafy stems 2.5- 7 mm thick, glabrous or minutely (0.05 mm) papillate- puberulent, terete; stipules 3-6(-9) mm long, to 6 mm broad, broadly ovate rounded to obtuse, splitting in age or with a short ( 1 mm) sinus, usually glabrous, decidu- ous. Leaves with petioles 5-35 mm long, 1 .3-3 mm thick, glabrous or minutely papillate-puberulent; leaf blades 1 2-26(-32) cm long, 5-1 1(-1 3) cm broad, elliptic-oblong to elliptic or oblong, apex acuminate with a narrowed tip 4-15 mm long, base obtuse to cuneate (acute) base, drying chartaceous, dark green or brown above (distinct- ly paler beneath), glabrous above and below or minutely papillate-puberulent beneath, 2° veins 8-14/side. Inflo- rescences terminal, solitary or 3, (7-) 12-26 cm long, 8- 20 cm broad, pyramidal open panicles with mostly op- posite branching, peduncles 2-8 cm long, 1 .2-6 mm thick, minutely papillate-puberulent, bracts of primary branch- es l-8(-12) mm long and ca. 1 mm broad, triangular to elliptic, flowers congested in cymose clusters of 3-10, bracteoles 1.5-3 mm long, elliptic, pedicels 0-2 mm long. Flowers glabrous externally, hypanthium 0.7-1.3 mm long, obconic, calyx lobes 0.2-0.5(-0.8) mm long, broad- ly obtuse; corolla short-funnelform, white, greenish white or yellowish, tube 2-4 mm long and 1.5-2 mm diam., lobes 5, 1-1.7 mm long and 0.7-1 mm broad, obtuse or rounded. Fruits 3-5 mm long and 3-6 mm diam., oblong or globose, blue, blue-black, or purple; pyrenes 3 mm long, with 4-5 dorsal ridges.
Plants in open secondary growth and light gaps in evergreen rain forest formations of the Carib- bean slopes and lowlands, 20-1000 m elevation. Flowering primarily in March-July in Costa Rica; fruiting in July-November. This species ranges from southern Nicaragua to Bolivia.
Psychotria luxurians is recognized by its large essentially glabrous leaves usually drying greenish, the short and broad stipules that are entire or with two very short rounded lobes, large minutely pa- pillate-puberulent inflorescences with many small elliptic bracts, numerous small flowers, and small blue fruit. Ants may live within the stems of this species (Beach 1480 CR). Note that the persisting bracts subtending the flowers can be mistaken for calyx lobes. This species can be confused with P. angustiflora and P. berteriana, but those species do not have their flowers surrounded by as many
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conspicuous bracteoles, and the flowers differ. Steyermark (1972) treated this as a variety of P. berteriana with larger bracts, but the stipule mor- phology is different in the two. Psychotria tapan- tiensis of higher elevations is closely related with larger bracteoles, corollas, and fruits.
Psychotria macrophylla Ruiz & Pav., Fl. Peruv. 2: 56. 1799. P. anomothyrsa K. Schum. & J. D. Smith, Bot. Gaz. 35: 3. 1903. P. macrophylla var. angustissima Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 203. 1940. P. macrophylla ssp. anomothyrsa (K. Schum. & J. D. Smith) Steyerm., Mem. New York Bot. Gard. 23: 563. 1972. Figure 12.
Herbs or subshrubs, 0.5-1 .5(-2) m tall, leafy stems 3- 1 1 mm thick, semisucculent, usually unbranched, gla- brous, becoming terete; stipules 2-5 mm long, triangular, usually with a narrow divergent succulent slightly bi- lobed apex, glabrous. Leaves with petioles 2-8 cm long, 1.2-3 mm thick, glabrous or rarely puberulent in early stages; leaf blades 12-32 cm long, 4-14 cm broad, elliptic to elliptic-oblong, narrowly to broadly oblong, or lan- ceolate, apex acuminate with tip 8-18 mm long (rarely gradually narrowed and acute), base obtuse to acute and slightly decurrent on petiole, drying membranaceous to thin-chartaceous, dark greenish to dark brownish above, usually glabrous above and below, 2° veins 8-15/side, occasionally loop-connected near the margins. Inflores- cences axillary, usually I/node, 3-15 cm long (to 25 cm in fruit), 4-6 cm wide, usually open pyramidal panicles with opposite branching, peduncles 0.7-8(-12) cm long, glabrous or occasionally puberulent with hairs 0.1-0.4 mm long, bracts 0.5-2(-10) mm long, triangular, flowers solitary or in small glomerules of 2-3 along the distal axes, sessile or subsessile. Flowers glabrous or puberulent externally, hypanthium ca. 0.5 long, obconic, calyx ca. 0.7 mm long, lobes ca. 0.2 mm high, broadly triangular; corolla tubular-sal verform, white, tube 2-4 mm long and 0.6-1.5 mm diam., lobes 4 or 5, 1-2 mm long; stamens 4-5, anthers 1.2 mm long. Fruits 5-7 mm long and 3- 4 mm diam., oblong, white and arenchymatous; pyrenes ca. 6 mm long, with 4-5 rounded ridges.
Understory plants of wet sites in evergreen for- est formations, from 20 to 2200 m elevation in Costa Rica. The species appears to be more com- mon in highland forests than in those below 500 m elevation; it is rare in sandy streamside soils at La Selva. Flowering and fruiting throughout the year (but flowering mostly in January-August). The species ranges from southern Mexico to Bolivia.
Psychotria macrophylla is recognized by its usu- ally short stature with succulent unbranched main stems, the axillary inflorescences, small subsessile flowers on branched inflorescences, white fruit with four to five rounded ribs. This species is variable;
leaves range from narrowly lanceolate to broadly elliptic in different plants; most plants are glabrous but a few are minutely puberulent. Some dried collections have crinkled leaf margins. The ar- rangement of flowers on the distal branches of the inflorescence varies greatly, from open-cymose to spicate. Steyermark (1974) interpreted some of this variation in Venezuela as hybridization between P. macrophylla and P. uliginosa, but this seems doubtful and it may be simply that individual plants of P. macrophylla can vary greatly in the expression of a variety of morphological features. Compare Psychotria aggregata, with densely ag- gregated flowers, slightly larger calyx lobes, and pyrenes with a single dorsal ridge. See the discus- sions under P. capacifolia, P. siggersiana, and P. aggregata. This group of species was recently stud- ied by Molly Nepokroeff (wis, 1992).
Psychotria marginata Sw., Prodr. 43. 1788. P. ni- caraguensis Benth. in Oersted, Vidensk. Med- del. Dansk. Naturhist. Foren. Kjobenhavn 1852: 34. 1853. Figure 61.
Shrubs or treelets, (0.5-)l-3(-4) m tall, leafy stems 1 .3-4 mm thick, terete, glabrous; stipules 5-14 mm long, to 5 mm broad, narrowly ovate and acute, glabrous, usually drying reddish brown, caducous. Leaves often clustered distally, petioles 4-22(-35) mm long, 1-1.8 mm thick; leaf blades 6-17 cm long, 2-6 cm broad, narrowly obovate to elliptic obovate, oblanceolate or elliptic-oblong, apex acute to acuminate with tip 3-10 mm long (rarely obtuse), base acute to cuneate and de- current on petiole, drying chartaceous or stiffly charta- ceous and grayish, glabrous above, glabrous beneath or minutely (0.05 mm) papillate-puberulent on the veins beneath, 2° veins 7-13/side, weakly loop-connected in the distal part of the leaf, margin thickened along the edge, usually ciliolate, midvein usually with small ex- panded flaps-like domatia near the vein axils beneath. Inflorescences terminal and solitary, 7-17 cm long, 5- 1 2 cm broad, open pyramidal panicles with thin opposite branches diverging at 90° angles, peduncles 2-7 cm long, 0.6-1.7 mm thick, glabrous, bracts 1-3 mm long, tri- angular, flowers 2-6 in distal cymes, pedicels l-3(-9) mm long. Flowers distylous, glabrous externally, hypan- thium ca. 0.6 mm long, obconic, calyx tube ca. 0.4 mm long, lobes 0.2 mm high, broadly triangular or obscure; corolla short-funnelform, white or yellowish, tube 2-3 mm long, 0.7-1.3 mm diam., lobes 5, 1-1.5 mm long; anthers 0.8-1.1 mm long. Fruits 3.5-6 mm long, 3-6 mm diam., subglobose, red or purple; pyrene ca. 3 mm long, with 4-5 ridges.
Plants in secondary growth of evergreen forests and partly deciduous forest formations, from 10 to 800 (-1 100) m elevation. Flowering and fruiting throughout the year (fruiting most often in Feb-
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ruary-May in Costa Rica). At La Selva, this spe- cies flowers in the wettest part of the year: July- November. The species ranges from Mexico and the West Indies to Peru and Bolivia.
Psychotria marginata is recognized by its small stature, acute triangular stipules, usually obovate leaf blades often with pronounced domatia and ciliolate margins, small flowers, and small fruit on long thin pedicels in open inflorescences. The red fruit and leaves drying grayish (less often reddish) are characteristics of subgenus Psychotria. Com- pare this species to P. laselvensis, which lacks cil- iolate leaf margins and has a shortly truncated or rounded leaf base.
Psychotria maxonii Standl., Proc. Biol. Soc. Wash. 37: 53. 1924. Figure 60.
Small epiphytic shrubs with stems 0.2-0. 5(-l) m long, often pendant, leafy stems 0.6-2 mm thick, succulent, glabrous and often becoming pale grayish; stipules with united truncated membranaceous tube 1-2 mm long, glabrous, persisting and sometimes becoming thickened and pale-colored at base. Leaves subsessile or with pet- ioles 1-3 mm long and 0.3-0.5 mm thick, glabrous; leaf blades 12-23(-28) mm long, 3-5(-6) mm broad, nar- rowly lanceolate to linear-lanceolate, apex tapering grad- ually and acute, base acute or obtuse and slightly de- current on petiole, drying stiffly chartaceous or subcoriaceous dark gray above, glabrous above and be- neath, 2° and 3° veins obscure in dried material, margins slightly revolute. Inflorescences terminal or axillary, I/node, 2-4 cm long, 2-4 cm wide, open few-flowered 3-branched pyramidal panicles or cymes with slender opposite branches often red in color, bracts ca. 0.4 mm long, peduncles 3-12 mm long, 0.2-0.5 mm thick, gla- brous, drying black, pedicels filiform, 1-5 mm long. Flowers glabrous externally, hypanthium ca. 1 mm long, obovoid to oblong, calyx ca. 0.7 mm long (including lobes), lobes 4, ca. 0.5 mm long and triangular; corolla salverform, white and often tinged with red or pink, tube 2.5-4(-6) mm long, 0.4-0.8 mm diam., lobes 4, 1-1.5 mm long, obtuse; stamens 4, anthers ca. 1 mm long. Fruits ca. 4 mm long and 4 mm diam., globose, becoming orange or red and finally black; pyrenes smooth.
Epiphytic plants in montane cloud forest for- mations, from 1500 to 2300 m elevation. Flow- ering in February-March, May, and July-August; fruiting in January-March, July, and October. The species ranges from the northern part of the Mese- ta Central (near San Ramon) along the Caribbean escarpment and continental divide to the high- lands of western Panama.
Psychotria maxonii is recognized by the epi- phytic habit, small narrow succulent leaves, small thin few-flowered inflorescences, and small flowers with slender corolla tubes. The leaves dry grayish
and the fruit become red, then black (as do species of subgenus Psychotria). Panamanian collections appear to have more ovate leaves than are found in Costa Rica. No other Central American species of Psychotria has such small and narrow leaves. This species is closely related to P. guadalupensis and shares the same habitats in Costa Rica with that species. It is possible that the two species are actually conspecific, with P. maxonii being no more than a very unusual form of P. guadalupensis; see the discussions under/*, guadalupensis and P. pith- ecobia.
Psychotria mexiae Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 296. 1929. Palicourea nigres- cens M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 1 1(50): 136. 1844, not Psychotria nigrescens De Wild, 1924. P. schippii Standl. & Steyerm., Publ. Field Mus. Bot. Hist., Bot. Ser. 23: 24. 1943. Figure 62.
Shrubs or small trees, 2-5(-10) m tall, leafy stems 2- 4 mm thick, glabrous (in Costa Rica) or sparsely puber- ulent; stipules 1 2-20(-40) mm long, 1 .5-3(-5) mm broad at the base, narrowly lanceolate and sheathing the apex (as in Ficus), glabrous, caducous. Leaves with petioles 3-20 mm long, 1.3-2 mm broad, glabrous; leaf blades 8-16(-20) cm long, 2.5-7(-8.5) cm broad, elliptic to nar- rowly elliptic, apex acuminate with tip 10-17 mm long, base acute to obtuse and decurrent on petiole, drying thinly chartaceous to stiffly chartaceous, dark above, gla- brous above and below (rarely with hairs along midvein beneath), 2° veins 10-13/side, domatia with small hairs often present in the vein axils beneath. Inflorescences terminal, 1-3, 3-6 cm long, 2-5 cm wide, rounded to broadly pyramidal panicles with usually 4 branches at the first node, peduncles 8-28 mm long, 1 .5-2 mm thick, bracts and bracteoles not apparent, cymules of 3-8 flow- ers, pedicels 0.5-1 mm long. Flowers glabrous or pu- berulent externally, calyx tube ca. 0.3 mm long cupulate, lobes little developed; corolla funnelform, white (drying pink), tube 2.5-4.5 mm long, ca. 1 mm diam., lobes 5, 1.5-2 mm long, ca. 1 mm broad at the base; stamens 5, anthers 0.8-1 mm long. Fruits 4.5-6 mm long, 3.5-5 mm diam., ellipsoid to subglobose, becoming red; py- renes with 4-5 ridges.
Plants of lower montane evergreen forest for- mations, at about 600-700 m elevation in Costa Rica. Fruiting in January (Standley & Valeria 45770 & 46375 us). The species ranges from Mex- ico to the region of Tilaran in northern Costa Rica.
Psychotria mexiae is recognized by its elliptic leaves with small domatia, fused stipules forming a cap over the apex, short inflorescences, longer corolla tubes often drying pink, and spherical fruit with shallow longitudinal grooves when dried. The
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leaves drying grayish and the red fruit are char- acteristic of subgenus Psychotria. This species re- sembles P. tenuifolia, P. panamensis, and P. li- monensis. This species is only known in Costa Rica from the two collections cited above.
The red fruit and tendency of the leaves to dry grayish or reddish brown are characteristics of the subgenus Psychotria. Close to P. neilii with leaves abruptly truncate and rounded. It may be confused with material of P. mortoniana.
Psychotria micrantha H.B.K., Nov. gen. sp. 3: 363 (quarto), pi. 284. Nov. 1 8 1 9. P. rufescens Roem. & Schult., Syst. Veg. 5: 192 . Dec. 1819, non P. rufescens H.B.K. Figure 63.
Shrubs or small trees, 2-8 m tall, leafy stems 3-8 mm thick, densely hirsutulous with erect reddish brown hairs 0.3-0.7 mm long; stipules 9-18 mm long, 3-8 mm broad at base, triangular to narrowly ovate, with 2 narrow lan- ceolate acuminate lobes 1-7 mm long, reddish brown puberulent, deciduous. Leaves with petioles 6-26 mm long, 1 .2-3 mm thick, densely hirsutulous with reddish brown hairs; leaf blades 12-30 cm long, 4-13 cm broad, oblong to elliptic, or narrowly elliptic-oblong, apex acute to short-acuminate with tip 5-10 mm, base obtuse to cuneate and slightly decurrent on petiole (rarely slightly auriculate), drying thin- to stiff-chartaceous, grayish to dark reddish brown, pubescent above and below with hairs 0.2-0.9 mm long, longest and most dense on the midvein beneath, 2° veins 14-22/side and loop-con- nected distally, 3° veins often subparallel. Inflorescences terminal (pseudoaxillary) and solitary (rarely 2), 5-15 cm long, to 12 cm broad, pyramidal to rounded open panicles usually with 4 closely set branches (2 longer and 2 shorter) at first node of 1° and 2° axes, peduncles 4- 10 cm long, 0.8-1.5 mm thick, densely hirsutulous with reddish brown hairs 0.3-0.5 mm long, bract, 2-3 mm long, triangular, flowers sessile in distal clusters or capit- ulae of 3-8. Flowers densely puberulent externally, hy- panthium and calyx 1.5-2 mm long, lobes ca. 0.2 mm long; corolla white, salverform, tube 1.5-2.5 mm long, 0.5-1 mm diam.. lobes 5, 1.3-2 mm long; stamens 5, anthers 0.8-1.2 mm long. Fruits 3-7 mm long, 3—4 mm diam., ellipsoid to subglobose, red, hirsutulous; pyrenes with 4-5 low ridges.
Rarely collected plants of evergreen rain forest formations, from 250 to 700(-1000) m elevation in Costa Rica. Flowering in May-September in southern Central America; fruits have been col- lected primarily in late May-December. In Costa Rica this species has only been collected from the Zona Protectora above La Selva, the drainage area of the Rio Renventazon, and the General Valley. The species ranges from Guatemala, Nicaragua, and central Costa Rica to Venezuela, Peru, and Bolivia.
Psychotria micrantha is recognized by its dense hirsutulous surfaces with reddish brown hairs, large bifid stipules, large leaves with many secondary veins, open paniculate inflorescences with flowers in dense distal clusters, and small hirsutulous fruit.
Psychotria microbotrys Ruiz ex Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 204. 1930. Figure 58.
Shrubs or small treelets, l-3(-4) m tall, leafy stems 1.7-6 mm thick, quadrangular, glabrous; stipules united around the stem for 1-4 mm or apparently separate, 7- 2 1 mm long, 2-5 mm broad at base, ovate, deeply bifid on each side with 2 prominent narrowly ovate or tri- angular lobes on each side (4/node), these acute and often minutely bifid, glabrous, persisting. Leaves with petioles 7-25(-38) mm long, 1-2.5 mm thick, glabrous, drying dark; leaf blades 11-27 cm long, 4-1 3 cm broad, ovate- elliptic, elliptic or oblong, apex short-acuminate with tip 5-13 mm long, base obtuse to acute and slightly decur- rent on petiole, drying thin-chartaceous, yellowish green or brownish green, glabrous on both surfaces, 2° veins 7-12/side. Inflorescences terminal and solitary, 3-6 cm long, 2.5—4 cm wide (to 15 x 10 cm and purplish in fruit), pyramidal paniculate with opposite or alternate branching, peduncles 2-6 cm long, 1-1.8 mm thick, gla- brous, bracts 2-4 mm long, linear, branches often di- chotomous or trichotomous distally, flowers alternate or in groups of 2-4, bracteoles ca. 1 mm long, pedicels 0- 1 mm long. Flowers glabrous externally, hypanthium ca. 0.8 mm long and 0.5 mm diam., cylindrical, calyx tube 0.3-0.4 mm long, calyx lobes minute or obscure; corolla salverform, yellow, white, or greenish, tube 1.8-3.2 mm long, 0.4-0.8 mm diam., lobes 5, ca. 1 mm long. Fruits ca. 3-5 mm long and 3-5 mm broad, 8 mm diam. and spongy when fully ripe, oblate to subglobose, green to grayish blue or white, branches of the infructescence of- ten purplish; pyrenes ca. 2.5 mm long, with 4-5 longi- tudinal ridges and transverse ribs.
Understory plants of evergreen forest forma- tions; from 50 to 1000 m elevation on the Carib- bean slope and lowlands, and on the Pacific slope of southern Costa Rica from 500 to 1 500 m ele- vation. Flowering in February-March and May- July; fruiting primarily in July-January. The spe- cies ranges from Costa Rica to Peru.
Psychotria microbotrys is recognized by its larger glabrous leaves, large and deeply two-lobed (or almost separate) persisting stipules, larger inflo- rescences, small flowers, and small fruit becoming spongy white at maturity. The transverse ridges between the ribs on the immature dried fruit and the tendency of the distal inflorescence branches to be dichotomous with alternating flowers or flower clusters are additional distinctions. The majority of Costa Rican collections come from the
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Caribbean lowlands of northern Costa Rica and from the General Valley-San Vito area of the Pa- cific slope. Similar to P. dejlexa, which has much smaller stipules.
not correlated with ecology or geography. This is our only species of the largely African subgenus Tetramerae (Hieron) R. Petit. Compare P. pubes- cens, also of deciduous forest formations but with yellowish flowers.
Psychotria microdon (DC.) Urban, Symb. Ant. 9: 539. 1928. Rondeletia microdon DC., Prodr. 4: 408. 1830. Figure 59.
Shrubs, subshrubs, or small trees (sometimes clam- bering), 0.4-3(-5) m tall, leafy stems 1-3 mm thick, gla- brous or with thin hairs ca. 0.2 mm long, greenish and becoming pale gray; stipules 1-2 mm long, ca. 2 mm broad, broadly triangular, obtuse or rounded, glabrous or puberulent, pale yellowish and thick in texture, often persisting. Leaves usually closely clustered distally, pet- ioles 3-15(-60) mm long, 0.5-1.2 mm thick, glabrous or puberulent; leaf blades 4-15 cm long, 1.5-6 cm broad, obovate to elliptic-obovate, elliptic-oblanceolate, ellip- tic, or ovate-elliptic, apex acuminate with tip 3-14 mm long, base acute to obtuse and decurrent on petiole, dry- ing membranaceous or thin-chartaceous, green to dark brownish green above, glabrous above, glabrous or pu- berulent beneath with thin whitish hairs 0.2-0.4 mm long, usually with thin hairs grouped near the vein axils beneath (domatia), 2° veins 3-7/side. Inflorescences ter- minal, solitary or several, 3-10 cm long, 2-8 cm broad, open rounded panicles with opposite branching or 3 pri- mary branches, peduncles 2-7 cm long, 0.6-1 .7 mm thick, glabrous or minutely puberulent, bracts ca. 1 .4 mm long, flowers in distal cymes of 2-5, pedicels 0.5-4 mm long. Flowers glabrous or minutely papillate-puberulent ex- ternally, hypanthium 1-1.5 mm long, obconic, calyx tube 0.8-2 mm long, 2-3 mm broad, cupulate to campanu- late, lobes 5, minute; corolla salverform, white, tube (4-)6- 10 mm long, 1-2 mm diam., lobes 5, 2-6 mm long, to 2.2 mm broad at base. Fruits 7-10 mm long, 5-7 mm diam., ellipsoid, becoming red-orange; pyrenes with 2- 3 ridges.
Plants of forest understory and thickets in sea- sonally very dry and deciduous woodland for- mations, from near sea level to 200 m elevation (rarely to 600 m). Flowering in May-August; fruit- ing in July-October and January. The species rang- es from Mexico and the West Indies to Venezuela and Peru.
Psychotria microdon is recognized by its decid- uous forest habitat, smooth grayish stems, thin often obovate leaves clustered at the ends of branchlets, broad stiff stipules, thin but well-de- veloped calyx tube with minute or absent lobes, and larger corollas blooming in the first half of the wet season. The leaves sometimes dry grayish green (resembling species of subgenus Psychotria). Spec- imens whose lower leaf surfaces are puberulent have been called P. microdon variety meridionalis (Steyermark, 1 974), but this is a minor distinction,
Psychotria monteverdensis Dwyer & C. Hamilton, Phytologia 64: 230. 1988. Figure 61.
Shrubs or small trees, 2-5 m tall, leafy stems 2.5-4 mm thick, glabrous; stipules with a sheath 7-15 mm long and 4-6 mm broad, triangular to ovate with 2 aris- tate lobes 2-4 mm long, glabrous, drying reddish brown, caducous. Leaves subsessile, petioles 1-5 mm long, 1.2- 2.2 mm broad, glabrous, sulcate above; leaf blades (5-)7- 16 cm long, (l-)2-4.5 cm broad, oblanceolate to nar- rowly elliptic, apex acute or acuminate with tip ca. 5 mm long, base cuneate and slightly rounded (auriculate) at petiole, drying stiffly chartaceous, grayish green or pinkish gray, glabrous above and below, 2° veins (5-)8- 10/side and united distally by an arcuate submarginal vein. Inflorescences terminal and solitary, 4—10 cm long, 3-4 cm wide, open pyramidal panicles, peduncles 15- 50 mm long, 2-2.5 mm thick, glabrous or pilosulous, bracts 4-8 mm long, triangular, ciliate, flowers usually in distal glomerules of 3-6, bracteoles 1-4 mm long, ovate, ciliate, pedicels 0-1.5 mm long. Flowers glabrous externally, hypanthium ca. 2 mm long, obconic, calyx tube ca. 1 mm long, cupular, lobes 5, 0.5-1 mm long, triangular, often ciliolate; corolla funnelform, greenish yellow, tube 2-3(-5) mm long and l-2(-3) mm diam., lobes 5, ca. 2 mm long, with keel-like appendages; sta- mens 5, anthers ca. 1.2 mm long. Fruits 7-8 mm long (not including the 1-2-mm-high calyx), 5-7 mm diam., ellipsoid; pyrenes 4-5-ridged.
Plants of evergreen lower montane rain forest formations, at 900-1600 m elevation. Flowering in May-July; fruiting in June-July, September, and November. This endemic species is only known from the eastern slopes of Volcan Miravalles, the Monteverde Nature Preserve, and the eastern part of the Cordillera de Talamanca.
Psychotria monteverdensis is recognized by its sheathing stipules with slender biaristate tips, sub- sessile and oblanceolate leaves with prominent submarginal veins (often slightly auriculate at the base), few-branched inflorescences with flowers in distal glomerules, and short thick corollas with keel-like structures on the large corolla lobes.
Psychotria mortoniana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 355. 1938. Figure 54.
Shrubs or small trees, 1.5-8(-10) m tall, leafy stems 1 .3-8 mm thick, densely pubescent with straight, curved or crooked hairs 0.3-0.9(-1.5) mm long; stipules 6-18
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
261
mm long, to 9 mm broad near the base, narrowly ovate, bifid and often splitting into 2 acuminate lobes 2-8 mm long, pubescent, persisting. Leaves with petioles 10-45 mm long, 0.8-2.5 mm thick, with straight or crooked yellowish hairs 0.3-1 mm long; leaf blades 13-30 cm long, 5-12 cm broad, elliptic to elliptic-obovate or el- liptic-oblong, apex long-acuminate with tip 1 1-27 mm long, obtuse to acute at the base, drying membranaceous to thin-chartaceous, dark olive green or brownish green, pubescent on the major veins above, densely pubescent beneath with pale-colored hairs (to 1 .2 mm long on mid- vein), 2° veins (14-)17-23/side, weakly loop-connected near the margin in the distal part of the leaf. Inflores- cences terminal and solitary, 9-16 cm long, 8-14 cm broad, open pyramidal panicles with opposite primary branching, peduncles 1-9 cm long, 1.5-3 mm thick, densely villous or hirsutulous with yellowish brown hairs, bracts subtending the proximal pair of branches 6-12 mm long and 2-3 mm broad, narrowly triangular, flow- ers mostly sessile in crowded distal clusters of 3-10, partly enclosed by the persisting bracteoles 6-12 x 2-3 mm. Flowers glabrous externally, hypanthium ca. 1 mm long, obconic, calyx ca. 0.5 mm long, lobes ovate, obtuse; corolla tubular, white to pale yellowish, tube 3-5 m long and 1-1.5 mm diam., lobes 0.8-1.3 mm long and 0.5 mm broad at base, rounded. Fruits 3-6 mm long and 3-4 mm diam., ellipsoid, black, glabrous; pyrenes with 4-5 angles.
Plants of evergreen forest formations of the Pa- cific slope, from 1 00 to 1100m elevation. Flow- ering in May and July-August; fruiting in August- September and December-January. The species is only known from southern Puntarenas province in Costa Rica and from the nearby Chiriqui High- lands in Panama.
Psychotria mortoniana is recognized by its dense pubescence, large leaves with many secondary veins large bifid stipules, larger inflorescences with flow- ers in dense distal clusters, and sessile fruit partly enclosed by persisting elliptic bracteoles. The rounded bracteoles and the tendency to dry dark brown are additional distinctions. This species has only been collected from the General Valley, around Golfo Dulce and near San Vito in southern Costa Rica. Specimens of this species may resem- ble P. pilosa and P. micrantha.
Psychotria neillii C. Hamilton & Dwyer, Phyto- logia 64: 231. 1988. Figure 63.
Shrubs, or small trees, 1-4 m tall, leafy stems 1.5-4 mm thick, hirsutulous with reddish brown hairs ca. 0.7 mm long; stipules 7-16 mm long, 3-5 mm broad at base, basal sheath ca. 4 mm long, apex biacuminate with nar- row tips 3-5 mm long, puberulent, deciduous. Leaves subsessile, petioles 2-8 mm long, ca. 2 mm thick, densely puberulent, terete; leaf blades 8-15(-18) cm long, 3- 6.5(-7.5) cm broad, obovate to elliptic-obovate or ob- lanceolate, apex abruptly short-acuminate with tip 3-8
mm long, base gradually narrowed and rounded-auric- ulate or subcordate at petiole, the lamina drying thin- to stiffly chartaceous, grayish or reddish brown, glabrous above, with thin reddish hairs 0.2-0.8 mm long beneath (especially along the mid vein), 2° veins 8-12/side. In- florescences terminal or becoming axillary, solitary, 7- 13(-15) cm long, to 8 cm broad, open panicles, usually with 2 nodes bearing 2-4 lateral branches, peduncles 3- 8(-10) cm long, 0.8-1.7 mm thick, with short reddish hairs, bracts ca. 4 mm long, bracteoles ca. 0.5 mm long, pedicels 0-1 mm long. Flowers puberulent externally, calyx tube ca. 0.3 mm long, calyx lobes ca. 0.2 mm long, triangular; corolla yellowish white, tube 1.5-2 mm long, ca. 1 mm diam., lobes 5, ca. 1 mm long; stamens 5, anthers 0.8 mm long. Fruits 5-7 mm long, 3.5-5 mm diam., obovoid-oblong and truncated at apex, reddish purple at maturity.
Plants of lowland Caribbean rain forest for- mations, from 80 to 400 m elevation. Flowering in January-February and April; fruiting in Feb- ruary and April-July. The species ranges from southernmost Nicaragua to southernmost Costa Rica.
Psychotria neillii is recognized by the densely hirsutulous young stems, subsessile obovate leaves rounded at the base, thin-puberulent inflorescence axes, small flowers and obovoid-oblong fruit with truncated apex. The leaves usually drying grayish and the red fruit are characteristics of subgenus Psychotria. This species is similar to P. micrantha, which has flowers and fruits in denser clusters, acute leaf bases, and larger stipules. Compare also P. jimenezii from higher elevations, usually gla- brescent, with differing stipules and inflorescences.
Psychotria nervosa Sw., Prodr. 43. 1 788. P. undata Jacq., Hort. Schoenbr. 3: 5, pi. 260. 1798. P. elongata Benth. in Oersted, Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 32. 1853. P. quiinifolia Dwyer, Ann. Missouri Bot. Card. 53: 108. 1966. Figure 63.
Shrubs or rarely small treelets, 0.5-1 .5(-3) m tall, leafy stems 0.9-4 mm thick, sparsely to densely pubescent with usually crooked reddish hairs 0.3-1 mm long; stip- ules 3-9(-l 1) mm long, 2-4(-6) mm broad, narrowly to broadly ovate (sometimes tubular near the base), round- ed distally to obtuse, drying reddish brown, sparsely to densely pubescent, caducous. Leaves usually clustered at the tips of branchlets, petioles 2-22 mm long, 0.7-1.5 mm thick, usually pubescent, with adaxial margins con- tinuous with the leaf margins; leaf blades 7-13(-19) cm long, 2-5(-7) cm broad, elliptic-obovate to elliptic or obovate, apex acute to acuminate with tip 3-10 mm long, base acute and often long-decurrent on petiole, drying thin- to stifF-chartaceous, grayish to pinkish gray or red- dish brown, sparsely puberulent above, sparsely to densely pubescent beneath (especially on the midvein) with hairs
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0.3-0.7 mm long, 2° veins 7-13/side and often weakly loop-connected near the margin. Inflorescences terminal (pseudoaxillary), solitary or several, 1-5 cm long, to 5 cm broad, cymose or trichotomous, peduncle 5-18 mm long, hirsutulous with reddish brown hairs, bracts to 0.5 mm long, flowers closely clustered and sessile or sub- sessile in distal cymules of 3-5. Flowers with hypanthi- um ca. 0.5 mm long, densely hirsutulous, calyx 0.5-1 mm long, calyx lobes 0.2 mm long or obscure; corolla funnelform, white, sparsely puberulent externally, tube 2-4 mm long, 1-2 mm diam., lobes 5(-6), 1-2 mm long; stamens 5, anthers 0.7-1 mm long. Fruits 6-8 mm long, 2.5-4 mm diam. ellipsoid to oblong sparsely hirsute with thin erect hairs, becoming red-orange or red; pyrenes with 3-5 ridges.
Plants of seasonally very dry and deciduous woodland formations, semideciduous forest, and (less often) evergreen forest formations, from 20 to 800 m elevation (to 1 200 near San Vito). Flow- ering and fruiting throughout the year (flowering primarily in June-July, and fruiting in July-Jan- uary in Costa Rica). The species ranges from Mex- ico and the West Indies to northern South Amer- ica.
Psychotria nervosa is recognized by its decidu- ous woodland habitat (less often in evergreen for- ests in Costa Rica), reddish pubescence on most parts, rounded to globose terminal buds, thin usu- ally ovate stipules, leaves narrowed to the base and decurrent on the petiole, and small inflores- cences. The red fruit and tendency for the foliage to dry reddish brown are characteristics of sub- genus Psychotria. This species is sometimes con- fused with P. pubescens, but that species has per- sisting two-lobed stipules.
Psychotria officinalis (Aubl.) Raeuschel ex Sandw., KewBull. 1931:473. \93\ . Nonatelia officinalis Aubl., Hist. pi. Guiane 1: 182, pi. 70, f. 1. 1775. P. involucrata Sw., Fl. Ind. Occ. 1:413. 1797. Figure 59.
Shrubs or small treelets, l-3(-8) m tall, trunks to ca. 2.5 cm diam., leafy stems 1.3-5 mm thick, glabrous or rarely minutely (0.2 mm) puberulent, terete; stipules 2- 4(-6) mm long, 3-6 mm broad, often with a short (0.5- 1 mm) cupular sheath, truncated distally and with 2 narrow lobes 1-3 mm long separated by a U-shaped sinus 1-2 mm wide, usually glabrous and persisting. Leaves with petioles 4-1 2(-20) mm long, 1 .2-2 mm thick, usually glabrous, slightly sulcate above with 2 adaxial ridges; leaf blades 7-17(-25) cm long, 3-8(-12) cm broad, ovate-elliptic to ovate-oblong, lanceolate-elliptic or el- liptic, apex acute to acuminate with tip 6-20 mm long, base obtuse to acute and slightly decurrent on petiole, drying thin-chartaceous, dark yellowish green to olive green, glabrous above, glabrous below or minutely (0. 1
mm) puberulent along major veins, 2° veins 6-10/side and strongly ascending. Inflorescences terminal and sol- itary (rarely 2), (1.5-)2-6 cm long (to 10 cm in fruit), 2-
4 cm broad, panicles with 2-4 opposing pairs of short lateral branches or sometimes subcapitate, peduncles 1- 4(-5) cm long, ca. 1 .4 mm thick, sparsely and minutely puberulent, flowers sessile in distal clusters of 3-8, sub- tended by bracts 2-8 mm long and 1-3 mm broad, lan- ceolate to elliptic, yellow becoming purplish in fruit. Flowers distylous, usually sparsely puberulent with thin whitish hairs 0.1-0.2 mm long externally, hypanthium 0.5-0.9 mm long (obscured by the bracts), calyx cup ca. 0.4 mm high, lobes 0.2 mm high or obscure; corolla funnelform, white or yellow, tube 2-5 mm long, 1.2-2 mm diam., lobes 5, 1-2 mm long and 1.2 mm broad at base; anthers ca. 1.5 mm long. Fruits 4—6 mm long, 4-
5 mm diam., becoming subglobose, becoming dark pur- ple or black, infructescence and pedicels often purplish; pyrenes 2-3 mm long, with 4-5 ridges.
Plants of shaded understory in evergreen forest formations, 50-1200 m elevation. At La Selva it is usually found on slopes and ridges in forest un- derstory. Flowering and fruiting throughout the year (flowering primarily in March-July). The spe- cies ranges from southern Mexico and the West Indies to northeastern Brazil and the Guianas.
Psychotria officinalis is recognized by the small terminal-branched inflorescences with prominent bracts subtending the distal flower clusters, the minutely and sparsely puberulent little flowers, the short two-lobed stipules, and black or purple fruit. The primary inflorescence branches are not sub- tended by bracts, but the distal flower groups are sometimes subtended by a larger central bract and two lateral braces adnate near the base to form a small involucre. The conspicuous bracts and ten- dency of the inflorescence branches to become purplish is reminiscent of the closely related P. hoffmannseggiana with condensed inflorescences. Specimens in early stages of flowering may be mis- taken for that species as well as P. brachybotrya and P. platypoda.
This is one of our most common species of Ru- biaceae, but its range in Costa Rica appears to be restricted to the Caribbean lowlands and lower Caribbean escarpment (to 900 m), the General Valley (ca. 500-900 m), and the San Vito area (to 1 200 m). Except for the San Vito area (near the Panama border), the species has not been collected above 900 m in Costa Rica. The name P. involu- crata has been applied to material of this species and P. hoffmannseggiana in the past. Breeding behavior was studied by Bawa and Beach (1983).
Psychotria orosiana Standl., J. Wash. Acad. Sci. 15: 288. 1925. Figure 61.
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263
Shrubs l-2(-3) m tall, leafy branchlets 0.7^4 mm thick, glabrous, drying gray or blackened; stipules 2-5 mm long, to 3 mm broad at base, narrowly to broadly triangular and with 2 closely set minutely reddish brown puberulent awns with blunt tips, caducous. Leaves clustered distally, sessile or subsessile, petioles 0-5 mm long, glabrous; leaf blades 4-9(-13) cm long, 1.5-3(-5) cm broad, elliptic- obovate, oblong-obovate to elliptic or oblanceolate, apex obtuse to acute or short-acuminate with tip 3-10 mm long, base acute to cuneate (rarely slightly truncated) and long-decurrent on petiole, drying membranaceous to thin- chartaceous, dark gray or blackish above, glabrous above and below, 2° veins 5-7/side, pit domatia often present in the vein axils beneath. Inflorescences terminal and solitary, 3-7 (-10) cm long, 3-6 cm broad, open panicles with few slender opposite primary and secondary branches, peduncles l-4.5(-7) cm long, ca. 0.5 mm thick, glabrous, bracts ca. 1 mm long, triangular, flowers in distal cymes of 3-6, pedicels 0-3 mm long, bracteoles ca. 0.5 mm long. Flowers glabrous externally, hypan- thium ca. 1 mm long, obconic, calyx ca. 0.5 mm long, lobes to 0.3 mm long, triangular or obscure; corolla fun- nelform, white or yellow-white, tube 2-5 mm long, 1- 1.5 mm diam., lobes 5, 1-1.4 mm long; stamens 5, an- thers 0.7-1 mm long. Fruits 4-7 mm long and 4-6 mm diam., ellipsoid, smooth, drying black; pyrenes with 4- 5 ridges.
Plants of evergreen montane forest formations in the central Cordilleras, from (400-) 1 200 to 1 900 m elevation. Flowering in January-July and Sep- tember; fruiting in March, June, and August-Oc- tober. In Costa Rica this species has only been collected from near Monteverde, San Ramon, Car- pintera, and Tablazo and the southern drainage of the upper Rio Reventazon. This species is only known from Costa Rica and western Panama.
Psychotria orosiana is recognized by its smaller obovate leaf blades usually decurrent at the base and often without a well-developed petiole (and often drying blackish gray above), domatia (when present), montane habitat, glabrous parts except stipules with two puberulent awns, and open slen- der-branched inflorescences with small flowers. The tendency of leaves to turn grayish and the red fruit are characteristics of subgenus Psychotria. This species resembles P. graciliflora but has larger in- florescences and longer corolla tubes. It also re- sembles P. laselvensis (q.v.).
Psychotria panamensis Standl., Contr. U.S. Natl. Herb. 18: 132. 1916. P. magna Standl., loc. cit. 131. 1916. P. compressicaulis K. Krause, Bot. Jahrb. Syst. 54, Beibl. 119: 44. 1916. P. gran- distipula Standl., J. Wash. Acad. Sci. 18: 276. 1928, non P. grandistipula Merrill, 1923. P. molinae Standl., Ceiba 1: 46. 1950. Figures 62 and 64.
Small trees or shrubs, 2-12(-20?) m tall, trunks 4-20 cm diam., leafy stems 1.5-8 mm thick, glabrous or mi- nutely (0. 1 mm) puberulent in early stages; stipules 5- 25 mm long, to 55(-80) mm when subtending inflores- cences, 2-7 (-12) mm broad, lanceolate with a long nar- rowed apex often curved back in late stages, united to form a cap over the apex and splitting on 1 or 2 sides, glabrous (ciliolate along the interior edge), greenish white to yellowish in life but drying dark reddish brown, ca- ducous. Leaves differing greatly in size in the highland and lowland varieties (see below), with petioles 5-53 mm long, 1-2.7 mm thick, glabrous or very minutely (0.05 mm) puberulent, opposing petioles often unequal; leaf blades 6-28 cm long, 2-16 cm broad, elliptic to ovate- elliptic or obovate-elliptic, apex acuminate (acute, ob- tuse) with tip 3-15 mm long, base gradually narrowed and acute or obtuse, slightly decurrent on petiole, drying chartaceous, dark grayish to dark reddish brown, gla- brous above, glabrous or minutely (0.05 mm) papillate- puberulent beneath, 2° veins (5-)8-19/side (sometimes loop-connected near the margin). Inflorescences termi- nal and solitary (or 3), 5-10 cm long (to 16 cm in fruit), 5-10 cm wide, open pyramidal panicles branches, pe- duncles 5-35(-50) mm long, 1-2.5 mm thick, usually glabrous, bracts ca. 1 mm long and 2-3 mm broad, cy- mules of 2-5 flowers, pedicels 0-1 (-4) mm long and puberulent. Flowers monomorphic and short-styled, gla- brous externally, hypanthium 0.7-1.5 mm long, calyx 0.5-0.8 mm long and 2 mm diam., calyx lobes not de- veloped or obscure; corolla white or greenish yellow, tube 2-3 mm long and 1.5-2 mm diam., lobes 5, 1-3 mm long and triangular, rotate or reflexed; stamens 5, anthers 0.7-1 mm long. Fruits 5-9 mm long and 4-7 mm diam., ellipsoid to subglobose, yellowish becoming red; pyrene ca. 7 mm long, with 3-5 ridges.
Plants usually found in high rainfall formations, 20-2100 m elevation. Flowering in March-Oc- tober in Costa Rica and Panama (flowering pri- marily in July-August at La Selva); fruiting throughout the year. The species ranges from Mex- ico to Colombia.
Psychotria panamensis is recognized by its habit (usually a small tree), the often long and unequal petioles, large caducous stipules, that enclose the shoot apex as in Ficus, and the flowers with short thick perianth tube and subentire calyx. The nu- merous secondary veins are usually quite straight and not as arcuate as in other species. The ten- dency of foliage to dry grayish or dark reddish brown and the fruit turning red are characteristics of subgenus Psychotria. A majority of Costa Rican collections are from higher-elevation cloud forests and belong to variety panamensis with smaller narrower leaves than the lowland plants (see be- low). This species is closely related to P. sarapi- quensis, which shares the same type of stipule but has narrower leaves with fewer secondary veins, and to P. cocosensis, with larger flowers. Compare also P. nervosa with enlarged stipules below the
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inflorescences and lacking the acute terminal buds of P. panamensis.
Hamilton distinguished four varieties, of which three are found in southern Central America: va- riety compressicaulis (K. Krause) C. Hamilton, va-
riety magna (Standl.) C. Hamilton, and variety panamensis. The following key separates the three varieties. However, it is possible that the two va- rieties found in Costa Rica will prove to be two closely related but distinct species.
la. Dried fruits 4.5-5 mm long, 3.5-4 mm diam.; Caribbean lowlands of eastern Panama
var. magna
Ib. Dried fruits 5.5-8 mm long, 4.5-9 mm diam 2
2a. Leaf blades 15-28 cm long, 5.5-15 cm broad, with 12-19 secondary veins on each side; fruit 5.5-
7 mm long; lowland Caribbean Nicaragua to Panama var. compressicaulis
2b. Leaf blades usually 9-15 cm long, 4-6 cm broad, with 8-16 secondary veins on each side; fruit
usually 7-9 mm long; Mexico to Panama, mostly above 500 m elevation var. panamensis
Psychotria parvifolia Benth. in Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1852: 35. 1853. Figure 60.
Shrubs or small treelets, 0.7-3(-5?) m tall, leafy branchlets 0.5-3 mm thick, at first densely puberulent with minute (0.1-0.2 mm) reddish brown hairs, soon glabrescent; stipules 2-6 mm long, 1-2 mm broad at the base, united and lanceolate or oblong with acute tip, splitting down 1 side, minutely puberulent and drying reddish brown, caducous. Leaves clustered at the ends of branchlets, petioles 2-14 mm long, 0.3-0.6 mm thick, puberulent with minute reddish brown hairs; leaf blades l-3.5(-5) cm long, 0.5-1 .4(-2) cm broad, elliptic to ovate- elliptic or rhombic, apex acute or obtuse, base acute and slightly decurrent on petiole, drying membranceous to thin-chartaceous, dark grayish green or dark brown above, glabrous above, glabrous beneath except for the minute (0.1 mm) reddish brown hairs along the midvein, 2° veins 2-5/side and obscurely loop-connected near the margin. Inflorescences terminal or pseudoaxillary, 1-3/ node, 5-20 mm long, to 8 mm broad, cymose or 3-branched, peduncles 2-8 mm long, 0.4-0.8 mm thick, minutely reddish brown puberulent, bracts ca. 1 mm long and glabrous, flowers often in triads, pedicels 0-3 mm long. Flowers with hypanthium 0.7-1 mm long, obconic or turbinate and minutely (0.05 mm) papillate- puberulent or glabrous, calyx 0.3-0.5 m long, with mi- nute lobes or subentire distally; corolla white, salver- form, glabrous externally, tube 2-4 mm long and 1-1.5 mm diam., lobes 4, 1-2 mm long, triangular; stamens 4, anthers ca. 0.7 mm long. Fruits 4-6 mm long, 4-5 mm diam., ellipsoid to subglobose, becoming orange- red; pyrenes with 4-5 rounded ridges.
Understory plants of lower montane evergreen wet forest formations, from 1 200 to 2200 m ele- vation in Costa Rica. Probably flowering through- out the year (mostly in July-August); fruiting in October-May. The species ranges from central Costa Rica to western Panama.
Psychotria parvifolia is recognized by its small
flat-topped stature, very small leaves with few sec- ondary veins, puberulent stems, calyptrate stip- ules, and small few-flowered inflorescences. The tendency of the leaves to dry grayish beneath and the reddish fruit are characteristics of subgenus Psychotria. These plants are often conspicuous for their many horizontal branchlets and small leaves, especially in forest interiors where such small leaves are rare on woody plants. The Panamanian col- lections tend to have inflorescences with thinner axes, longer (3 mm) pedicels, and longer corolla tubes. Compare P. graciliflora with different stip- ules.
Psychotria phanaerandra (Standl. & Steyerm.) Lorence, Novon 2: 260. 1992. Palicourea phan- aerandra Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 252. 1947. Psychotria luteotuba Lorence, Bol. Soc. Bot. Mex. 47: 55. 1987.
Shrubs, 1-6 m tall, leafy stems 0.6-4 mm thick, gla- brous and becoming grayish (rarely minutely papillate- puberulent); stipules with a tube and 2 linear lobes ca. 0.5 mm long, deciduous. Leaves with petioles 2-8 mm long, 0.4-0.9 mm thick, glabrous, drying dark; leaf blades 4-11 cm long, 1.3-4.5 cm broad, elliptic to narrowly elliptic-oblong or lanceolate, apex tapering gradually and long-acuminate with tip 3-18 mm long, base attenuate, drying membranaceous to firmly chartaceous, grayish green to dark brown above, glabrous above and below, 2° veins 5-8/side. Inflorescences solitary and terminal, 3-5 cm long, 1.5-3 cm broad, cymose-paniculate with few opposite or alternate branches, peduncle 1 5-22 mm long, 0.4-0.7 mm thick, usually glabrous, braces 0.5-1 .4 mm long or absent, pedicels 0-2 mm long. Flowers gla- brous externally (rarely sparsely papillate-puberulent), hypanthium 0.5-0.8 mm long, 0.6 mm diam., calyx limb
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
265
ca. 0.5 mm long with minute (0. 1-0.2 mm) lobes; corolla white to lilac, pale purple or rose-lilac, tubular-funnel- form, tube (3-)4-8 mm long, ca. 1 mm diam., lobes 4, 2.5-5 mm long; stamens 4, anthers 1.8-2.5 mm long. Fruits 4-6 mm long, 5-6 mm diam., ovoid or globose- oblate, black; pyrenes with 3 prominent ridges.
Plants of wet evergreen forest on the Caribbean slope at 600-900 m elevation. It ranges from 20 to 2100 m elevation in Mexico and Guatemala. Flowering in March-June and December (throughout its range); fruiting in July-December and February. The species is also known from Mexico, Guatemala, and Honduras.
Psychotria phanaerandra is recognized by its small leaves, small slender few-flowered inflores- cences, lilac corolla, and smooth glabrous fruit be- coming black. Our material differs from northern collections by the more slender petioles, thinner translucent leaves with longer apices, lower-ele- vation wet forest habitat and lilac (not yellow or white) flowers. Compare with material placed in P. deflexa.
Psychotria pilosa Ruiz & Pav., Fl. Peruv. 2: 60. 1799. P. costaricensis Polak., Linnaea 41: 571. 1877. Figure 54.
bright blue or purple; pyrenes ca. 2.5 mm long with 3- 4 ridges.
Understory plants in evergreen rain forest for- mations of the Caribbean slope, General Valley, and Golfo Dulce area, from 20 to 900 (rarely to 1500) m elevation. Flowering in March-August; fruiting in August-March. This species ranges from Nicaragua to Peru.
Psychotria pilosa is recognized by its long pilose hairs on almost all surfaces, large two-lobed per- sisting stipules, large leaves with arcuate submar- ginal vein, densely flowered inflorescences, small flowers subtended by broad persisting bracts, and bright blue fruit. There is considerable variation in the form of the inflorescences and prominence of floral bracts among different individual plants. This species is rarely collected on the Pacific slope. Individual collections with conspicuous blue-vi- olet coloring on the leaf undersurfaces are occa- sionally collected but they appear to be no more than an unusual form. This species is one of our more distinctive species of Psychotria. Sterile col- lections may resemble P. mortoniana, P. micran- tha, and P. pittieri, but they have shorter pubes- cence and smaller bracts. Compare the closely similar Palicourea standleyana with differing flow- ers and fruits.
Shrubs or subshrubs, l-2(-3) m tall, leafy internodes 1.4-7 mm thick, terete, hirsute to pilose with erect or ascending thin hairs 1.3-2 mm long; stipules 8-25 mm long, 4-10 mm broad at the base, with 2 narrowly tri- angular or subulate lobes 4-12 mm long, sparsely pu- bescent distally, persisting. Leaves distant along the stem, petioles 12-40 mm long, 0.8-2.2 mm thick, densely pi- lose with thin straight or crooked hairs to 2 mm long; leaf blades ( 1 0-) 1 2-27 cm long, (3-)4- 1 3 cm broad, ovate- elliptic to elliptic-oblong or elliptic-obovate, apex grad- ually to abruptly acuminate or caudate-acuminate with tip to 2 cm long, base obtuse to cuneate and slightly decurrent on petiole, drying membranaceous to thin- chartaceous, dark olive green or brownish (rarely deeply blue-violet beneath), pilose above and below with hairs to 1.5 mm long, 2° veins 9-18/side, loop-connected dis- tally to form an arcuate submarginal vein. Inflorescences terminal or pseudoaxillary, solitary, becoming pendant, 5-14 cm long, 2-8 cm broad, congested-pyramidal to subcylindrical panicles with opposite branches and densely clustered flowers (or condensed and subcapitate), peduncles 1-8 cm long, 1-2.5 mm thick, densely pilose with yellowish brown hairs, bracts to 8 mm long and 4 mm broad, bracteoles 3-6 mm long, ovate and glabrous to pilose abaxially, flowers sessile and enclosed within the persisting bracts. Flowers monomorphic, pilose ex- ternally, calyx ca. 1 mm long, dentate; corolla funnel- form, white, sparsely puberulent externally, tube 3-6 mm long and 1.5 mm diam., lobes 5, 1-3 mm long. Fruits ca. 4 mm long (not including the 1 mm persisting calyx) and 3-5 mm diam., to 10 mm when succulent, pilose,
Psychotria pithecobia Standl., Publ. Field Col- umb. Mus., Bot. Ser. 8: 187. 1930. Figure 60.
Shrubs to l(-2) m tall, epiphytic (terrestrial), stems sometimes vine-like, leafy stems 1.5-6 mm thick, gla- brous and drying pale yellowish; stipules with a mem- branaceous tubular truncated sheath 2-4 mm long, ca- ducous or becoming indurated at base. Leaves with petioles 4-12 mm long, 1-1.8 mm thick, glabrous; leaf blades 4—12 cm long, 2-4.5 cm broad, elliptic-oblong to ovate-elliptic or elliptic, apex acute or short-acuminate with tip to 1 cm long, base acute to obtuse and slightly decurrent on petiole, drying grayish green to dark olive green, glabrous on both surfaces and with short (0. 1-0.2 mm) linear cystoliths that may resemble hairs, 2° veins 5-1 I/side and arising at angles of 40-60° from the mid- vein. Inflorescences terminal (rarely axillary) and soli- tary, 4-14 cm long, to 8 cm broad, open panicles with distant opposite branching and flowers in distal cymose groups, often red, becoming pendant, peduncles 2-6 cm long, 0.7-1.3 mm thick, glabrous, drying black, bracts 1.5-5 mm long and subulate to linear, pedicels 1-5 mm long. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx tube ca. 0.3 mm long, calyx lobes 0.2-0.7 mm long, triangular, obtuse or acute; corolla salverform, white or pink, tube 2-3.5 mm long, ca. 1 mm diam., lobes 4, 1-1.5 mm long. Fruits ca. 4.5 mm long and 3.5 mm diam., oblong to obovoid, orange or red at maturity.
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Plants of evergreen lower montane cloud forest formations, from (500-)800 to 2200 m elevation. Flowering in December-July; fruiting in every month. This species (as presently denned) ranges from the Cordillera de Tilaran in Costa Rica to the western highlands of Panama.
Psychotriapithecobia is recognized by its usually epiphytic habit, small tubular stipules (in early stages), succulent leaves with distinctive venation, large open reddened inflorescences, and small flowers with four-locular ovaries. The leaves dry- ing grayish and the red fruit are characters shared with species of subgenus Psychotria. Specimens placed here may be no more than a distinctive variety of P. guadalupensis (DC.) Howard. They differ by the large leaves tapering gradually to the apex and with a greater number of ascending sec- ondary veins clearly visible and elevated on the dried surfaces. The smaller corollas in specimens placed here are consistent with Steyermark's ( 1 972) description of P. guadalupensis in Venezuela but differ from Costa Rican material being called P. quadalupensis (q.v.). While these differences are distinctive in many collections, there may be a few intermediate collections. Costa Rican material placed under the names P. pithecobia and P. gua- dalupensis appears to have exactly the same eco- logical preferences and the same local distribution; this also suggests that they may represent different morphological forms of the same species.
Psychotria pittieri Standl., Contr. U.S. Natl. Herb. 1 8: 1 32. 1 9 1 6. P. dispersa Standl., J. Wash. Acad. Sci. 18: 184. 1928. Figure 55.
Shrubs or subshrubs, 0.5-1.5(-2.5) m tall, much- branched, leafy stems 1-3 mm thick, terete, sparsely to densely pubescent with thin whitish or yellowish hairs 0.2-1 mm long; stipules 3-7 mm long, with a tube 0-2 mm long, broadly ovate with 2 narrow awns 1.5-7 mm long separated by a U-shaped sinus, pubescent, persist- ing. Leaves with petioles 4-12(-22) mm long, 0.5-1 mm thick, sparsely to densely pubescent to hirsutulous; leaf blades 6-1 1(-15) cm long, 2-4(-5) cm broad, elliptic to elliptic-obovate or elliptic-oblong, apex acuminate with tip 5-14 mm long, base acute to attenuate and slightly decurrent on petiole, thinly chartaceous, drying brown- ish or green, glabrous or with short hairs along the mid- vein above, sparsely to densely puberulent on the veins beneath with hairs 0. 1-0.4 mm long, 2° veins 8-1 I/side. Inflorescences terminal or becoming pseudoaxillary, sol- itary, usually pendant, 3-10 cm long, 2-4 cm wide, py- ramidal open panicles, distal branching cymose, pedun- cles 1-3 cm long, 0.5-1 mm thick, hispidulous, bracts 4-8 mm long, linear, flowers mostly sessile, bracteoles ca. 2 mm long. Flowers minutely puberulent externally, hypanthium ca. 1 mm long and 0.8 mm diam., turbinate,
calyx lobes 0.2-0.5 mm long, acute; corolla white to yellowish or greenish, tubular-salverform, tube 2-4 mm long, 0.8-1.5 mm diam., lobes 5, 0.5-0.7 mm long. Fruits 3-4 mm long, 2.5-4 mm diam. (6 mm in life), ellipsoid or globose, minutely papillate-puberulent, becoming bright blue or blue-purple; pyrenes 2-3 mm long, with 4-5 ridges.
Plants of wet evergreen forest understory on the Caribbean slope and in the central Cordilleras, from near sea level to 1 100 m elevation. Flowering pri- marily in May-July; fruiting throughout the year (mostly in October-March). The species ranges from Belize and Guatemala to central Panama.
Psychotria pittieri is recognized by its small stat- ure, pubescent stems and peduncles, long stipular awns, smaller leaves, short inflorescences that are often pendant, small flowers, and bright blue fruit. It is one of our more common species. This species is similar to P. acicularis of the Osa Peninsula, with long slender bracts, and P. steyermarkii, which has more glabrous stems and peduncles and much larger bracts.
Psychotria platypoda DC., Prodr. 4: 510. 1830.
Shrubs or subshrubs, 1-3 m tall, leafy stems 1-3 mm thick, glabrous; stipules with a cupulate base 1-2 mm long and linear-lanceolate or oblong-lanceolate lobes 0.5- 2.5(-4) mm long (4/node), glabrous, usually persisting. Leaves with petioles 5-18 mm long, 0.8-1.8 mm thick, glabrous; leaf blades 8-19 cm long, 4-8 cm broad, nar- rowly elliptic to elliptic-lanceolate or narrowly ovate, apex tapering gradually and acuminate, base tapering gradually and slightly decurrent, drying chartaceous, ol- ive green above, glabrous or with few hairs along the midvein beneath, 2° veins 6-10/side and arcuate-as- cending. Inflorescences solitary and terminal, 12-25 mm long, 7-20 mm broad, capitulate or condensed cymose with 3-5 short branches, with 1 2-60 flowers, peduncles 3-12 mm long, ca. 1.5 mm thick, glabrous or minutely hirsute, conspicuous bracts 7-10 mm long, 6-14 mm broad, rounded distally, white tinged with green to pur- ple in color, subtending the branches of the inflorescence, flowers often in groups of 10-12 subtended by 1 larger and 2 smaller bracts. Flowers glabrous externally, hy- panthium ca. 0.8 mm long, calyx ca. 0.5 mm long, calyx teeth 0.1-0.2 mm long; corolla white, tubular-funnel- form, tube 3-6 mm long, lobes 5, 1.2-1.8 mm long, rounded distally. Fruits 3-5 mm long, 3-6 mm diam., blue to dark violet or black; pyrenes smooth with 3-5 planar ridges.
Plants of evergreen forest formations, 50-800 m elevation. Immature fruits were collected in No- vember, mature fruits in January. The species ranges from Costa Rica to Peru and eastern Brazil.
Psychotria platypoda is recognized by the broad colorful bracts that subtend the young compact
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
267
inflorescences. The small flowers, black or bluish fruits, glabrous leaves tapering gradually at both ends, and two-toothed stipules are additional characteristics. This species is quite variable in South America, and there is the possibility that hybridization occurs with P. officinalis or P. hoff- mannseggiana (q.v.). Rarely collected in Costa Rica; Chilamate de Sarapiqui, Valle del General, Rincon de Osa.
mentosd) is recognized by the usually shrubby habit, conspicuous pubescence, solitary terminal heads subtended by two bright red bracts, the yel- lowish corolla, and the blue fruit. This is a very common and conspicuous plant in the evergreen lowlands, but it is rarely collected above 1 000 m elevation in Costa Rica. It is one of the species that made Central American members of Cepha- elis seem so distinctive.
Psychotria poeppigiana Muell. Arg. in Mart., Fl. Bras. 6(5): 370. 1881. Tapogomea tomentosa Aubl., Hist. pi. Guiane 160, pi. 61. 1775, not Psychotria tomentosa Hemsl. Cephaelis tomen- tosa (Aubl.) Vahl, Ecolog. Amer. 1:19. 1796. C. hirsuta Mart. & Gal., Bull. Acad. Brux. (11) 1: 135. 1844. Evea tomentosa (Aubl.) Standl., Contr. U.S. Natl. Herb. 18: 123. 1916. Figure 17.
Shrubs or herbaceous subshrubs, 0.7-3(-6) m tall, leafy stems 1.5-6 mm thick, terete, pilose to villose with thin ascending or erect straight hairs ca. 2 mm long; stipules with a tubular sheath 2-8 mm long (but this often difficult to see or becoming torn) and linear awns 4-16 mm long, hirsute, the dark interior colleters to 3 mm long. Leaves with petioles 4-18(-30) mm long, 0.7-1.5 mm thick, hirsute with thin yellowish hairs; leaf blades 8-18(-24) cm long, 3-8(-l 1) cm broad, narrowly ovate-elliptic to lanceolate or elliptic-oblong, apex tapering gradually and acute or acuminate with tip to 20 mm long, base obtuse to acute or cuneate and slightly decurrent on petiole, drying thin-chartaceous or chartaceous, dark green above, with thin hairs 1-1.5 mm long on the surfaces (some- times breaking off near the base) and longer (to 2 mm) hairs along the midvein above and below, 2° veins 6- 1 2/side. Inflorescences terminal and solitary, the invo- lucrate capitula ca. 2 cm long and 3-8 cm broad, pe- duncles 1 .3-8(-l 3) cm long, ca. 1.5 mm thick and dense- ly villous or hirsute, the 2 opposite basal bracts ca. 3 cm long and 3-4 cm broad, united at the base, broadly ovate and acute or mucronate, bright red or orange red, flowers sessile and closely crowded. Flowers distylous, puberu- lent externally, hypanthium 1.5-2 mm long, calyx lobes 5, 0.6-2 mm long, with a linear gland in each sinus; corolla tubular, yellow or yellowish white, with ascend- ing hairs, tube 10-14 mm long, narrowly funnelform, lobes 5, 2-3 mm long, narrowly triangular; anthers 1.8- 2.5 mm long. Fruits 1-2 cm long, 5-10 mm diam., ob- ovoid or oblong, fleshy, bright blue; pyrenes 5-8 mm long, 1-2 mm broad, with 3-5 ridges.
Plants of forest shade and forest edges in ever- green rain forest formations, from near sea level to 800(-1 100) m elevation. Flowering throughout the year; probably also fruiting throughout the year. The species ranges from Mexico and Central America to Brazil and Bolivia.
Psychotria poeppigiana (formerly Cephaelis to-
Psychotria polyphlebia J. D. Smith, Bot. Gaz. 33: 253. 1902. Figure 13.
Herbs or small subshrubs, 1 5-50 cm tall, rhizomatous, usually with a single unbranched stem, leafy stems 1.5- 6 mm thick, glabrous or at first hirsute with hairs to 1 mm long, usually quickly glabrescent and drying black, terete; stipules 1-3 mm long, 4-6 mm broad, broadly triangular, obtuse, glabrous or puberulent, inconspicu- ous and deciduous. Leaves with petioles l-4(-7) cm long, 1-2.7 mm broad, sparsely puberulent to densely hirsute with hairs 0.5-1.5 mm long; leaf blades 7-16 cm long, 3-7 cm broad, oblong to obovate-oblong or obovate, apex bluntly obtuse to rounded, base cuneate to acute and decurrent on petiole, drying thin- to stiffly charta- ceous, grayish green to dark olive green above, glabrous or with stiff straight hairs 0.5-1 .3 mm long above, hirsute beneath with thin usually straight hairs, 2° veins 15-257 side and usually loop-connected close to the leaf margin, surface of the leaf becoming transversely corrugated. In- florescences solitary in leaf axils (usually I/node), 2-8 cm long, 15-30 mm broad, capitate-hemispheric, pe- duncles 1-6 cm long, 1-2 mm thick, glabrous or hirsu- tulous, lower bracts ca. 8 mm long and 1.5 mm broad, flowers sessile or subsessile, enclosed by subequal ovate to lanceolate bracts to 6 mm long and 3 mm broad. Flowers 5-parted, calyx minutely denticulate; corolla white, thin and translucent, tubular-funnelform, tube 4- 5 mm long, 1 mm diam., lobes 5, 1-1.5 mm long. Fruits 6-9 mm long, 4-6 mm diam., oblong, becoming orange or red and finally black, glabrous; pyrenes 5-7 mm long, with 3-5 ridges.
Plants of the deeply shaded forest floor in ev- ergreen rain forest formations primarily on the Caribbean slope, from 20 to 1200 m elevation. Flowering in February-August; fruiting through- out the year. The species ranges from Nicaragua to Colombia.
Psychotria polyphlebia is recognized by its small size, stiff corrugated leaves held horizontally over the dark forest floor, many secondary veins, the usually long-pedunculate capitate inflorescences from the axils of leaves, and small flowers hidden by bracts. The tendency of the undersides of leaves to dry grayish and the red or orange fruits are characters usually associated with subgenus Psy- chotria. This is one of the most distinctive species
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of Psychotria in Costa Rica; no other is so short in stature with such distinctive leaves.
Psychotria pubescens Sw., Prodr. 44. 1788. P. glauca Polak., Linnaea 41: 569. 1877. Figure 59.
Psychotria psychotriifolia (Seem.) Standl., Contr. U.S. Natl. Herb. 18: 133. 1916. Cephaelis psy- chotriaefolia Seem., Bot. voy. Herald 138. 1854. Figure 63.
Shrubs or subshrubs, 0.5-2(-3) m tall, leafy stems 1.5- 8 mm thick, at first densely puberulent with reddish brown or yellowish hairs ca. 0.2 mm long, glabrescent, terete; stipules with a truncate tube 4-14 mm long and 2-4 mm diam., awns 1-5 mm long, narrow, 2/side, mi- nutely puberulent along the midrib and awn, drying red- dish brown, usually splitting down 1 side and deciduous. Leaves with petioles 3-10(-20) mm long, 0.8-2 mm broad, minutely (0.05-0.2 mm) papillate-puberulent; leaf blades 5-1 5(-20) cm long, 2.5-7 cm broad, oblanceolate to narrowly obovate or elliptic-oblanceolate, apex acute or obtuse, base gradually narrowed and cuneate, decur- rent on petiole, drying chartaceous or stiffly chartaceous, grayish to pinkish or orange-brown, glabrous above, mi- nutely papillate-puberulent along major veins and with longer (0.2 mm) hairs distally and along the leaf margin, 2° veins 8-14/side and loop-connected near the margin to form a submarginal vein. Inflorescences terminal or pseudoaxillary, 1-4/node, 10-30 mm long, pyramidal panicles of glomerules or with 3 capitula on a primary peduncle (rarely with heads in an umbellate configura- tion), peduncles 3-10 mm long (to 18 mm in fruit), 0.8- 1 .8 mm thick, sparsely to densely puberulent, lower bracts biaristate (resembling the stipules) and 3-5 mm long, often forming an involucre, flowers sessile or subsessile and partly enclosed by the narrowly ovate bracteoles ca. 3 mm long. Flowers distylous, hypanthium and calyx ca. 2 mm long, calyx lobes ca. 1 mm long; corolla rotate, pale green or white, tube 1-2 mm long, lobes (4-)5, 1- 1.7 mm long and 0.6 mm broad at base; stamens 5, anthers 0.5-0.7 mm long. Fruits 5-6 mm long (not in- cluding the 0.5-2 mm long calyx), 3-4 mm diam., ellip- soid, longitudinally grooved, glabrous and becoming bright red; pyrenes ca. 4.5 mm long, with 3-5 ridges.
Plants of Caribbean lowland evergreen forest formations, from near sea level to 600(-800) m elevation. Flowering and fruiting throughout the year (flowering February-September in Costa Rica). The species ranges from southern Nicaragua to Colombia and Venezuela.
Psychotria psychotriifolia is recognized by the slightly oblanceolate leaves with arcuate submar- ginal veins, puberulent young stems, tubular bi- aristate stipules, small flowers in dense bracteate or involucrate capitula, ellipsoid fruit with per- sisting calyx, and lowland habitat. The inflores- cences often become pseudoaxillary. The tendency of vegetative parts to dry reddish brown or gray and the red fruit are characteristics of subgenus Psychotria.
Shrubs, l-2(-3) m tall, leafy stems 0.7-4 mm thick, minutely papillate-puberulent to hirtellous with hairs 0.2- 0.8 mm long or sometimes glabrous; stipules 3-8 mm long, 2-4 mm broad at the base, with a truncate to tri- angular basal portion ca. 2 mm long and 2 narrow distal lobes l~4 mm long (4/node). Leaves with petioles 4-28 mm long, 0.5-1.5 mm thick, with adaxial margins con- tinuous with the lamina margins, usually minutely pa- pillate-puberulent; leaf blades (5-)7-15(-18) cm long, (1.3-)2-5(-7) cm broad, ovate-elliptic, to elliptic-lan- ceolate or elliptic-oblong, apex acuminate to acute (ob- tuse) with a tip 3-10 mm long, base cuneate to acute (obtuse) and decurrent on petiole, drying membrana- ceous to thin-chartaceous and dark green, sparsely and minutely (0. 1-0.2 mm) puberulent or subglabrous above and below, 2° veins 9-12(-15)/side, weakly loop-con- nected near the margin distally. Inflorescences terminal and solitary (or 3), 2-6 cm long, 1.5-5 cm broad, com- pact (at first) to open-corymbose panicles with opposite branching, peduncles 6-30 mm long, ca. 1 mm thick, densely puberulent to glabrous, bracts 2-6 mm long, linear, flowers 2-3(-5) in cymes or triads, pedicels 0-2 mm long. Flowers usually very minutely (0.05 mm) pa- pillate-puberulent externally, hypanthium ca. 0.6 mm long and 0.5 mm diam., calyx tube ca. 0.2 mm long and 1 mm broad with lobes 0.4 mm long; corolla funnelform, white or yellowish, tube 2-4 mm long, 0.8-1 .8 mm diam., slightly constricted in the middle, lobes 5 or 6, ca. 1.5 mm long. Fruits 5-6 mm long and 5-6 mm diam., oblate to globose, with prominent longitudinal ridges, purple or black.
Plants of shaded sites in deciduous or partially deciduous forest formations, from 20 to 1200(-1500) m elevation on the Pacific slope of central and northern Costa Rica. Flowering in May-August; fruiting in July-January. This spe- cies is found from Mexico to Panama and in the West Indies.
Psychotria pubescens is recognized by the usual presence of minute hairs on young stems and in- florescences, short two-awned stipules, thin leaf blades with decurrent base, small flowers, and sea- sonally dry habitat. Despite the specific name, some plants may be nearly completely glabrous. Like- wise, there is great variation in leaf form between different plants, from small and lanceolate to larger and more obovate or oblong. The inflorescences are usually rounded distally at anthesis. This spe- cies can be confused with P. microdon, which also lives in deciduous vegetation but has longer peri- anth parts, red fruits, and a more clambering habit.
Psychotria quinqueradiata Polak., Linnaea 4 1 : 570. 1877. Mapouria obovata Oerst., Amer. Centr.
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17, t. 14, f. 3 and 4. 1863. P. morae Polak., loc. cit. 570. 1877. P. obovata (Oerst.) Hemsl., Biol. centr. amer. Bot. 2: 50. 1881, non P. obovata Ruiz & Pav. 1799. Figure 65.
Shrubs, 0.5-2.5 m tall, leafy stems 1.2-4 mm thick, glabrous, older twigs becoming pale gray and often with whitish lenticels; stipules 4-6 mm long and 2-5 mm broad but to 15 mm long and 8 mm broad below the inflorescences, lanceolate to narrowly ovate or broadly obovate beneath the inflorescences, acute, glabrous and drying dark reddish brown, usually caducous. Leaves with petioles 1.5-6(-10) mm long, 1-2 mm thick, gla- brous; leaf blades (3.5-)5-13(-15) cm long, (1.5-)2- 5.5(-7) cm broad, elliptic-obovate to obovate, oblan- ceolate, elliptic or subpandurate, apex acute to short- acuminate with tip 3-5 mm long, base cuneate or acute and decurrent on petiole or the blade sometimes rounded and subcordate at base, drying thin-chartaceous or char- taceous, grayish or pinkish brown, glabrous above and below, 2° veins 5-7/side, often with minute pit domatia or tufts of hairs in the vein axils beneath. Inflorescences terminal and 1-5, 1-3 cm long and elongating to 5 cm in fruit, to 1.5 cm wide, subcapitate or densely cymose to umbellate, peduncles 2-35 mm long, 0.4-1 mm thick, glabrous, bracts and bracteoles not evident, flowers in distal triads, diads or glomerules of 5-15, pedicels 0-1 mm long. Flowers glabrous externally, hypanthium ca. 1 mm long, calyx tube ca. 0.4 mm long, lobes slightly dentate; corolla white, funnelform, tube 3.5-5 mm long, 1-2 mm diam., lobes 5, 1.5-3 mm long; stamens 5, anthers 1-1.5 mm long. Fruits 6-9 mm long, 4-6 mm diam., oblong or ellipsoid, bright red or orange; pyrenes ca. 4 mm long and 3 mm broad, with 3-5 low ridges.
Plants of forest understory in both seasonally deciduous woodland and wet evergreen rain forest formations, from near sea level to 1400 m ele- vation. This species is rarely collected in the Ca- ribbean lowlands of Costa Rica; it forms a dis- tinctive understory in parts of Parque Nacional Rincon de la Vieja. Flowering throughout the year (primarily in January-July); fruiting in August- February in Costa Rica. The species ranges from Veracruz, Mexico, southward through Central America to western Panama.
Psychotria quinqueradiata is recognized by the usually short-petiolate and obovate leaves, the broad stipules below the young inflorescences, the capitate to umbellate- fasciculate inflorescences, and subsessile flowers. The tendency of the leaves to dry gray or reddish and the bright red fruit are characteristics of subgenus Psychotria, The stip- ules are often much enlarged below the young in- florescences. As in some other species of Psycho- tria, the petioles and leaf bases can vary greatly in different individuals: from subsessile to longer- petiolate and from decurrent to slightly auriculate. This species resembles P. carthaginensis but differs
in venation and inflorescences. Compare also P. venosa and P. chagrensis.
Psychotria racemosa (Aubl.) Raeuschel, Nomen. ed. 3, 56. 1797. Nonatelia racemosa Aubl., Hist, pi. Guiane 180, pi. 72. 1775. P. racemosa (Aubl.) Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792 fide Dwyer 1980. Rudgea thyrsiflora J. D. Smith, Bot. Gaz. 61: 375. 1916. Figure 59.
Shrubs or herbaceous subshrubs, 0.5-2(-4) m tall, leafy stems 1.5-5 mm thick, sparsely to densely puberulent with ascending hairs 0.1-0.3 mm long; stipules with a short (1-3 mm) sheath and 2 long (6-14 mm) narrow (0.3-0.6 mm) awns on each side (4/node), with indurated glabrous yellowish colleters ca. 1.5 mm long interior to the sheath, puberulent and persistent. Leaves with pet- ioles 5-20(-35) mm long, 0.9-1.5 mm thick, densely puberulent (except for the adaxial midvein); leaf blades 9-21 (-26) cm long, 3-7 (-1 1) cm broad, elliptic to ellip- tic-oblong or ovate-oblong, apex acuminate with tip 5- 1 5 mm long, base obtuse to acute and slightly decurrent on petiole, drying chartaceous, green to brownish green, glabrous above, minutely (0. 1-0.2 mm) puberulent along the veins beneath, 2° veins 7-12/side, the 3° veins sub- parallel and prominent beneath. Inflorescences terminal or becoming axillary, l(-2)/node, 3-7 cm long and 1.5- 2.3 cm broad (to 10 x 4 cm in fruit), pyramidal or hemispheric panicles with usually short (often distinctly cymose) lateral branches, peduncles 1-2 cm long and 1.5 mm thick, densely yellowish puberulent, bracts to 6 mm long, linear, bracteoles ca. 2 mm long, pedicels 0-2 mm long. Flowers minutely puberulent on the exterior, hy- panthium 0.8-1 mm long, obconic, calyx ca. 0.4 mm long, lobes ca. 0.3 m long and obtuse; corolla tubular- salverform, white, greenish or yellow, tube 1-3 mm long, 0.6-1.2 mm diam., lobes 4 or 5, 0.5-1 mm long. Fruits 4-5 mm long, 5-7 mm diam., oblate to globose, glabrous, becoming orange or red and finally purple or black, fleshy; pyrenes usually 5, ca. 4 mm long, with 3 sharp dorsal ridges.
Understory plants in evergreen lowland forest formations on both the Caribbean and Pacific slopes, from near sea level to 800 m elevation (rarely found above 400 m on the Caribbean slope). Flowering in March-July; fruiting throughout the year in Costa Rica. This species is found through- out the Caribbean lowlands in Costa Rica and in evergreen forests of the Pacific slope of southern Costa Rica. The species occurs in southern Mexico and ranges from Nicaragua to Brazil and Bolivia.
Psychotria racemosa is recognized by the minute puberulence on younger stems and leaf undersides, the persisting stipules with long stiff narrow awns, subparallel tertiary veins, relatively short puber- ulent inflorescences, small puberulent flowers, five- locular ovaries, and fruit with usually five seeds.
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FIELDIANA: BOTANY
The five-locular gynoecium is unique among Cen- tral American species of Psychotria.
Psychotria remote Benth., J. Bot. 3: 225. 1841. Mapouria remota (Benth.) Muell. Arg., Flora 59: 459. 1876, and in Mart., Fl. Bras. 6(5): 407. 1881. P. alboviridula K. Krause, Notizbl. Bot. Gart. Berlin 6: 208. 1914. Figure 66.
Shrubs or small treelets, 1.5-5(-8) m tall, leafy stems 1 .5-5 mm thick, glabrous and drying dark reddish brown; stipules 3-6 mm long, 1-3 mm broad at base, triangular, tip acute or acuminate (biacuminate), glabrous, cadu- cous. Leaves with petioles 5-24 mm long, 1-2 mm thick, glabrous, often with prominent adaxial margins forming a slight sulcus; leaf blades 10-24 cm long, 3-7.5(-9) cm broad, elliptic to elliptic-oblong or narrowly ovate-ob- long, apex acuminate with tip 4-1 3 mm long, base acute or obtuse and slightly decurrent on petiole, drying stiffly chartaceous or subcoriaceous, dark grayish or reddish brown, glabrous above and below, 2° veins 7-10/side, axils of the veins often with pit domatia 0.3-1 mm long beneath. Inflorescences terminal or becoming axillary, 6-17 cm long, 4-10 cm broad, open panicles with 2-5 pairs of lateral branches, peduncles 2-7 cm long, 0.9-2 mm thick, glabrous and drying dark reddish brown, bracts 1-3 mm long, subulate to triangular, flowers sessile or subsessile in distal cymes of 3 or clusters. Flowers gla- brous externally, hypanthium 0.8-1.5 mm long, turbi- nate, calyx tube ca. 0.5 mm long, subtruncate or with lobes to 0.3 mm long; corolla campanulate, white or yellowish green, tube 1.5-2 mm long, 1-1.5 mm diam., lobes 5, 1-1.2 mm long, triangular; stamens 5, anthers 0.3-0.6 mm long. Fruits 7-9 mm long, 4-6 mm diam., ellipsoid or oblong, becoming orange to maroon red; pyrenes with 4-5 shallow ridges.
Plants of evergreen rain forest formations along the Caribbean lowlands, from near sea level to 200 m elevation (to 1000 m in Panama). Flowering in March-April and September-October; fruiting in March-May, July, and September. The species ranges from Costa Rica to Peru and Brazil.
Psychotria remota is recognized by its glabrous parts often drying dark in color, stiff leaves usually with conspicuous pit domatia beneath, small flow- ers on inflorescences with slightly winged branch- es, and flowers and fruits usually disposed at 90° to each other. The first branch pair of the inflo- rescences is usually much larger than the others. The tendency of leaves to dry grayish or reddish brown and the red fruit are characteristics of sub- genus Psychotria. This species is rare in Costa Rica. None of our other species of Psychotria have such large pit domatia, but these are not always present. Compare P. panamensis with very different stip- ules.
Psychotria rosulatifolia Dwyer, Ann. Missouri Bot. Gard. 67: 423. 1980. Figure 65.
Small treelets, ca. 3 m tall, leafy stems 3-6 mm thick, glabrous and grayish; stipules 8-16 mm long, 5-8 mm broad at the base, triangular to ovate, bifid and acute with a distal sinus 0-1 mm deep, glabrous, caducous. Leaves sessile or subsessile, petioles 0-7 mm long, ca. 3 mm thick, glabrous, drying dark; leaf blades 1 5-32 cm long, 4-11 cm broad, very narrowly elliptic to narrowly elliptic-oblong or lanceolate-elliptic, apex acute to sub- acuminate, base rounded and subcordate (slightly auric- ulate), drying stiffly chartaceous, dark grayish, glabrous above and below except for minute (0.1-0.2 mm) thin hairs along the midvein beneath, 2° veins 18-22/side and arising at angles of 90-100° from the midvein (to the apex), united by a prominent submarginal vein 1-2 mm from the leaf edge. Inflorescences terminal and sol- itary, ca. 38 cm long and 30 cm broad, an open pyramidal panicle with widely spaced opposite branches, with 2 very short and 2 long lateral branches at the first node, peduncle 5-18 cm long and 2 mm thick, sparsely and minutely puberulent, bracts ca. 3 mm long, flowers in cymules or 3-7, pedicels 1-3 mm. Flowers subglabrous externally, buds yellow, hypanthium 0.5 mm long and 0.3 mm diam., calyx tube 0.2-0.3 mm long, lobes 0.1- 0.2 mm long; corolla tubular, white, tube 3-6 mm long, lobes 3—4 mm long. Fruits 5-6 mm long, 3-4 mm diam., ellipsoid, red; pyrenes with 4-5 long ridges.
Plants of the wet Caribbean slope of the south- ern Cordillera de Talamanca, collected at ca. 700 m elevation (Herrera 3306 CR, MO) with immature flowers in July. The species is known only from this collection in Costa Rica and several collec- tions from Darien, Panama.
Psychotria rosulatifolia is unusual because of its long narrow subsessile (sometimes auriculate) leaves, secondary veins arising at angles of 90- 100°, well-developed submarginal vein, and large open inflorescences with two small and two long lateral branches at the first node. The leaves are sometimes disposed in distal rosettes.
Psychotria sarapiquensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1360. 1938. Figure 62.
Shrubs or small trees, 2-7 (-9) m tall, leafy stems 1- 4(-6) mm thick, glabrous and often drying grayish green; stipules 5-12(--45) mm long, 2-5 mm diam., calyptrate, narrowly conical to lanceolate, acute at the apex, gla- brous, drying reddish brown, caducous. Leaves with pet- ioles 9-35(-45) mm long, 0.8-1.8 mm thick, glabrous; leaf blades 6-1 2(-l 5) cm long, 2-4(-7) cm broad, ovate- elliptic to narrowly elliptic or lanceolate, usually apex tapering gradually and acuminate or acute with tip 3-14 mm long, base acute to cuneate and decurrent on petiole, drying stiffly chartaceous, grayish green to dark green above, glabrous above and below, 2° veins 6-8/side. In- florescences terminal 1 or 3 (2-)4-13 cm long, to 10 cm
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broad, open pyrimidal panicles with opposite branching or 3 main branches, peduncle (l-)2— 4 cm long, 0.8-1.9 mm thick, glabrous, bracts 0.5-1 mm long, broad, flow- ers in distal umbels or clusters of 3-5, pedicels 1-3 mm long. Flowers glabrous externally, hypanthium ca. 1.3 mm long, obconic, calyx 0.5-1 mm long, ca. 2 mm broad at the truncated distal margin; corolla funnelform green- ish white, tube l-3(-4) mm long and 2 mm diam., lobes 5, 1.5-3 mm long and 1.3 mm broad at the base, tri- angular; stamens 5, anthers 1.2-1.5 mm long. Fruits 7- 10 mm long, 4-8 mm diam., obovoid-oblong, becoming yellow then bright red; pyrenes with 3-5 rounded ridges.
Plants of lower montane forest formations of the Caribbean slope and near the continental di- vide, from (600-)1200 to 1600 m elevation (400- 900 m in Mexico). Flowering in January, March, July-August (Skutch 3330 us the type), and Oc- tober; fruiting in January-March and August. The species range is disjunct; it occurs in Veracruz, Mexico, and in the west-central cordilleras of Cos- ta Rica (Cordillera de Tilaran to Vara Blanca de Sarapiqui) and the western half of Panama.
Psychotria sarapiquensis is recognized by its long- petiolate narrowly elliptic leaves, glabrous parts, calyptrate caducous stipules enclosing the shoot apices, entire calyx, short corolla tubes, longer fruit, and restricted habitat (in Costa Rica). The ten- dency for the leaves to dry grayish and the fruit turning bright red are characteristics of subgenus Psychotria. The larger leaf dimensions come from material collected in Mexico. This species is close- ly related to P. panamensis, with similar Ficus- like stipules, but P. sarapiquensis has smaller leaves with fewer secondary veins. It can also be confused with P. chiriquina, P. orosiana, and P. tenuifolia.
Psychotria siggersiana Standl., J. Wash. Acad. Sci. 15: 289. 1925. P. morii Dwyer, Ann. Missouri Bot. Gard. 67: 400. 1980.
Herbs or succulent-stemmed subshrubs, 0.5-1 (-2) m tall, leafy stems 4-12 mm thick and usually unbranched, glabrous or sparsely puberulent and glabrescent, succu- lent and terete; stipules 3-7 mm long, ca. 8 mm broad, with a short collar around the stem, rounded deltoid, with a deciduous conical lateral appendage 1-2 mm long, usually glabrous, persistent. Leaves with petioles 4.5-10 cm long, 1.7-4 mm thick, typically with thin crooked hairs to 1 mm long (glabrous); leaf blades (12-)20-35 cm long, (6-)l 1-17 cm broad, ovate-elliptic to elliptic- oblong, apex acuminate with tip 5-10 mm long, base obtuse or cuneate, drying thin-chartaceous, dark brown- ish green above, with thin crooked hairs to 1.2 mm long on the upper surface, the hairs mostly shorter and denser beneath, 2° veins 8-1 I/side. Inflorescences axillary, sol- itary at a node, 3-15 cm long, pyramidal panicles with open opposite branching, peduncles l-3(-4) cm long,
1.3-2 mm thick, puberulent, bracts 5-8 mm long and 3 mm broad at the base, lanceolate to elliptic, flowers 3- 7 within distal glomerules and sessile, enclosed by broad bracteoles. Flowers with calyx 0.5-1 mm long, lobes del- toid-ovate; corolla funnelform, white or greenish white, tube ca. 3 mm long, sparsely short villous or glabrous externally, lobes 5, ca. 1 mm long. Fruits 5-6 mm long, globose, red; pyrenes ca. 4 mm long, dorsal surface smooth.
Plants of the wet lowland Caribbean rain forest formations at 100-1200 m elevation. Flowering in January, March (Standley 37210 us the holo- type), and August; fruiting in January, March, and October-November. This rarely collected species is known only from the Caribbean slope and low- lands of Costa Rica, western Panama, and Choco, Colombia.
Psychotria siggersiana is recognized by the short usually unbranched habit, large leaves with long hairs, axillary inflorescences with larger bracts broadly sheathing the distal floral glomerules, and the red fruits. Standley distinguished P. sigger- siana primarily by its pubescence, but the type also has unusually large floral bracts (as does P. aggregata, which looks quite similar). The red fruits with pyrenes that lack costae on their abaxial sur- face are distinctive, but specimens with young fruits can be difficult to separate from other members of the P. aggregata-macrophylla species alliance.
Psychotria sixaolensis C. Hamilton, Phytologia 64: 234. 1988. Figure 64.
Shrubs or subshrubs, 1^4 m tall, leafy internodes 4- 9 mm thick, young stems pubescent with reddish brown hairs 0.2-0.8 mm long but quickly becoming completely glabrous; stipules 10-25 mm long and 6-12 mm broad, ovate in outline and sheathing at base, acutely lobed for '/4-%, caducous. Leaves with petioles 10-35(-50) mm long, ca. 2 mm diam., poorly distinguished from the lamina base, puberulent; leaf blades (14— ) 16-3 Ocm long, (5.5-)8-14 cm broad, obovate to elliptic-obovate, ellip- tic or elliptic-oblong, apex abruptly narrowed and short- acuminate (obtuse) with tip ca. 5 mm long, base cuneate to acute, long-decurrent on petiole, drying chartaceous, dark reddish brown, glabrous above, with dense reddish brown hairs 0.3-0.7 mm long along the major veins, reddish brown tomentulose beneath, 2° veins 20-33/side, a well-defined slightly arcuate submarginal vein 1-2 mm from the edge. Inflorescences terminal and solitary, 5- 1 7 cm long, subcapitate or with 3 capitulae on short (5 mm) secondary branches, fruiting peduncles 7-14 cm long, 3-4.5 mm thick, bracts and bracteoles ca. 7 x 6 mm, ovate, reddish brown puberulent with hairs ca. 0.3 mm long, flowers sessile and densely crowded in distal glomerules of 8-20. Flowers puberulent externally, calyx tube 1-2 mm long, lobes 5, 2-3 mm long and 0.8 mm wide; corolla tubular, white or greenish white, tube ca.
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4 mm long and 2.5 mm diam., lobes 5, ca. 3 mm long and 1.5 mm broad, thickened near apex; stamens 5, anthers 1.3 mm long. Fruits 7-10 mm long (not including the calyx), 5-6 mm diam., ellipsoid with a persisting calyx 2-3 mm long, red at maturity; pyrene with 4-5 sharp ridges.
Plants of lowland Caribbean rain forest for- mations, from near sea level to 500 m elevation. Flowering in April and August; fruiting in August. This species is known only from southern Limon Province and Bocas del Toro Province in Panama.
Psychotria sixaolensis is recognized by its dense- ly pubescent leaves and inflorescences, large leaves with many secondary veins and well-developed submarginal vein, long-pedunculate subcapitate panicles with large ovate bracts, and flowers in dense glomerules. The red fruit and leaves drying reddish brown are characteristics of subgenus Psy- chotria.
Psychotria solitudinum Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 207. 1940. Figure 57.
Shrubs or small trees, 2-6 m tall, leafy stems 2-6 mm thick, glabrous, usually drying yellowish green; stipules 2-6 mm long, 2-4.5 mm broad, oblong to ovate, rounded with 2 short (0.5-1 mm) lobes separated by a small sinus, glabrous, persisting. Leaves with petioles 7-35(-50) mm long, 1-2 mm thick, glabrous, drying yellowish; leaf blades (7-) 10-21 (-30) cm long, (2.5-)4.5-10(-14) cm broad, obovate-oblong to ovate-elliptic, elliptic-oblong or ob- long, apex acuminate with tip 5-1 3 mm long, base acute to broadly obtuse (slightly round) and often slightly de- current on petioles, drying chartaceous or stiffly char- taceous, grayish green above, glabrous above, glabrous or with few thin hairs 0.2-0.6 mm long along the sides of the midvein beneath, 2° veins 12-18/side, arising at nearly 90° from the midvein, distally loop-connected to form an arcuate submarginal vein. Inflorescences soli- tary and terminal, 8-26 cm long, 10-20 cm broad, open pyramidal panicles with distant opposite lateral branches and distal cymose or dichotomous branches, peduncles to 8 cm long and 2 mm thick, usually glabrous, bracts 3-5 mm long, bracteoles 2-3 mm long, flowers mostly sessile, separated or in glomerules of 2-3. Flowers with thin white hairs 0.2-0.5 mm long on the outer surfaces (less often glabrous), hypanthium ca. 1 mm long, calyx tube ca. 0.5 mm long, lobes poorly differentiated; corolla funnelform-urceolate, pale green to yellow (white), tube 4-5 mm long, 0.7-1.5 mm diam. near base and 2-3 mm distally, lobes ca. 1 mm long, with distal projections in early stages. Fruits ca. 4 mm long (not including calyx) and 4-5 mm broad, oblate to obovoid or subglobose, bluish, purplish, or black in final stages, usually puber- ulent, persisting calyx ca. 1 mm high; pyrenes with 4-5 ridges.
Plants of wet evergreen forest formations of the Pacific slope (in Costa Rica), from 30 to 800 m
elevation. Flowering in January-August; fruiting in March-September. This species ranges from south-central Costa Rica to Colombia.
Psychotria solitudinum is recognized by its larg- er leaves with many secondary veins and a sub- marginal vein, ovate stipules with a small distal sinus, large open inflorescences with sessile flowers usually borne along dichotomous distal branches, puberulent yellow flowers, and small puberulent fruit. The leaves are subglabrous and tend to dry yellowish green. This species, which is restricted to the Pacific side of southern Costa Rica, appears to be closely related to P. angustiflora, P. branchia- ta, and P. tapantiana (q.v.). Specimens of P. mi- crobotrys and P. brachiata with young flowers are also similar.
Psychotria steyermarkii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 387. 1940. Figure 55.
Small shrubs, (0.5-) 1-2 m tall, leafy stems 0.5-3.5 mm thick, with thin whitish hairs 0.1-0.3 mm long often in longitudinal ridges or rarely glabrous; stipules with a sheath 1-4 mm long, truncated and with 2 filiform or narrowly lanceolate awns 2-5 mm long, puberulent at the base, usually persisting. Leaves with petioles 4- 1 4(-20) mm long, 0.4-0.8 mm thick, glabrous or minutely pu- berulent; leaf blades 3.5-1 1(-14) cm long, l-3(-3.5) cm broad, narrowly elliptic to narrowly elliptic-oblong or elliptic-lanceolate, apex tapering gradually and acumi- nate, tip 3-8 mm long, base acute and decurrent on petiole, drying chartaceous, dark greenish or brownish green above, glabrous or with a few hairs along the mid- vein above, minutely (0. 1-0.2 mm) puberulent along the major veins beneath or glabrescent, 2° veins 9-14(-17)/ side, loop-connected near the margin. Inflorescences ter- minal, usually solitary, 3-9 cm long, 2-3 cm broad, open pyramidal panicles with 2-3 pairs of lateral branches, peduncles 1.4-3 cm long, ca. 0.7 mm thick, puberulent, bracts and bracteoles 3-8 mm long, to 4 mm broad, ovate-lanceolate and forming a persisting involucre, sometimes marked with blue, glabrous, flowers sessile in glomerules of 2-5. Flowers glabrous or minutely pu- berulent externally, hypanthium and calyx obscured by the bracts and bracteoles, calyx lobes 0.4-1 mm long; corolla salverform, white or greenish white, tube ca. 3 mm long, ca. 0.7 mm diam., lobes ca. 1.5 mm long. Fruits 3.5-5 mm long, 3.5-5 mm diam., subglobose, bright blue; pyrenes with 3-5 planar angles.
Plants of evergreen lower montane forest for- mations and cloud forests, from 800 to 1800 m elevation. Flowering in May-August; fruiting in August-September and November-February in Costa Rica. The species ranges from the Cordillera de Tilaran to the Chiriqui Highlands and is found in southern Mexico, Guatemala, and Nicaragua.
Psychotria steyermarkii is distinguished by its
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small leaves, cloud forest habitat, tubular awned stipules, small racemose inflorescences, and small flowers in distal bracteate clusters. The fruits be- come bright blue and very attractive. This species has been mistaken for P. graciliflora of subgenus Psychotria with bright red fruits and P. pittieri with larger leaves and lower-elevation habitat. This species may also be mistaken for P. goldmanii of similar habitat with longer narrow leaves and dif- ferent stipules. The Guatemalan type (Steyermark 33700 F) has unusually large leaves, but a smaller- leaved fruiting collection from Guatemala (Stey- ermark 33461 F) is very similar to Costa Rican material. Herbarium material annotated P. cor- nigera Dwyer (ined.) is this species. Compare P. hazenii.
Psychotria stock wellii Hamilton, Phytologia 64: 235. 1988. Figure 62.
Small trees or shrubs, 2-10 m tall, leafy stems 2-6 mm thick, glabrous or sometimes pilosulous, drying dark; stipules 9-1 5 mm long, 3.5-7(-9) mm broad, ovate, acute and usually bilobed with a short sinus (1 mm), glabrous, caducous. Leaves with petioles 5-25 mm long, 1-3 mm broad, glabrous; leaf blades 8-20 cm long, 2.5-8 cm broad, elliptic-obovate to elliptic or oblanceolate, apex acuminate with tip 5-10 mm long, base gradually nar- rowed and cuneate or acute, decurrent on petiole, drying chartaceous or thin-chartaceous, very dark above, gla- brous on both surfaces (or the midvein minutely puber- ulent beneath), 2° veins 10-15/side, often with a weaker 2° between the major 2° veins and parallel with them. Inflorescences terminal (pseudoaxillary), solitary, 9— 18(-24) cm long, 5-12 cm broad, open pyramidal pan- icles with opposite 1°, 2°, and 3° branches usually borne at angles greater than 90° from the apex, peduncles 5-9 cm long, ca. 2 mm thick and glabrous, bracts and brac- teoles 0.5-2 mm long, triangular, flowers in distal cymes or triads, pedicels 0.5-2 mm long, minutely papillate- puberulent. Flowers with hypanthium 0.5-1 mm long, obconic, calyx tube 0.3-0.8 mm long, lobes 5, 0.3-0.7 mm long, minutely puberulent distally; corolla tubular- campanulate, cream white or greenish white, glabrous externally, tube 4-5 mm long, 1.5-2 mm diam., lobes 5, ca. 2 mm long and 1 mm broad at the base; stamens 5, anthers 1-1.2 mm long. Fruits 5-6 mm long, 4-6 mm diam., ellipsoid to oblate, red; pyrenes with 4-5 ridges.
Plants of montane rain forest formations, from ( 1000-)! 700 to 2300(-2700?) m elevation. Flow- ering in January-October; fruiting in October and January-April. The species is known in Costa Rica only from the northwestern part of the central highlands (Monteverde to Vara Blanca). It is also found in the Chiriqui Highlands of western Pan- ama.
Psychotria stockwellii is recognized by the nar-
rowly obovate leaves that dry dark above, many secondary veins, broadly ovate stipules, larger in- florescences with many orders of opposite branch- es, and higher-elevation habitat. The leaves drying pinkish gray or dark reddish gray beneath and the bright red fruit are characteristics of subgenus Psy- chotria. The type has the 1° branches of the inflo- rescence arising at angles of more than 1 00°, but this may have been a negatively geotropic response on a pendant branch. Recent collections from Pan- ama suggest that this species may intergrade with P. sylvivaga (q.v.). Compare P. chiriquina, which has inflorescences with two unequal pairs of branches (4/node) at the first node.
Psychotria subsessilis Benth. in Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 34. 1853.
A badly damaged isotype (Oersted 11613 us) gives the impression that this may be an earlier name for P. panamensis. It does not appear to be P. grandis as annotated by Urban in 1913 and cited by Standley in 1938. It comes from an ele- vation in which the transition between the two subspecies of P. panamensis may occur. However, those subspecies are quite different and may be worthy of specific rank. Thus, there is both a no- menclatural problem and one of species delimi- tation here. The name and collection were not included in the monograph by Hamilton (1989).
Psychotria suerrensis J. D. Smith, Bot. Gaz. 27: 337. 1899. Figure 18.
Shrubs, l-3(-4) m tall, leafy stems 1-5 mm thick, glabrous or minutely (0.1-0.2 mm) puberulent and gla- brescent, often drying orange or reddish, terete; stipules with a short (1-3 mm) truncate sheath and 2 narrow lobes or awns 3-6 mm long on each side, glabrous or minutely puberulent, persisting. Leaves with petioles 3— 8(-12) mm long, 1-1.8 mm thick, glabrous or very mi- nutely (0.1 mm) puberulent; leaf blades 10-20(-25) cm long, 3-8(-l 1) cm broad, elliptic to ovate-elliptic or ovate- oblong, apex acuminate with tip 5-20 mm long, base obtuse or acute and slightly decurrent on petiole, drying chartaceous to stiffly chartaceous, greenish to brown above, glabrous above, sparsely and minutely puberulent on the veins beneath with thin whitish hairs 0. 1-0.2 mm long, 2° veins 6-1 I/side, the larger veins often drying pinkish red beneath. Inflorescences terminal and solitary (or 3), 2.5-4 cm long, 3-5 cm broad, densely bracteate hemispheric (subcapitate) cymes or panicles (sometimes subumbellate), peduncles 7-28 mm long, 1 .3-2 mm thick, minutely puberulent, bracts 12-20(-32) mm long, 3- 6(-14) mm broad, oblanceolate to elliptic-obovate, green
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to rose or dark purple in fruit, flowers sessile and hidden within bracts. Flowers distylous, with thin whitish hairs externally, hypanthium ca. 1 mm long, turbinate, calyx tube 0.3-0.8 mm long, lobes 0.3-0.5 mm long; corolla tubular-funnelform, white or white with pinkish coloring (violet), usually drying pinkish, tube 9-13 mm long, 0.8- 1.2 mm diam. in lower half, 2 mm diam. distally, lobes 5, 2-4 mm long, ca. 1.5 mm broad at the base, triangular; anthers 2.5 mm long. Fruits 4-5 mm long (not including the 0.4-1 -mm-high calyx) and 4-5 mm diam., subglo- bose or oblong, bright blue, purple or black, somewhat rugose when dried, glabrous or sparsely puberulent; py- renes 3-3.5 mm long, smooth or with 3-5 ridges and slight transverse ribbing.
A common species of lowland rain forest for- mations, from 20 to 600 m elevation on the Ca- ribbean slope and from 300 to 500 m on the Pacific slope (to 1000 m in Honduras). Flowering pri- marily in February-May; fruiting mostly in June- February. The species ranges from Honduras to central Panama.
Psychotria suerrensis is recognized by its short- petiolate leaves that often dry greenish and with veins reddish beneath, small two-lobed stipules, unusual subcapitate inflorescences with many large brightly colored persisting bracts, and longer co- rollas. The freshly cut pith turns red, as it does in Simira. While common in the Caribbean low- lands, this species is rare on the Pacific slope where it has not been collected below 300 m elevation. This species resembles P. hoffmannseggiana (with smaller bracts and inflorescences). Breeding biol- ogy was studied by Bawa and Beach (1983).
Psychotria suerrensis is closely related to P. lu- pulina Benth., a variable and wide-ranging species of South America (cf. Steyermark, 1 974, pp. 1 539- 1 544). If P. lupulina is circumscribed very broadly, it is probable that P. suerrensis will be included as a northern subspecies with consistently larger co- rollas and slightly ridged pyrenes.
Psychotria sylvivaga Standl., J. Wash. Acad. Sci. 18:274. 1928. Figure 62.
Shrubs or subshrubs, 1-4 m tall, leafy stems 1.2-4 mm thick, glabrous or sparsely puberulent (except for a ring of reddish hairs at the node); stipules 6-1 5 mm long, 3-7 mm broad, ovate, acute to obtuse, sometimes with a tubular sheath 10 mm long, glabrous and drying red- dish brown, caducous. Leaves with petioles 6-26(-35) mm long, 0.6-1.8 mm thick, glabrous; leaf blades (6-)9- 17 long, (2.5-)3-5.5 cm broad, elliptic-oblanceolate to narrowly elliptic or elliptic-oblong, apex acute or short- acuminate, base tapering gradually and cuneate to acute, slightly decurrent on petiole, drying membranaceous to chartaceous, dark grayish or reddish brown, glabrous above, glabrous or minutely papillate-puberulent on the
veins beneath, 2° veins 8-12/side. Inflorescences ter- minal or becoming axillary, solitary, 5-16 cm long and 3-10 cm broad, usually an open panicle, peduncle 4- 7(-12) cm long and 1 mm thick, to 10 cm in fruit, gla- brous or sparsely puberulent, bracts 1-4 mm long, su- bulate, pedicels 1-2 mm long. Flowers glabrous on the outside, hypanthium ca. 0.7 mm long, obconic, calyx ca. 0.8 mm long, lobes ca. 0.4 mm long, obtuse; corolla white, tubular-salverform, tube 3-6 mm long, narrow (1.3 mm) or expanded distally to 2 mm diam., lobes 1- 2 mm long, triangular. Fruits 5-9 mm long, not including the persisting (1-3 mm) calyx, 5-6 mm diam., subglo- bose, red.
Plants of montane forest formations, from 1300 to 2700 m elevation. Flowering in January-Sep- tember; fruiting in November-December and March. The species ranges from the Cordillera de Tilaran to the Chiriqui Highlands of Panama.
Psychotria sylvivaga is recognized by its higher- elevation habitat, leaves usually widest above the middle and with ca. 1 0 pairs of secondary veins, larger narrowly ovate unlobed stipules, and longer corolla tube. The leaves drying dark grayish or reddish brown and the red fruit are characteristics of subgenus Psychotria. Recent collections from Panama make it appear that this species is more variable than previously thought. This includes material collected in the Cerro de la Muerte at about 2500 m elevation with more broadly elliptic leaves and somewhat larger flowers. In addition, a broader interpretation might require the inclu- sion of P. stockwellii.
Psychotria tapantiensis C. M. Taylor, sp. nov. Fig- ure 57.
Arbuscula 1-4 m alta, ubique dense pilosula. Folia laminis ellipticis, 8.5-23.5 cm longis, 4-10 cm latis, ve- nis secundariis paribus 12-17 dispositis; petiolo 1-4 cm longo; stipulis vagina 2-3 mm longa atque lobulis del- toideis 1.5-2 mm longis munitis. Inflorescentia ex panic- ula terminali pyramidali 8-12 cm longa, 5.5-12 cm lata, pedunculo 5-8 cm longo insidente constans; bracteis flo- res subtentibus ex ellipticis ovatis, 2-5 mm longis. Flores limbo calycino usque ad 0.5 mm longo, dentato; corolla infundibuliformi, tubo 5-7 mm longo, lobulis 1-1.5 mm longis. Fructus diametro 3-4 mm, atropurpureus.
TYPUS-/?. L. Wilbur 307 4 l(hololypus DUKE), 10 Aug. 1980, Reserva Tapanti, 1 1 km above the Tapanti bridge, Cartago, Costa Rica.
Shrubs or small trees, (l-)2-4 m tall, leafy stems 2-8 mm thick, minutely puberulent to pilosulous with straight or curved hairs 0. 1-0.3 mm long; stipules 2-6 mm long, sheath 2-3 mm long with 2 triangular lobes on each side 1.5-4 mm long, obtuse to rounded, glabrous. Leaves with petioles 1-4 cm long, 1.2-2 mm thick, minutely puber- ulent; leaf blades (8-)10-20(-23.5) cm long, (4-)4.5-
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9(-10) cm broad, elliptic, ovate-elliptic to elliptic-ob- long, apex acuminate with tip 8-1 5 mm long, base obtuse to acute, drying chartaceous, dark yellowish green or brownish green above, minutely (0.05-0. 1 mm) puber- ulent above (barely visible but soft to the touch), softly puberulent beneath with yellowish hairs 0. 1-0.4 mm long, 2° veins 1 1-17/side. Inflorescences terminal, pyramidal open panicles 8-25 cm long, 6-17 cm broad, peduncles 5-10(-15) cm long, 1-3 mm thick, minutely puberulent, bracts 6-8 mm long, 2-3 mm broad, lanceolate, flowers sessile in open distal cymes, bracteoles 2-5 mm long and 2-3 mm broad, often persisting in fruit. Flowers usually glabrous externally, hypanthium ca. 0.5 mm long, calyx lobes ca. 0.3 mm long, obtuse; corolla whitish to yellow, yellow-green or pink, tubular-funnelform, tube 5-7 mm long, 1.3-1.8 mm diam., lobes 1-1.5 mm long, trian- gular. Fruits 3-4 mm long, 3—4 mm diam., globose to oblate, becoming black or dark purplish; pyrenes with 4-5 sharp ridges, concave between them.
Plants of lower montane cloud forest formations on the Caribbean slope, at 1400-1600 m eleva- tion. Flowering in March and June-August; fruit- ing in May-August. This species is known only from between Vara Blanca (Alajuela/Heredia) and the upper drainage area of Rio Grande de Orosi, Cartago.
Psychotria tapantiensis is recognized by the large many-branched inflorescences usually with large persisting bracteoles, larger leaves softly puberu- lent beneath, and small blackish fruit. The species, a member of subgenus Heteropsychotria, is closely related to P. luxurians of lower elevations with narrower bracts and more rounded stipules. It may be easily confused with P. berteriana with stipules with two-angled lobes and smaller bracts. Early collections were mistakenly annotated as P. pali- coureoides. The species is common around the Re- fugio Nacional Tapanti.
Psychotria tenuifolia Sw., Prodr. 43. 1788. P. ses- silifolia Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11: 228. 1844. P. granadensis Benth. in Oerst., Vidensk. Meddel. Dansk. Na- turhist. Foren. Kjobenhavn 1852: 34. 1853. Fig- ure 65.
Shrubs or subshrubs, 0.3-2 m tall, leafy stems 1-5 mm thick, minutely (0.05-0. 1 mm) papillate-puberulent or glabrous; stipules 6-14 mm long, 4-6 mm broad at base, triangular to ovate with 2 closely spaced lobes 1- 6 mm long, the lobes narrowly triangular to linear, drying reddish brown, deciduous. Leaves often closely clustered distally, petioles 2-25(-40) mm long, 0.6-2 mm broad, glabrous; leaf blades (3.5-)6-20(-23) cm long, (1.5-)2- 6.5(-7.7) cm broad, narrowly elliptic to oblanceolate, elliptic or lanceolate, apex usually tapering gradually and acute (obtuse or short-acuminate), base cuneate to acute
and decurrent on petiole, drying membranaceous to thin- chartaceous, grayish or dark grayish brown above, gla- brous above, glabrous or sparsely to densely puberulent beneath with thin whitish hairs 0.05-0.2 mm long, 2° veins 7-13/side, loop-connected distally to form a thin- arcuate submarginal vein. Inflorescences terminal or be- coming axillary, 1-6/node, 1.5-4 cm long, 2-3 cm broad, open paniculate, peduncles 5-20 mm long, 0.5-1.3 mm thick, minutely puberulent, bracts and bracteoles 0.5-1 mm long, subulate, flowers sessile or on short (1 mm) pedicels in dense distal clusters of 3-5. Flowers minutely papillate-puberulent or glabrous externally, hypanthium ca. 0.5 mm long, calyx 0.2-0.5 mm long, lobes ca. 0.3 mm long, obtuse; corolla salverform, white or yellowish green, tube 1.5-2 mm long, 0.8-1 mm diam., lobes 5, 1-1.8 mm long, ca. 1 mm broad at the base, triangular, barbate in throat; stamens 5, anthers 0.5-0.7 mm long. Fruits 4-5 mm long, 3-4 mm diam., usually ellipsoid, red; pyrenes with ca. 5 low rounded ridges.
Plants of both deciduous woodlands and ever- green premontane moist forest formations, from near sea level to 800(-1 100) m elevation in Costa Rica (to 2000 m elsewhere). Flowering in April- August in southern Central America; fruiting throughout the year but primarily in July-Decem- ber. The species ranges from Mexico to eastern Panama and occurs in southern Florida, Cuba, and Hispaniola.
Psychotria tenuifolia is recognized by its short stature, thin (often narrow) leaves with slender arcuate submarginal vein, biacuminate deciduous stipules, usual presence of thin whitish hairs, clus- ters of short umbelliform panicles, small ellipsoid fruit, and preference for deciduous woodlands (in Costa Rica). The tendency of leaves to dry grayish and the bright red fruit are characteristics of sub- genus Psychotria. This is a variable species, in both morphology and ecology. It has rarely been col- lected outside of lowland Guanacaste in Costa Rica; it has not been collected in the eastern half of Costa Rica or the western half of Panama. Compare P. pubescens.
Psychotria trichotoma M. Mart. & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11: 227. 1844.
This species does not appear to occur in Costa Rica. It ranges from southern Mexico to central Nicaragua and is found also in Venezuela and Ec- uador. The records for Panama were misidentifi- cations (Hamilton, 1989). This species belongs to subgenus Psychotria and is part of a group includ- ing P. carthaginensis, P. chagrensis, P. limonensis, P. quinqueradiata, and P. tenuifolia. This species is recognized by the larger (10-26 x 6-15 cm)
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leaves with ca. 152° veins per side, larger (7-16 cm) inflorescences, and larger (6-8 mm) obovoid fruit.
Psychotria turrubarensis Burger & Q. Jimenez, Brenesia (in press).
Shrubs, ca. 4 m tall, leafy stems 1.3-3.5 mm thick, glabrous, drying pale grayish; stipules 12-16 mm long, 3-4 mm broad at the base, lanceolate, forming a calyp- trate sheath over the shoot apex and splitting along 1 side, broadly ovate (ca. 10 x 10 mm) below the young inflorescences, drying dark reddish brown, caducous. Leaves opposite or apparently in a whorl of 4 when an internode fails to develop, petioles 5-18 mm long, 1.1- 2.3 mm broad, glabrous and drying dark; leaf blades 7- 13 cm long, 3-5.5(-6.5) cm broad, elliptic to elliptic- oblong or narrowly elliptic-obovate, apex acute to blunt- ly obtuse, base acute to cuneate and slightly decurrent on petiole, drying stiffly chartaceous, grayish, glabrous above and below, 2° veins 5-9/side, deep rounded pit domatia (0.2-0.4 mm) usually present in vein axils be- neath. Inflorescences terminal or pseudoterminal, soli- tary, sessile capitula, 8-16 mm long, 10-15 mm broad, at first enclosed by the broad bracteate stipules, brac- teoles to 2 mm long, difficult to see among the villous- arachnoid hairs, flowers sessile or subsessile. Flowers with corolla appendages separate in bud, hypanthium ca. 1 mm long, calyx with villous yellowish hairs, lobes 5, ca. 1 mm long; corolla white, salverform, glabrous externally, corolla tube to 5 mm long, 2 mm diam. dis- tally, with dense thin hairs within the throat, lobes ca. 3 mm long and 1 mm broad, with a thickened rounded appendage ca. 0.3 mm diam. Fruits 7-8 mm long, 6-9 mm thick, subglobose, red at maturity and subsessile, surface smooth; pyrene 7 x 6.5 mm, flat on the inner (adaxial) face, with 5 longitudinal sulci abaxially.
Plants of evergreen forest on the Pacific slope in central Costa Rica at about 1600 m elevation. Flowering and fruiting on 6 November 1989 (Q. Jimenez et al. 935 CR holotype, F). This species is known only from the Faldas del Cerro Bares in the Zona Protectora de Turrubares, San Jose Prov- ince.
Psychotria turrubarensis is distinguished by its small sessile capitate inflorescences, united stip- ules forming a calyptrate Ficus-\\ke sheath over the vegetative shoot apices, lack of puberulence on vegetative parts, well-developed pit domatia, and appendage at the tips of the corolla lobes. The red fruit and leaves drying grayish are character- istics of subgenus Psychotria. The Cerros de Tur- rubares are a conspicuous feature at the south- western perimeter of the Meseta Central, but they are isolated and have been little explored. It is surprising that so small an area should harbor such a distinctive new species.
Psychotria uliginosa Sw., Prodr. 43: 1788. Figure 12.
Herbaceous subshrubs, 0.5-1 .5 mm tall, stems usually unbranched and succulent, leafy stems 2-12 mm thick, terete and often thickened at the nodes, glabrous; stipules 3-6 mm long in early stages, shorter and broader (to 10 mm) in age, with a short (2-A mm) sheath, broadly ob- tuse or truncate distally, glabrous and persisting. Leaves well spaced along the stem, petioles 2-6 cm long, 1.5-3 mm thick, glabrous; leaf blades (ll-)15-35 cm long, (3-)5-13 cm broad, narrowly elliptic-oblong to elliptic- obovate or elliptic-oblanceolate, apex gradually or abruptly narrowed and short-acuminate, tip 3-10 mm long, base cuneate or acute and slightly decurrent on petiole, slightly succulent in life but drying chartaceous, dark green above and pale green beneath in life but be- coming dark brown above when dried and much paler beneath, glabrous above and below, 2° veins 9-15/side. Inflorescences axillary, usually I/node, 2-5 cm long at anthesis, to 10 cm long in fruit, 2-8 cm broad, open pyramidal paniculate with usually 3 main branches and flowers in dense distal congested clusters, peduncles 1- 6 cm long, ca. 1.5 mm thick, glabrous, bracts 2-4 mm long, narrowly triangular, flowers sessile. Flowers gla- brous or rarely sparsely and minutely puberulent exter- nally, hypanthium and calyx 1.5-2.5 mm long, obconic or cylindrical, calyx tube truncate or with minute (0.2 mm) lobes; corolla white or pinkish, tube 1-2.5 mm long and ca. 0.7 mm diam., lobes 0.5-1.5 mm long. Fruits 7-10 mm long, 6-8 mm diam., oblong or ovoid, bright red or orange-red; pyrenes ca. 8 mm long, with 1 prom- inent ridge and 2 thickened margins.
Plants of shaded understory in wet evergreen rain forest formations on both the Caribbean and Pacific slopes, from near sea level to 900(-1 100) m elevation. Probably flowering throughout the year; fruiting in all months. This species ranges from Mexico and the West Indies to Ecuador and the Guianas.
Psychotria uliginosa is recognized by its short usually unbranched semisucculent stems, short truncated stipules (in age), large glabrous leaves dark above and pale beneath, axillary inflores- cences with congested little flowers, and bright red fruit. This distinctive species is uncommon on the Pacific slope. The general aspect of these plants is similar to P. macrophylla and P. aggregata, but those species have white fruits and the leaves dry- ing greenish without so great a color contrast be- tween the upper and lower surfaces.
Psychotria umbelliforims Dwyer & Hayden, Ann. Missouri Bot. Card. 55: 44. 1968. Figure 47.
Shrubs or small treelets to 2 m tall, leafy stems 2-6 mm thick, densely pubescent with stiff yellowish hairs ca. 0.5 mm long; stipules united around stem 1-2 mm,
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with 2 slender distal awns 4-1 1 mm long, puberulent, deciduous. Leaves with slender petioles (4-) 7-2 5 mm long, 1-2.2 mm thick, densely pubescent; leaf blades (6-)8-18(-26) cm long, (3-)5-8(-9) cm broad, ovate-el- liptic to elliptic-oblong, apex tapering gradually and acu- minate or acute with tip to 2 cm long, base acute to obtuse, drying chartaceous, dark green above, glabrous above and with small (0.1-0.2 mm) whitish cystoliths visible, pubescent beneath with thin yellowish hairs 0.3- 0.6 mm long especially on major veins, 2° veins 7-97 side, distal veins arcuate-ascending. Inflorescences ter- minal and solitary, 5-8 cm long, 3-6 cm broad, umbel- liform, peduncles 3-5 cm long, 1.5-2 mm thick, densely pubescent, with 3 primary branches 1-2 cm long bearing terminal capitulae 6-10 mm broad, bracts ca. 7 mm long and 3 mm broad at the base, flowers sessile in glomerules of 3-8, subtended by bracteoles ca. 4 mm long, ovate- triangular. Flowers puberulent externally, calyx ca. 2.5 mm long, lobes to 0.8 mm long, obtuse or acute; corolla salverform, yellowish white and greenish, tube ca. 12 mm long, lobes ca. 8 mm long; stamens 5, anthers ca. 2 mm long. Fruits 15-18 mm long, oblong, glabrous and smooth, black; pyrenes smooth.
Plants of evergreen lower montane forest for- mations and lowland rain forest formations, from near sea level to 1 500 m elevation. Flowering in July-August. Rarely collected plants of the Carib- bean slopes and lowlands; known only from Costa Rica and Panama.
Psychotria umbelliformis is recognized by the pilosulous hairs on nearly all surfaces (except the upper leaf surfaces), longer petioles, umbelliform inflorescences with three distal capitula or glom- erules of sessile flowers, and longer corollas. This species is poorly known. We have seen only the following collections from Costa Rica: Croat 36753 CR, MO, us, and Standley 36775 us.
18 mm long, ca. 1 mm thick, glabrous, bracts 2-3 mm long, linear, flowers subsessile in distal cymes, pedicels 0-1 mm long. Flowers glabrous externally, hypanthium ca. 0.8 mm long, cylindric or obconic, calyx tube ca. 0.3 mm long, truncate; corolla tubular, white, tube 1.8-3.5 mm long and ca. 1 mm diam., lobes 4, 0.4-1 .5 mm long; stamens 4, anthers 0.8-1.1 mm long. Fruits 4-5 mm long and 5-6 mm diam., smooth and bilobed or globose, yellow or orange and becoming black and lustrous.
Understory plants in lower montane cloud for- est formations on the Caribbean slope and con- tinental divide, from 800 to 1600(-1900) m ele- vation. It has been collected at about 450 m elevation on Volcan Orosi, a pattern of unusually low elevation seen in a number of other species. Flowering in May-July; fruiting in October-March. In Costa Rica nearly all our collections coming from the Cordillera de Tilaran and from north of San Ramon, Alajuela. The species ranges eastward into Panama.
Psychotria valeriana is recognized by the very unusual stipular morphology, lack of pubescence, small inflorescences with little flowers, restricted flowering period, and fruits usually without prom- inent longitudinal ribs. The single stipular awn is occasionally deeply bifid but always arises at one point in the center of the distal stipule margin. The tooth-like appendage on the inner (adaxial) surface of the awn is unique among our species. Unlike most of our other species of subgenus Het- eropsychotria, the fruit is orange during later stages. This species is very similar to P. acuminata of lower elevations, but the stipular morphology helps to separate the two.
Psychotria valeriana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1363. 1938. Figure 56.
Shrubs, (l-)1.5-3(-4) m tall, leafy stems 0.7-3 mm thick, glabrous and smooth, intern odes usually becoming elongated (4-1 5 cm); stipules with short-truncate collar- like sheath 0.3-2 mm long and 1 central distal awn 2-4 mm long that may be split (bifid) near the tip (rarely bifid to the base), often with 1 (rarely 2) thickened "teeth" ca. 1 mm long borne on the adaxial side from the base of the awn, glabrous, persisting. Leaves with petioles 4- 9(-12) mm long, 0.5-1.3 mm thick, glabrous; leaf blades 7-1 3(-l 5) cm long, 2.3-4(-5.5) cm broad, narrowly ovate- elliptic to elliptic, lanceolate-oblong or narrowly elliptic- oblong, apex acuminate or narrowly acute with tip 5-12 mm long, base acute (obtuse) and slightly decurrent on petiole, drying thin-chartaceous and olive green to green- ish brown above, glabrous above and below, 2° veins 5- 8/side. Inflorescences terminal and solitary (3), 1.5-5 cm long, 1.5-3 cm broad, small open pyramidal panicles with usually 2 pairs of opposite branches, peduncles 4-
Psychotria viridis Ruiz & Pav., Fl. Peruv. 2: 61, pi. 210. 1799. P. glomerata H.B.K., Nov. gen. sp. 3: 362 (quarto). 1 8 1 9. P. microdesmia Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 36. 1853. Figure 65.
Shrubs, 1.5-5 m tall, leafy stems 1.5-4 mm thick, glabrous; stipules 7-14 mm long, 3-5 mm broad, ob- ovate, acuminate and fimbriate along the distal edge, glabrous, membranaceous and drying reddish brown. Leaves often closely clustered distally, petioles 3-9 mm long, 0.5-1.3 mm thick, glabrous; leaf blades 5-15 cm long, 2-5.5 cm broad, elliptic-obovate to oblanceolate or obovate, apex acute or short-acuminate, base tapering gradually and acute or cuneate, decurrent on the petiole, drying thin-chartaceous, pinkish gray or light to dark gray above, glabrous above and below, 2° veins 5-7/side, large (1-2 mm) pit domatia often present in the axils of distal veins beneath. Inflorescences terminal and soli- tary, 4-10 cm long, spiciform with verticillate groups of flowers or with l(-2) pair(s) of opposite lateral branches
278
FIELDIANA: BOTANY
above the peduncle, peduncles 10-30 mm long, ca. 1 mm thick, glabrous, flowers sessile in verticils of 4-8 separated by 5-15 mm. Flowers glabrous externally, hy- panthium ca. 0.5 mm long, calyx ca. 0.3 mm long, sub- truncate; corolla urceoate, white, tube ca. 1.5 mm long and 1 mm diam., lobes 5, ca. 1 mm long. Fruits 4-6 mm long, 4-6 mm diam., globose or ellipsoid, turning red; pyrenes with 4-5 rounded ridges.
Rarely collected plants (in Costa Rica) of ev- ergreen Caribbean rain forest formations and in the General Valley, from near sea level to 700(-900) m elevation in Costa Rica. Flowering in March, and October-January; probably fruiting through- out the year. The species ranges from Cuba, Belize, and Guatemala to Bolivia.
Psychotria viridis is distinguished by the thin narrowly obovate leaves with large domatia, broad thin stipules often fimbriate and caudate-acumi- nate at the apex, and the unusual inflorescences with few or no lateral branches and the flowers in well-separated verticils. No other Costa Rican Psychotria has such an inflorescence. The red fruit and leaves drying grayish are characteristics of subgenus Psychotria. The few Costa Rican collec- tions come from the Caribbean lowlands in central and southern Costa Rica (not known from La Sel- va) and the slopes of the Talamanca mountains. It is more common in South America.
Psychotria sp. A, Cocos Island, subgenus Hetero- psychotria, not matched.
Shrubs or small trees, leafy stems 0.9-2 mm thick, glabrous, with minute longitudinal striations; stipules 1- 2 mm long, with a small U-shaped sinus 0.7-1.2 mm deep separating the 2 acute lobes, glabrous, persisting or breaking off. Leaves with petioles 3-15 m long, 0.4-1 mm thick (dried), glabrous; leaf blades 5.5-10 cm long, 1.7-3.7 cm broad, narrowly ovate-elliptic to elliptic-lan- ceolate or elliptic, apex gradually narrowed and acu- minate with tip 5-10 mm long, base obtuse to acute and slightly decurrent on petiole, drying membranaceous to thin-chartaceous, greenish brown above, glabrous above or below, 2° veins 5-8/side. Inflorescences solitary and terminal, 5-9 cm long, open paniculate with 2 lateral branches at first node, peduncles 3-5 cm long, 0.5-1 mm thick, glabrous, bracts to 9 mm long and linear, flowers in distal thin-branched dichasia of 3-5 or short helicoid cymes, pedicels 0-4 mm long. Flowers not seen at an- thesis, hypanthium ca. 1 mm long and 0.7 mm diam., calyx cupulate, lobes 0.5-1 mm long, triangular and acute. Fruits ca. 4 mm long (not including the 1 mm long persisting calyx), ca. 3.5 mm diam., ovoid-ellipsoid, black; pyrenes 4-5 -angled.
This species is presently known from only a sin- gle collection from Cocos Island: Holdridge 5168
(CR, F, us) collected in late February 1963. The species is characterized by its thin smaller elliptic- lanceolate leaves, short stipules with small distal lobes, few-flowered inflorescences with slender pe- duncles, and very slender elongate branches and pedicels. The well-developed calyx lobes and lack of puberulence are also noteworthy. It appears to be a species of Colombia.
Psychotria sp. B, subgenus Psychotria aff. P. pa- pantlense (Oerst.) Hemsl.
Small trees, 1-4 m tall, leafy stems 0.9-3.5 mm thick, glabrous and pale grayish, smooth; stipules sheathing, 4-9 mm long, acute (or biaristate), reddish when dried, caducous. Leaves with petioles 5-35 mm long, 0.5-1.2 mm thick, glabrous; leaf blades 6-11 cm long, 1.8-4 cm broad, narrowly elliptic-oblong to narrowly oblong, acu- minate at the apex, tapering gradually to the cuneate attenuate base and decurrent on the petiole, drying thin- chartaceous and grayish green, glabrous above and be- low, 2° veins 7-9/side. Inflorescences not seen at anthe- sis, 5-9 cm long, apparently with only 2 lateral branches at the first node, with few branches and few (1-4) fruits, peduncles 2-3 cm long, 1 .6 mm thick, glabrous. Flowers not seen at anthesis. Fruits 6-10 mm long, 4-6 mm diam., ellipsoid, purple at maturity.
The thin long-decurrent gray-green leaves and open few-fruited infructescences are reminiscent of P. papantlense, but that species usually has in- florescences with four branches at the first node and is not known south of Belize and southern Mexico. This description is based on Haber et al. 4440 (CR, MO) from 1 000 m elevation near Tilaran; fruiting in April. This may be an atypical southern extension of P. papantlense or perhaps a new spe- cies.
Randia Linnaeus
Shrubs, trees, or lianas, bisexual or unisexual (dioe- cious), glabrous or puberulent, nodes often with 1^4 ax- illary or extraaxillary spines; stipules interpetiolar, small and triangular to tubular (intrapetiolar) at the base, often imbricate on the short-shoots. Leaves opposite or fas- ciculate on short-shoots, sessile or petiolate, membra- naceous to coriaceous, entire, rarely with domatia of tufted hairs in the vein axils. Inflorescences terminal or axillary (rarely cauliflorous), of solitary flowers or with few fasciculate or rarely cymose flowers, sessile or ped- icellate, often subtended by bracteate stipules. Flowers unisexual and dioecious or monoecious, less often bi- sexual, 4-7-parted, hypanthium cupulate to tubular, truncated distally and with slender or expanded (folia- ceous) calyx lobes; corolla salverform or funnelform, white or pale yellow, glabrous or villous within, tube short to
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
279
pearance of bilocularity, (2) many discoid seeds imbedded in sweet pulp becoming brown or black, (3) a seed coat with thick-walled testa cells having a smooth or papillose inner wall, and (4) pollen in permanent tetrads (rarely diads or monads).
Species ofRandia are characterized by the usu- ally terminal solitary flowers or few-flowered in- florescences, salverform white or yellowish corol- las with convolute lobes in bud, and hard-surfaced baccate fruit with many horizontal discoid seeds. A number of species have conspicuous short-shoots with tightly clustered distal leaves, and many spe- cies have conspicuous sharp-tipped spines. The great variability of leaf size and form, dioecy, and spiny branches that discourage collectors make this a taxonomically difficult genus.
long, lobes (4-)5-6(-7), convolute in bud; stamens usu- ally 5, inserted in the distal part of the tube, filaments short or absent; ovary basically 1-locular, ovules many in 3-4 tiers on the 2 fleshy parietal placentas. Fruits baccate, 1 -locular or becoming 2-locular, globose to el- lipsoid with a thick coriaceous pericarp; seeds discoid, arranged horizontally in a gelatinous pulp.
Randia is now considered to be a Neotropical genus of 60-70 species, ranging from southern Florida and Texas through Central America and the West Indies to Bolivia. Our treatment follows the annotations and advice of David Lorence (PTBG). He distinguishes Randia from other genera of the Gardineae by the following combination of characters: (1) a unilocular ovary with parietal pla- centas that may abut in fruit and give the ap-
Key to the Species of Randia
la. Plants lianas or vines 2
1 b. Plants trees or shrubs 4
2a. Plants glabrous, spines usually solitary and recurved R. altiscandens
2b. Plants puberulent to pilose, spines opposite or in 4's at each node 3
3a. Spines usually 4/node; <5 corolla tubes ca. 8 cm long, 9 corolla tubes ca. 5 cm long
R. vazquezii
3b. Spines usually opposite; $ corolla tubes 4-5 cm long, 9 corolla tubes 3-3.5 cm long
R. retroflexa
4a. Fruits cauliflorous and more than 4 cm diam.; flowers cauliflorous or axillary, more than 5 cm long; leaf blades often exceeding 25 cm in length and with 10-16 pairs of 2° veins; plants of wet
evergreen forests [10-700 m elevation] 5
4b. Fruits usually borne in among the leaves, less than 3 cm diam. if cauliflorous; flowers axillary or terminal, 2-10 cm long; leaf blades rarely exceeding 25 cm in length or with more than 10 pairs
of 2° veins (except R. genipoides); plants of wet evergreen to dry deciduous forests 6
5a. Corolla tube 5-8 cm long, flowers cauliflorous; leaf blades cuneate at the base . . . R. mira 5b. Corolla tube ca. 4.5 cm long, flowers often axillary; leaf blades slightly rounded or subtruncate
at the base R. pterocarpa
6a. Leaves of the same node differing greatly in size and the larger leaves more than 10 cm long
[hypanthium and calyx sometimes with minute red glands] R. gentryi
6b. Leaf blades about the same size at each node or the larger blades less than 5 cm long if differing
greatly in size 7
7a. Mature flowers present 8
7b. Mature fruit present 19
8a. Calyx lobes laminar, nearly as broad as long, obtuse or rounded distally and with a narrowed
petiole-like base [evergreen forests 800-2300 m elevation] R. calycosa
8b. Calyx lobes not laminar, narrowly triangular or awned, lacking a narrowed petiole-like base
9
9a. Corolla tube less than 2 cm long 10
9b. Corolla tube more than 2 cm long 14
lOa. Largest leaf blades less than 6 cm long, often appearing to be very unequal at the same
node or short-shoot, with 3-4 pairs of 2° veins 11
lOb. Larger leaf blades more than 6 cm long, usually equal or subequal at a node, with 2-
1 1 pairs of 2° veins 12
1 la. Calyx lobes minute or absent; leaf blades to long, often obovate; wet forests 20- 500 m elevation . . R. loniceroides
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FIELDIANA: BOTANY
1 Ib. Calyx lobes 1-2 mm long; leaf blades to 6 cm long, usually elliptic; dry deciduous
forests 0-300 m elevation R. thurberi
12a. Calyx lobes 0-1 mm long; corolla tube 3-7 mm long; evergreen and deciduous for- mations [0-900 m elevation] R. aculeata
12b. Calyx lobes 1-9 mm long, corolla tube 8-18 mm long; wet evergreen formations 13
13a. Calyx lobes 1-4 mm long, corolla tube 10-18 mm long; 0-1 100 m elevation
R. grandifolia
13b. Calyx lobes 5-9 mm long, corolla tube 8-14 mm long; 1000-2300 m elevation
R. calycosa
1 4a. Largest leaf blades less than 6 cm long, often very different in size at the same node [corolla
tube 25-45 mm long, wet evergreen forests] R. brenesii
14b. Largest leaf blades more than 10 cm long, usually similar in size at each node 15
1 5a. Corolla tube 20-50 mm long 16
15b. Corolla tube 50-90 mm long 18
16a. Calyx lobes 1-3 mm long; cloud forests, 1300-1600 m elevation R. matudae
16b. Calyx lobes 3-6 mm long; lowland evergreen forests, 0-800 m elevation 17
1 7a. Petioles to 1 0 mm long, leaf blades rarely more than 6 cm broad; widespread
R. armata
1 7b. Petioles ca. 20 mm long, leaf blades usually more than 1 0 cm broad; rarely collected
plants of the Caribbean lowlands R. grayumii
1 8a. Calyx lobes ca. 4 mm long; leaf blades to 1 6 cm long, elliptic to ovate R. pittieri
18b. Calyx lobes 1 1-25 mm long; leaf blades to 35 cm long, oblanceolate R. genipoides
1 9a. (from 7b) Fruits 10-15 mm long, globose 20
19b. Fruits 1 5-100 mm long, globose to ovoid or ellipsoid 21
20a. Leaf blades 6-30 mm long, rounded to bluntly obtuse at the apex, 2° veins 3— 4/side; spines
usually absent; wet forests, 20-500 m elevation R. loniceroides
20b. Leaf blades 8-90 mm long, acute to bluntly obtuse at the apex, 2° veins 3-8/side; spines
usually present; dry or wet forests, 0-1700 m elevation R. aculeata
21a. Fruits 15^0 mm long 22
21b. Fruits 40-100 mm long 28
22a. Leaf blades not exceeding 6 cm in length 23
22b. Larger blades regularly exceeding 6 cm in length 25
23a. Leaf blades to 4 cm long, often differing greatly in size at the same node [wet evergreen
forests, 100-1600 m elevation] R. brenesii
23b. Leaf blades to 6 cm long, usually equal or subequal at the same node 24
24a. Persisting calyx 0-2 mm high; dry deciduous forests, 0-300 mm elevation
R. thurberi
24b. Persisting calyx 6-15 mm high; evergreen montane forests, 1000-2300 m elevation . .
R. calycosa
25a. Leaf blades often oblanceolate to obovate; stipules with a short awn 1-2 mm long; evergreen
forests, 1000-2300 m elevation R. calycosa
25b. Leaf blades never oblanceolate, sometimes obovate; stipules acute or with a narrow tip;
evergreen or partly deciduous forests, 0-1 100 m elevation 26
26a. Spines absent; leaf blades 1 7-28 cm long, minutely (0.05 mm) puberulent or glabrous beneath; stipules somewhat striate; sepal lobes (on fruits) to 5 mm long and narrow; Caribbean lowlands
R. grayumii
26b. Spines usually present, and without the above combination of characters; widespread species
27
27a. Spines often 4/node; leaf blades 3-18 cm long, usually puberulent and often with tufted
domatia in the vein axils; deciduous to evergreen formations R. armata
27b. Spines 2/node; leaf blades 7-30 cm long, glabrous and without tufted domatia in vein axils;
evergreen formations R. grandifolia
28a. (from 21b) Stipules 15-25 mm long; leaves closely clustered at the ends of branchlets, blades 14-
35 cm long, oblanceolate to narrowly elliptic [evergreen forests, 50-500 m elevation]
R. genipoides
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 28 1
28b. Stipules rarely exceeding 5 mm in length; leaves not more than 16 cm long if closely clustered at
the ends of stems, blades to 22 cm long and never oblanceolate 29
29a. Plants of cloud forests, 1300-1600 m elevation; spines absent, leaves and stems glabrous, leaf
blades stiffly chartaceous to subcoriaceous R. matudae
29b. Plants of lowland evergreen or partly deciduous forest formations, 0-1000 m elevation; spines
often 4/node, leaves and stems puberulent, blades membranaceous to thin-chartaceous 30
30a. Leaf blades with 3-7 pairs of secondary veins, 4-1 1 cm long; deciduous forests . . .R. monantha 30b. Leaf blades with 9-12 pairs of secondary veins, 7-16 cm long; evergreen and partly deciduous
forests R. pittieri
Randia aculeata L., Sp. PI. 1192. 1753. R. mitis L., Sp. PI. 1 192. 1753. R. karstenii Polak., Lin- naea 41: 568. 1877, fide Lorence in herb. Figure
22.
Shrubs or small trees, 1 .5-3(-5) m tall, dioecious, bark smooth and gray, leafy stems 1.2-4 mm thick, glabrous or rarely puberulent, spines 6-20 mm long, 1.5-3 mm diam. near the base, opposite and lateral, erect or as- cending, sometimes terminating the stem and Y-shaped; stipules 1-3 mm long, 1-3 mm broad, broadly triangular, thin, brownish, sparsely puberulent, without clear ve- nation. Leaves usually closely congested on short-shoots, opposing leaves subequal, smallest leaves sessile, peti- oles 0.5-12 mm long, 0.5-1 mm broad; leaf blades (0.8-) 1 .2-6(-9) cm long, (0.4-)0.8-3(-4) cm broad, ovate to elliptic-ovate, obovate or obovate-orbicular, apex bluntly obtuse to rounded, base tapering gradually and cuneate, decurrent on petiole, drying stiffly chartaceous, dark or pale grayish, glabrous (rarely puberulent), 2° veins 3-8/side. Inflorescences terminal on the short-shoots along the stem or apparently axillary, flowers usually solitary, pedicels absent or undifferentiated from the ovary base. Flowers functionally unisexual, usually glabrous externally, hypanthium 0.5-1 mm long, calyx tube be- coming 1-1.5 mm high, ca. 2 mm broad, calyx lobes 4- 5, ca. 0.5 mm long; corolla salverform, white, glabrous externally, tube 3-8 mm long, 1.5-3 mm diam., lobes 4-5(-6), 3-6 mm long, 2 mm broad at the base, acute. Fruits terminal on small short-shoots and appearing to be axillary, (8-)10-14 mm long, (7-)8-14 mm diam., subglobose to ovoid, greenish becoming white, persisting calyx tube 1-2 mm high and 1.5-2 mm broad, pericarp ca. 1 mm thick.
Plants of partly deciduous and evergreen forest formations on both the Caribbean and Pacific slopes, from near sea level to 1700 m elevation. Flowering in May; fruiting in January, June, and November. This species ranges from Mexico, Florida, and the West Indies to Colombia and Venezuela.
Randia aculeata is recognized by the small dis- tally rounded leaves closely clustered on short- shoots, presence of paired spines, smaller flowers, and white fruit. Our collections are essentially gla- brous but R. aculeata variety dasyclada Steyerm. often has hirtellous stems; it is found along the Caribbean coast of Panama. Specimens formerly
assigned to Randia karstenii have thinner leaves that dry dark and have been found at higher ( 1 000- 1700 m) elevations or in evergreen forests. We follow Lorence in including this material under R. aculeata. More typical material of R. aculeata is found in the drier parts of the Meseta Central where it is called espino bianco. Considerable morpho- logical variation occurs throughout the range of this species, making it a difficult species complex (D. Lorence, pers. comm.).
Randia altiscandens (Ducke) C. M. Taylor, comb, nov. Basanacantha altiscandens Ducke, Arq. Inst. Biol. Veg. Rio Janeiro 2: 172. 1935. R. pepoformis Dwyer, Ann. Missouri Bot. Gard. 67:451. 1980. Figure 23.
Lianas to over 30 m high, dioecious, leafy stems 2-6 mm thick, glabrous, terete, yellowish becoming grayish with age, with usually solitary recurved or retrorse spines 10-20 mm long, 3 mm thick at the base; stipules 3-5 mm long, 2-3 mm broad at the base, triangular to oblong with obtuse apex, free or united to form a short (0.5 mm) tube. Leaves clustered near the ends of branchlets, pet- ioles 5-12(-20) mm long, 0.9-1.9 m broad, glabrous; leaf blades 10-18(-24) cm long, 4-9(-12.5) cm broad, elliptic to elliptic-obovate or obovate, apex obtuse to acute, base obtuse to cuneate and slightly decurrent on petiole, drying chartaceous to subcoriaceous, grayish above, glabrous above and below, 2° veins 5-8/side and loop-connected at the margin in the distal half of the blade, 3° veins subparallel and raised on both surfaces. Inflorescences terminal, of 1-3 $ flowers or solitary 1 flowers, peduncles of the flower groups 0-10 mm long, pedicels 1 8-22 mm long, ca. 2 mm thick, glabrous, un- differentiated from the hypanthium. Flowers unisexual, glabrous externally, hypanthium 2-3 mm diam., contin- uous with the pedicel, calyx tube ca. 1 mm long, calyx lobes 5-6, 5-30 mm long, 1-6 mm broad, narrowly el- liptic-oblong to lanceolate-oblong, with a primary vein and parallel secondaries, acute at the apex, narrowed at the base, stiffly chartaceous; corolla salverform, white, yellow and pilose within the tube, tube 30-50 mm long, 3-5 mm diam., to 10 mm at the throat, lobes 5(-6), 18- 26 mm long, 14-18 mm broad, rounded distally; anthers 6.5-8 mm long. Fruits 5-9 cm long, 3-5.5 cm diam., ovoid to cylindrical, glabrous and lenticellate, drying black, borne on pedicels to 3 cm long, persisting calyx
282
FIELDIANA: BOTANY
ca. 6 mm long and 8 mm diam., pericarp 3-5 mm thick; seeds ca. 1 1 x 8 mm, oblong-ellipsoid.
Plants of rain forest formations on both the Ca- ribbean and southern Pacific slopes in Costa Rica, 0-400 m elevation, to 1 300 in Panama. Flowering in January, March, and May-October; fruiting in February and August. This species is known from the La Selva research site, the Hitoy Cerere reserve in the Caribbean lowlands, and Corcovado Na- tional Park (Osa Peninsula); it ranges southward into the Amazon basin.
Randia altiscandens is distinguished by its climbing habit, usually solitary retrorse spines, prominent subparallel 3° venation, large mostly solitary unisexual flowers, and large oblong fruit. Both corollas and fruit dry black, but the leaves and calyx lobes dry pale grayish. The calyx lobes vary greatly in length in different plants.
Randia armata (Sw.) DC., Prodr. 4: 387. 1830. Gardenia armata Sw., Prodr. 51, 1788. Mus- saenda spinosa Jacq., Stirp. Am. 70. 1 763. Ran- dia spinosa (Jacq.) Karst., Fl. Colomb. 2: 128. 1869, not R. spinosa (Thunb.) Poir., 1812. Fig- ure 22.
Shrubs or small trees, 3-6(-10) m tall, dioecious, leafy stems 1-4 mm thick, glabrous or puberulent with minute (0.2-0.3 mm) thin whitish hairs, glabrescent, spines pres- ent or absent, usually 4/node, 6-18(-25) mm long, 1-2 mm thick at the base, ascending; stipules 4-9 mm long, 2-4 mm broad at the base, brownish, with many parallel veins, glabrous (longer hairs within at the base), often persisting. Leaves usually closely crowded at ends of stems or short-shoots, petioles 3-6(-l 1) mm long; leaf blades (2.5-)4-l 5(-l 9) cm long, ( 1 .5-)2-6(-10) cm wide, broad- ly elliptic, elliptic-oblong, ovate, elliptic-obovate, or ob- ovate, apex abruptly acute to gradually acuminate, base obtuse or acute and slightly decurrent on petiole, drying membranaceous to chartaceous and pale to dark greenish or brown above (much paler beneath), essentially gla- brous above, with thin whitish hairs ca. 0.2 mm long on the veins beneath, 2° veins 5-8/side, tufted domatia pres- ent in some vein axils beneath. Inflorescences of terminal solitary 2 flowers or 2-4(-8) $ flowers, subtended by per- sisting stipules, sessile or on pedicels to 9 mm long and 0.5 mm thick, sparsely to densely puberulent with whit- ish hairs. Flowers unisexual, with gardenia-like odor, hypanthium 1-2 mm long, obconic, to 10 mm long and ellipsoid in 9 flowers, calyx tube ca. 0.5 mm long, calyx lobes 4-5, 3-6(-8) mm long, l-2.5(-3.5) mm broad, lan- ceolate to obovate, striate, yellowish, minutely serrate, glabrous or rarely puberulent; corolla tubular-salver- form, glabrous or minutely puberulent distally on the tube, tube 22-28 mm long, 0.8-2 mm diam. (5 mm at the throat), lobes 5, 9-12 mm long, ca. 6 mm broad, ovate-lanceolate to rhombic; anthers 3.5-5 mm long. Fruits 18-35 mm long, 20-28 mm diam., oblong-ellip-
soid to ovoid, glabrous or sparsely and minutely puber- ulent, surface smooth and becoming yellow or orange, persisting calyx 1-2 mm high and ca. 6 mm broad, peri- carp ca. 1 mm thick; seeds 5-10 mm broad and 2 mm thick.
Plants of partly deciduous and evergreen forest formations and along stream sides in deciduous forests, from near sea level to 800 m elevation. Flowering in February-May; fruiting in all months. The species ranges from Mexico and the West In- dies to Colombia and Brazil.
Randia armata is recognized by the nodes often with four spines, the stiff striate persisting stipules, closely clustered leaves, tufted domatia in some vein axils, and varied lower elevation habitats. Leaf size and shape can vary greatly in different sites. On Barro Colorado Island, Panama, the leaves are deciduous before flowering (Croat, 1978). Common names reported for Costa Rica are cru- cillo, crucito,Jlordecruz,jagua macho, limoncillo, miel quema, mostrenco, and palo de cruz.
Randia brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1365. 1938. Figure 22.
Shrubs or small trees, 1-5 m tall, branches mostly held horizontally, dioecious, leafy stems 1 .2-4 mm thick, at first with small (0.2-0.4 mm) ascending stiffbrownish hairs, glabrescent, terete, spines 6-18 mm long, 0.5-2 mm thick, usually solitary; stipules 1-2 mm long, thin, brownish, glabrous, deciduous. Leaves opposite and of- ten appearing unequal (anisophyllous), congested on short-shoots or separate along the stem, petioles l-3(-5) mm long; leaf blades 4-40 mm long, (2-)4-l 5 mm broad, elliptic or narrowly elliptic-rhombic to ovate-lanceolate (broadly ovate in the very small leaves), apex acute, base acute to cuneate and decurrent on petiole, drying char- taceous, dark grayish green, glabrous above, with few small (0.2-0.4 mm) straight ascending hairs on the mid- vein beneath, 2° veins 3-4/side, strongly ascending. In- florescences of solitary terminal flowers on axillary short- shoots, sessile and subtended by a few short (1 mm) bracteate stipules. Flowers unisexual, ca. 4 cm long, hy- panthium 1.5-2.5 mm long, calyx tube 3-4 mm long, 0.8-2 mm diam., calyx lobes 5, 3-5 mm long, 1.5-5 mm broad, ovate and acute, with a narrow petiole-like base, ciliate; corolla salverform, white, glabrous externally, tube 25-45 mm long, 1-3 mm diam., lobes 5, ll-15(-22) mm long, 4-6(-15) wide, elliptic-oblong to ovate-ob- long. Fruits 15-28 mm long, 14-25 mm diam., globose, glabrous, persisting calyx 2-5 mm high, 2-4 mm broad, pericarp 1-2 mm thick.
Plants of evergreen forest formations on the Ca- ribbean slope, along the continental divide, and in the evergreen Pacific lowlands from 20 to 1 600 m elevation. Flowering in February-June and Oc-
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tober; probably fruiting throughout the year. The species ranges from Nicaragua to Panama.
Randia brenesii is recognized by the small op- posite leaves that are very different in size, solitary flowers, foliaceous calyx lobes, globose fruit, and wet forest understory habitat. The leaves may be subequal but with two larger and two smaller leaves decussate and arising from the apex of the same short-shoot, giving the impression of anisophylly. The unusual calyx lobes distinguish this species from R. loniceroides. This species also resembles Psychotria parvifolia, but that species has subequal opposing leaves. Common names are cachitos and horquetilla de montana.
Randia calycosa Standl., Contr. U.S. Natl. Herb. 20: 201. 1919. Figure 22.
Shrubs or trees, 3-9 m tall, trunk to 25 cm diam., leafy stems 1.2-5 mm thick, with ascending brownish hairs 0.3-0.5 mm long, spines solitary or opposite 4-9(-15) mm long, 1.5-2 mm thick at the base; stipules 4-6 mm long, stipular tube ca. 2 mm long, apex obtuse and with a linear awn ca. 2 mm long, deciduous. Leaves opposite along the stems or sometimes clustered on short-shoots, petioles 3-8 mm long, 0.7-1.4 mm thick, glabrous or sparsely puberulent; leaf blades (2-)4- 1 3 cm long, ( 1 . 3-)2- 5 cm broad, elliptic-obovate to oblong-obovate or ovate, apex acute to acuminate with tip to 7 mm long, base tapering gradually and acute or cuneate, slightly decur- rent on petiole, drying thin- to stiffly chartaceous and usually dark grayish above, glabrous above and below or with short (0.5 mm) thin whitish hairs on the midvein beneath, 2° veins 5-9/side. Inflorescences of solitary ter- minal flowers on short-shoots, subtended by cupulate bracteate stipules 2-3 mm long. Flowers with hypan- thium 6 mm long and 4 mm diam., cylindrical, densely sericeous with brownish ascending hairs, calyx tube 4 mm deep, 3 mm diam., little differentiated from the hypanthium, calyx lobes 5-6, 5-9(-ll) mm long, 5-8 mm broad, broadly ovate and foliaceous with a petiole- like base; corolla white but drying dark, corolla tube 8- 24(-30) mm long, l-3(-4) mm diam., with whitish ap- pressed hairs ca. 0.5 mm long or glabrous, lobes 5, 10- 1 5(-30) mm long, 5 mm broad at the base. Fruits 30- 40 mm long, 25-35 mm diam., oblong-globose, persist- ing calyx 6-15 mm high.
Plants of lower montane evergreen forest for- mations, from 800 to 2300 m elevation. Flowering in February and April-May; fruiting in May, Au- gust, and November. This species ranges from the Cordillera de Tilaran to Veraguas, Panama.
Randia calycosa is recognized by its laminar calyx lobes, short spines, average-sized leaves, rel- atively large corolla lobes, and higher-elevation evergreen habitats. The flowers appear to vary greatly in size; for example, Almeda et al. 5078 F, from Monteverde, has corollas that are 6 cm long.
Randia formosa (Jacq.) K. Schum. is now called Rosenbergiodendron formosum (Jacq.) F. Faegerl. (Lorence in herb.); it ranges from central Panama to Peru. The corolla tubes are 8-10 cm long and densely whitish sericeous externally. The corolla lobes are 3-7 cm long and elliptic. The green fruits become 3 cm long and have broad white stripes down their sides and a narrow persisting calyx 5- 7 mm long.
Randia genipoides Dwyer, sp. nov. Figure 24.
Arbores ad 25 m altae. Folia anguste obovata vel el- liptica, ad 35 cm longa et 15 cm lata, venis lateralibus ca. 28; petiolis 5-8 mm longis; stipulis 9-25 mm longis. Inflorescentia flores femineas ca. 3 vel masculas 2-8 ge- rens. Flores hypanthio ca. 1 2 mm longo, calycis dentibus 5, 17-28 mm longis; corolla alba vel flavida pilis ca. 0.5 mm longis vestita, tubo 50-90 mm longo, lobis 5, 35- 50 mm longis. Fructus 6-8 cm longis, 4-7 cm crassus; seminibus ca. 10 mm latis.
TYPUS— G. Herrera 1 129 (holotypus CR, isotypi F, MO), from Upala, Alajuela, Costa Rica.
Small or medium-sized trees, 3-8(-25) m tall, dioe- cious, leafy stems 3-9 mm thick, glabrous and grayish, with prominent leaf scars, spines paired, 5-15 mm long, 1-3 mm thick, or often absent on distal twigs; stipules 9-25 mm long, lanceolate and acute, long (0.5-1 mm) whitish hairs present within, deciduous. Leaves closely clustered at the ends of stems, subsessile or with short (5-8 mm) petioles, ca. 2 mm broad; leaf blades 14-35 cm long, 5-12(-15) cm broad, oblanceolate to narrowly obovate or elliptic, apex abruptly narrowed and acu- minate with tip 6-10 mm long, base tapering gradually and cuneate, drying chartaceous, grayish or grayish brown above, glabrate above, minutely (0.2-0.4 mm) strigillose along the veins beneath, 2° veins 12-15/side, 3° veins often subparallel. Inflorescences of 3 terminal 9 flowers or 2-8 $ flowers at the tips of leafy branchlets, subsessile and subtended by lanceolate bracteate stipules 12-35 mm long and with thin hairs. Flowers unisexual, to 1 1 cm long, hypanthium ca. 12 mm long, not clearly dif- ferentiated from calyx, calyx tube 8-14 mm long, densely appressed-strigillose with hairs ca. 0.2 mm long, trun- cated distally and with 5 slender calyx lobes 1 7-28 mm long; corolla salverform, yellowish white with appressed- ascending whitish or yellowish hairs ca. 0.5 mm long, tube 50-90 mm long, 2-4 mm diam., lobes 5, 35-50 mm long and 7 mm broad near the base. Fruits 6-8 cm long, 4-7 cm diam., oblong-ellipsoid to subglobose, per- sisting calyx 5-10 mm long and 6-8 mm broad, sparsely and minutely puberulent; seeds ca. 1 0 mm broad, ellip- soid.
Plants of lowland rain forest formations, from 50 to 500 m elevation. Flowering in April-May and July; fruiting in June-September and Novem- ber (the type). The species is endemic to Costa Rica.
Randia genipoides is recognized by the larger
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narrowly obovate leaves closely clustered at the ends of stems, the greater number of secondary veins and subparallel tertiary veins, the large flow- ers sericeous or appressed-strigillose externally, long corolla lobes, and larger globose fruit. No other Costa Rican species of Randia has similarly clustered large oblanceolate leaves. Other collec- tions seen are Burger & Baker 9837 CR, F, Frankie 121a MO, Frankie 121c CR, F, and Zamora 998 CR. A fruiting collection (Herrera 4334 MO) from the Osa Peninsula is the only collection known from the Pacific lowlands.
Randia gentry! Dwyer, Ann. Missouri Bot. Gard. 67: 447. 1980.
Small trees to 10 m tall, leafy branchlets ca. 2-3 mm thick, glabrous; stipules not seen, caducous. Leaves strongly unequal at each node, petioles of the larger leaves 2-5 mm long, ca. 1.5 mm thick, glabrous; leaf blades of larger leaves 8-23 cm long, 2.2-7 cm wide, elliptic to oblong or narrowly elliptic-oblong, the smaller leaves subsessile with blades 1-4 cm long and ovate, apex acu- minate to acute (sometimes rounded in the small leaves), cuneate at the base, drying stiffly chartaceous, glabrous, 2° veins 8-12/side. Inflorescences axillary, peduncles to 5 mm long, bracts crowded and imbricate, flowers sol- itary, pedicels 0-3 mm long. Flowers glabrous externally except for minute red glands on hypanthium and calyx, hypanthium ca. 4 mm long, oblong, calyx tube 5-8 mm long, truncated, lobes 5, 2.5-3.8 mm long, ca. 1 mm wide; corolla white, carnose, tube 20-70 mm long, 2—4 mm diam., lobes 5, ca. 20 mm long, ovate-elliptic to oblong, strongly imbricate; stamens 5, anthers ca. 7.5 mm long; style ca. 12 mm long. Fruits 28-35 x 25-30 mm, oblong, rounded, glabrous, calyx ca. 3.5 mm long.
Plants of evergreen wet forest formations, 30- 1400 m elevation. The species has only recently been collected on the Osa Peninsula, with flowers in May (Hammel & Quintanilla 18217 MO) and immature fruit in August (Herrera 4133 MO). The species ranges to northwestern Colombia.
Randia gentryi is recognized by the nodes with opposite leaves differing greatly in size, the larger leaves over 1 2 cm long, the larger flowers, and the minute glandular dots on the lower parts of the flower.
Randia grandifolia (J. D. Smith) Standl., J. Wash. Acad. Sci. 18: 116. 1928. Basanacantha gran- difolia J. D. Smith, Bot. Gaz. 55: 436. 1913. Figure 23.
Shrubs or small trees, 2-6(-10) m tall, to 10 cm dbh, dioecious, leafy stems 1.5-8 mm thick, glabrous, the
outer grayish bark sometimes flaking off, spines 3-12 mm long, 1-2 mm thick, ascending, 1-2/node or often lacking; stipules 3-8 mm long, narrowed into a slender tip 1-2 mm long, brown, glabrous externally, with thin white hairs within. Leaves usually closely congested at the tips of branchlets, petioles 6-30 mm long, 0.8-2 mm broad, glabrous; leaf blades 7-22(-30) cm long, 3-10(-12) cm broad, elliptic, ovate, ovate-elliptic or elliptic-oblong to lanceolate, apex acute to short-acuminate, base obtuse to acute and slightly decurrent on petiole, drying thin- to thick-chartaceous, grayish green to yellowish green, glabrous above and beneath, 2° veins 6-1 I/side. Inflo- rescence terminal on leafy branchlets, of solitary 9 flow- ers or 1-5 $ flowers, subtended by triangular bracteate stipules ca. 3 mm long, flowers subsessile or with pedicels to 3 mm long. Flowers with hypanthium and calyx tube 4-10 mm long, 2-3 mm diam., calyx tube truncated distally with 5 slender (rarely expanded) calyx lobes 1- 4 mm long, sparsely puberulent with thin appressed hairs; corolla salverform, white, glabrous externally, tube 10- 18(-20) mm long, 2-3 mm diam., lobes 5, 8-12(-15) mm long, 2-9 mm broad. Fruits 3-4 cm long, 2.5-3.5 cm diam., globose to ellipsoid, slightly flattened on op- posite sides, greenish white becoming yellow or orange, pulp white to yellow, pericarp 1-2 mm thick; seeds ca. 8 mm long, rounded.
Plants of evergreen wet forest habitats on both the Caribbean and Pacific slopes of Costa Rica, from near sea level to 1 100 m elevation. Flowering in February-May and August; fruiting in April- January in Costa Rica. The species ranges from Mexico to Bocas del Toro, Panama.
Randia grandifolia is distinguished by its rain forest habitat, congested leaves varying greatly in size, shape, and texture, and flowers with relatively large corolla lobes. The calyx lobes may some- times be expanded. This is a fairly common spe- cies on the Caribbean slope but has been only rarely collected in the Golfo Dulce area on the Pacific side of Costa Rica (Tonduz 9982 us the type). It is often confused with R. armata.
Randia grayumii Dwyer & Lorence, sp. nov. Fig- ure 24.
Frutex vel arbor parva usque ad 4 m alta, ramulis inermibus. Folia chartacea, lamina elliptica, oblonga vel oblongo-elliptica, 18-30 x 8-16 cm. Inflorescentia staminata capituliformis, involucrata, flores 10-12 ge- rens. Flos lobis calycinis lineari-lanceolatis erectis, 3-4 x 0.5-0.7 mm; corolla infundibuliformi tubo gracili 22- 28 mm longo, 1-1.2 mm diametro; fauce sparsim villosa, 4-6 mm diametro.
TYPUS— Graynum & Schatz 5287 (holotypus MO, iso- typus ?CR), Costa Rica.
Dioecious shrubs or small spindly trees, 2-4 m tall, unarmed, leafy twigs 3-5 mm wide, often forked, finely hirtellous or glabrate, lenticellate, longitudinally wrin-
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285
kled, drying pale green or brown, internodes (0.5-)2-10 cm long; stipules 2.5-4.5 mm long, 2.5-3.5 mm broad, broadly ovate-triangular, the awn 0.5 mm long, brown- ish white, margins scarious, ciliolate, externally finely puberulent, internally with basal ring of sericeous white hairs and digitate white colleters. Leaves opposite, pet- ioles 1.5-3.5 cm long, 2-3 mm thick, often winged, finely hirtellous; leaf blades 18-30 cm long, 8-16 cm broad, elliptic, oblong or oblong-elliptic, apex deltoid with acu- men 1-2 cm long, base cuneate, usually abruptly atten- uate and decurrent, chartaceous and drying green, finely hirtellous above and beneath, glabrescent above, costa slender and sulcate above, to 2 mm wide and drying yellowish, 2° veins 9-12/side, festooned brochidodro- mous, forming an undulate submarginal vein. Male in- florescences terminal, 3-4 cm long and 4-5 cm broad (including corollas), condensed rounded capituliform 10- 12 flowered cymes, subtended by an involucre of ca. 12 overlapping ascending glumaceous ovate-oblong bracts in 4 rows, 3-12 x 2-6 mm, glabrous, margins ciliolate, pedicels 3-10 x 0.3-0.5 mm, finely hirtellous. Female inflorescences not seen at anthesis. Male flowers with hypanthium 0.8-1 .2 long, 1 .2-1.5 mm thick, calyx lobes 3-4 mm long, 0.5-0.7 mm broad, linear-lanceolate, stiff, erect, often involute, glabrous, yellowish white, margins ciliolate; corolla funnelform, white, drying black, tube 22-28 mm long, 1-1.2 mm wide medially, throat abrupt- ly flared to 4-6 mm wide, externally minutely puberu- lent, base of lobes and throat villous within, lobes 5, 7- 1 0 mm long, 5-7 mm broad, obovate-oblong, recurved; stamens with tips exserted, anthers 2.5 mm long, linear; style 22-28 mm long, stigma 0.5 mm long, clavate, tip exserted, disc conical, 0.5-0.6 mm long, glabrous. Fruits solitary, 3-5 cm diam., subglobose, borne on a short pedicel 2-5 x 3-4 mm, wall glabrous, tuberculate, 1- 1.5 mm thick, yellow or orange when ripe; seeds nu- merous, 8-10 mm diam., irregularly discoidal.
Understory plants of the Caribbean lowland ev- ergreen forest formations, 10-800 m elevation. Flowering in April-May; fruiting in January-April and September. The species ranges from near the Nicaraguan border in Costa Rica to central Pan- ama.
Randia grayumii is very closely related to R. armata and R. grandifolia but is distinguished by the large leaves with relatively few secondary veins and male inflorescences subtended by a small in- volucre of stipule-like bracts. Additional Costa Ri- can collections seen are Davidse & Herrera 31470 MO, Gomez et al. 21125 MO, Lent 524 F, Stevens 23659 MO, PTGB, and Stevens et al. 24831 MO. Panamanian collections seen are Kennedy et al. 3365 us and von Wedel 2342 us. [Descriptions provided by D. Lorence.]
Randia loniceroides Dwyer & Lorence, Bol. Soc. Bot. Mex. 47: 40. 1987. Figure 22.
Small shrubs, 0.5-1.5 m tall, dioecious, branching mostly horizontal, leafy branches 0.5-3 mm thick, with
ascending hairs ca. 0.3 mm long, glabrescent, spines to 6(-l 0) mm long and 0.5 mm thick, solitary, often absent; stipules 1-2 mm long, forming a short tube but splitting up and caducous. Leaves opposite along the stems or closely clustered on short-shoots, opposing leaves sub- equal or differing greatly in size, subsessile or with short- puberulent petioles 1-2 mm long; leaf blades 6-23(-30) mm long, 4-1 1(-15) mm broad, elliptic to obovate or narrowly obovate, apex bluntly obtuse or rounded, base tapering gradually and cuneate, decurrent on petiole, drying thin-chartaceous, dark grayish or grayish green, glabrous or with a few ascending hairs 0.2-0.3 mm long on the midvein above and below, 2° veins 3-4/side. In- florescences of solitary flowers, apparently axillary (ter- minal on very small axillary short-shoots), flowers sessile and subtended by small (1 mm) imbricated bracteate stipules. Flowers ca. 20 mm long, hypanthium 0.5-1.5 mm long, calyx tube ca. 1 mm deep and 1 mm broad, truncate or with narrow lobes; corolla salverform, white drying black, glabrous externally, tube 15-18 mm long, 0.5-1 mm diam., lobes 4 or 5, (3-)6-9 mm long, 3-6 mm broad. Fruits 10-15 mm diam., globose, becoming pale whitish green, surface smooth, pericarp ca. 1 mm thick.
Plants of lowland rain forest formations of the Caribbean slope, 20-500 m elevation (to 1 000 m in Mexico). Flowering in March-May; fruiting in January. This species is currently known only from Veracruz, Mexico, and northern Costa Rica.
Randia loniceroides is recognized by the very small leaves that may be subequal or strongly an- isomorphic, small stature with horizontal branch- ing, apparently axillary flowers and fruit, and rain forest habitat.
Randia matudae Lorence & Dwyer, Bol. Soc. Bot. Mexico 47: 42. 1 987. Casasia chiapensis Miran- da, Ceiba 4: 1 42. 1954, non R. chiapensis Standl. Figure 24.
Trees to 30 m tall, dioecious, leafy stems 3-10 mm thick, glabrous except above the stipule scars with slen- der hairs 0.5-1.5 mm long, bark of young twigs reddish brown and exfoliating in small pieces, spines absent; stipules 4-5 mm long, 3 mm broad at the base, triangular, glabrous externally. Leaves not closely crowded on the distal branchlets, petioles 6-22(-38) mm long, 1.3-3 mm broad; leaf blades 9-22(-29) cm long, 4-8(-12) cm broad, elliptic to elliptic-obovate or elliptic-oblong, apex acute or bluntly acute, base tapering gradually and cuneate or acute, slightly decurrent on petiole, drying stiffly char- taceous to subcoriaceous, usually very dark grayish above, glabrous above and below, occasionally with hairs or pits (domatia) in the vein axils beneath, 2° veins 5-97 side. Inflorescences of a solitary 9 flower or (1 1-)3-9 $ flowers, peduncles of the $ 6-10 mm long or absent, pedicels 4-8 mm long and not differentiated from the base of the hypanthium, glabrous, drying black. Flowers glabrous externally, drying black, hypanthium ca. 6 mm long, 2—4 mm diam., calyx tube 4—6 mm long, calyx
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lobes 4 (or with additional smaller lobes between), 1-3 mm long, narrow; corolla salverform, white, tube 30-40 mm long, 3-5 mm diam., lobes 4 or 5, 22-30(-40) mm long, ca. 10 mm broad near the base, acute. Fruits 5-7 cm long, 4.5-6.5 cm diam., globose to oblong, outer rind 5-10 mm thick (dried), persisting calyx to 8 mm high and 10 mm broad, pericarp 2-5 mm thick; seeds ca. 10 x 7 mm, elliptic.
Plants of wet evergreen cloud forest formations of the Cordillera de Tilaran at around 1300-1600 m elevation in Costa Rica but ranging from 20 to 2000 m in Guatemala. Flowering in June-August and October; fruiting in February, June, and Oc- tober. This species ranges from southern Mexico to northern Costa Rica.
Randia matudae is characterized by the large- sized tree habit, most plant parts turning dark on drying, larger leaves, and cloud forest habitat (in Costa Rica). Material of this species was incor- rectly referred to Genipa vulcanicola Standl. in the Flora of Guatemala (Standley & Williams, 1975); that species is now called Glossostipula concinna (Standl.) Lorence.
Randia mira Dwyer, Ann. Missouri Bot. Gard. 67: 450. 1980. Figure 24.
Shrubs or small treelets, 3-6(-8) m tall, dioecious, leafy stems 4-14 mm thick, glabrous, young stems with smooth reddish brown bark exfoliating in thin longitu- dinal strips, spines absent; stipules 6-10 mm long, to 10 mm broad at the base, triangular with a distal awn 0.5- 4 mm long, minutely appressed-puberulent. Leaves not usually closely spaced distally, petioles (5-) 12-25 mm long, 2-4 mm thick, minutely appressed-puberulent and glabrescent; leaf blades 24-44 cm long, 8-17 cm broad, elliptic to narrowly obovate or narrowly oblong-obovate, apex acute to acuminate, base gradually narrowed and cuneate, drying stiffly chartaceous to subcoriaceous, grayish green or dark olive green above, glabrous above, with few thin appressed hairs ca. 0.3 mm long on the surfaces beneath and glabrescent, 2° veins 10-16/side. Inflorescences with 1 -several flowers terminal on cau- liflorous short-shoots, flowers sessile and subtended by a small involucre of imbricate bracteate stipules ca. 3 mm long and with short (1 mm) awns. Flowers 10-15 cm long, minutely puberulent externally, drying black, hypanthium 3-4 mm long, ca. 2 mm thick, not clearly distinguished from the calyx, calyx tube 10-14 mm long, 5-7 mm diam., cylindrical with a truncated apex and short (1-3 mm) linear lobes; corolla tubular-sal verform, puberulent externally, tube 55-80 mm long, 3-5 mm diam., lobes 5-7, 45-70 mm long, 6-12 mm broad, lan- ceolate to narrowly elliptic-lanceolate, with narrowly acute apices; anthers 6-7. Fruits 4-7 cm diam., globose, yellow to yellow-orange, pericarp 5-10 mm thick; seeds 8-10 mm broad, ellipsoid.
Plants of lowland rain forest formations of the
Caribbean slope, 10-700 m elevation. Flowering in May-June (February-March in Panama); fruit- ing in October-January. The species ranges from northern Costa Rica to central Panama.
Randia mira is distinguished by the large cau- liflorous flowers with six to seven narrow corolla lobes, larger succulent or subcoriaceous leaves, lack of spines, thin exfoliating reddish brown bark on young stems, and lowland rain forest habitat.
Randia monantha Benth., PI. Hartw. 84. 1841. Basanacantha monantha (Benth.) Hook. f. ex Hemsl., Biol. centr. amer. Bot. 2: 39. 1881. B. subcordata Standl., Contr. U.S. Natl. Herb. 18: 133. 1916. R. subcordata (Standl.) Standl., Contra. U.S. Natl. Herb. 20: 201. 1919. Figure 23.
Shrubs or small trees, 3-6 m tall, dioecious, leafy stems 1-5 mm thick, with ascending whitish hairs ca. 0.5 mm long but quickly glabrescent, terete, brownish, spines 5- 15(-20) mm long, 1-2 mm thick, often 4/node, ascend- ing; stipules 2-5 mm long, 3-4 mm broad at the base, lanceolate with a narrowly acute apex, glabrous on the back and ciliolate along the edge. Leaves usually closely clustered at the ends of stems or on short-shoots, petioles 6-28(-35) mm long, 0.7-1.2 mm wide, with stiff hairs 0.3-0.9 mm long; leaf blades (2-)4.5-l 1 cm long, (l-)2- 6 cm broad, ovate to ovate-elliptic, broadly elliptic or elliptic-oblong, apex obtuse to acute (short-acuminate), often with a short acute tip, base cuneate to obtuse or acute or occasionally truncated and subcordate, drying membranaceous to chartaceous, dark or pale grayish above, with thin whitish hairs on both surfaces, 2° veins 3-7/side. Inflorescences of solitary 2 or few $ flowers, terminal on stems or short-shoots, flowers sessile and subtended by bracteate stipules. Flowers unisexual, the 3 to 7 cm long, appressed-puberulent externally, hypan- thium 5-15 mm long, ca. 4 mm diam., tubular, calyx tube 8-10 mm long, 3-4 mm diam., calyx lobes 5, 4- 8(-12) mm long, 1-1.8 mm broad, linear; corolla sal- verform, white to cream yellow, tube 30-45 mm long (to 60 mm in $ flowers), 0.8-2.4 mm diam., to 4 mm broad at the mouth, with thin erect whitish hairs exter- nally, lobes 5, 14-24 mm long (to 40 mm in $ flowers), 4-10 mm broad at the base, glabrous but with a ciliolate edge, ovate to ovate-lanceolate. Fruits 4-5(-6) cm long, 5-5.5 cm diam., globose to oblate, persisting calyx ca. 5 mm high and 6 mm broad, pericarp 3-4 mm thick (dried); seeds 8x5 mm, discoid, in a cream-colored pulp that becomes black.
Plants of seasonally very dry deciduous or partly deciduous forest formations of the Pacific slope, 20-900 m elevation (to 1400 m in Guatemala). Flowering in March-June; fruiting in all months except March and October. The species ranges from Mexico and Guatemala along the Pacific slope into central Costa Rica.
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Randia monantha is recognized by its deciduous forest habitat, stems usually with four spines at distal nodes, thin-pubescent leaves, larger puber- ulent unisexual flowers, and long calyx lobes. Cru- cilla is a common name.
Randia pittieri (Standl.) Standl., Contr. U.S. Natl. Mus. 20: 201. 1919. Basanacantha pittieri Standl., loc. cit. 18: 134. 1916. Figure 23.
Shrubs or small treelets, 1—4 m tall, leafy stems 2—4 mm thick, glabrous, brownish to grayish, terete, spines usually 4/node, 4-15 mm long, 1-3 mm at the base; stipules 7-12 mm long, 3-4 mm broad at the base, nar- rowly triangular and acute at the apex, brown, glabrous externally, thin white hairs within. Leaves closely clus- tered at the ends of stems, petioles 5-15(-24) mm long, 0.7-1 .5 mm broad, pubescent with stiff hairs; leaf blades 7-16 cm long, 4-10 cm broad, broadly ovate to elliptic- obovate or obovate, apex abruptly acute to gradually obtuse, often with a short (0.5-2 mm) sharp tip, base obtuse to slightly rounded, drying chartaceous, grayish above, with thin slightly curved hairs 0.2-1 mm long above and below, 2° veins 9-12/side. Inflorescences not seen, probably of solitary 9 flowers and several 3 flowers, terminal on the ends of leafy branchlets, pedicels of $ flowers 4-5 mm long. Flowers dioecious, calyx tube ca. 4 mm long, with subulate lobes 4-5 mm long, glabrous except for thin whitish hairs on the lobes and around the margin of the tube; corolla tubular-salverform, white, tube ca. 6 cm long, slightly expanded at the throat, lobes 25-30 mm long, lanceolate to elliptic lanceolate; anthers sessile in the throat, ca. 10 mm long. Fruits ca. 6 cm long and 5 cm diam., subglobose, with thin hairs or glabrescent, persisting calyx ca. 5 mm high.
Plants of evergreen rain forests and partly de- ciduous forests, from near sea level to 1000 m elevation. Flowering in February; fruiting in April. The species has been collected in the Caribbean lowlands (Pittier 16036 us the type), Cordillera de Guanacaste, and around the Meseta Central in Costa Rica.
Randia pittieri is distinguished by often having four spines at a node, thinner-textured pubescent leaves, corollas with long (6 cm) tube, and larger fruit. This species is not well collected; it is very similar to R. monantha and R. armata.
Randia pterocarpa Lorence & Dwyer, Bol. Soc. Bot. Mex. 47: 43. 1987.
Shrubs or small trees, 2-5 m tall, leafy stems 3-8 mm thick, glabrous, pale yellowish brown to dark brown, smooth or exfoliating, spines absent; stipules 3-6 mm long, intrapetiolar and forming a short adnate tube, tri- angular distally, glabrous, deciduous. Leaves opposite,
not closely congested distally, petioles 2-9 mm long, 2.5- 3.5 mm thick, drying dark, glabrous; leaf blades 16- 37(-50) cm long, 6-14(-17) cm broad, narrowly ovate- elliptic or elliptic-oblong, apex tapering gradually and subacuminate, base gradually narrowed but abruptly rounded or truncated at the petiole, drying stiffly char- taceous, dark grayish green, glabrous above and below, 2° veins 1 1-15/side. Inflorescences axillary or cauliflo- rous, of 1-5 flowers subtended by scarious bracteoles 1- 2 mm long, pedicels 1-4 mm long, not differentiated from the hypanthium. Flowers probably bisexual, gla- brous externally, calyx tube 1.5-3 mm long, 1.5-2.1 mm diam., truncated distally or with 5 small (0.2-1 mm) acute lobes; corolla salverform, white, tube 40-55 mm long, lobes 5, 25-38 mm long, elliptic-lanceolate. Fruits 5-10 cm long, 5-10 cm diam., obovoid, surface with 5 (9-11) longitudinal slightly elevated (1-3 mm) costae; seeds 10 x 8 mm, flattened, elliptic.
Plants of the very wet lower montane cloud for- ests of the Caribbean slope, 400-600 m elevation. The species is found in southern Mexico and in Costa Rica.
Randia pterocarpa is recognized by its long nar- row leaves, lack of pubescence, cauliflorous fruit with elevated costae, and limited altitudinal range (in Costa Rica).
Randia retroflexa Lorence & M. Nee, Brittonia 39: 371. 1987.
Lianas or shrubs with clambering branches, to over 13 m high with stems to 7 cm thick, dioecious, leafy stems 2-5 mm thick, pilose with spreading hairs 1-2 mm long, spines 5-10 mm long, 2-4 mm thick at the base, recurved (less often straight); stipules 6-10 mm long, 3-5 mm broad at the base, triangular to ovate, appressed-hirsute, persisting or deciduous. Leaves equal or subequal at each node, petioles 3-30 mm long, 1-2 mm thick, hirsute; leaf blades 6-23 cm long, 3-12.5 cm broad, elliptic to elliptic-obovate, apex acute to short- acuminate or obtuse, base obtuse to rounded or cuneate, drying membranaceous to chartaceous, appressed-hir- sutulous on both surfaces, 2° veins 5-9/side. Inflores- cences terminal, $ to 9 cm long and 20 cm wide (including corollas), candelabra-like and corymbose with 3-20 flowers, 9 flowers 2-6 sessile and subcapitate, peduncles 0-10 mm long. Flowers densely hirsutulous externally, hypanthium 2-3 mm long, obconical (3-6 mm long in 9), calyx tube 8-10 mm long, calyx lobes 5(-6), 1-12 mm long, 0.3-1 mm wide in <3, 2-4 mm wide in 9; corolla white, tube 4-5 cm long in $ (3-3.5 cm in 9), lobes 5, 1.5-4 cm long; anthers 10-12 mm long. Fruits solitary, to 7 x 6 cm, persisting calyx to 10 mm long and 7 mm diam.; seeds 8-10 mm diam., 1.5-3 mm thick.
Randia retroflexa is distinguished by its climb- ing or clambering branches, spine-bearing stems, hirsute pubescence, large flowers, and larger fruits.
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Two sterile collections (Haber et al. 4823 MO, Hammel & Haber 13921 MO) are provisionally placed here by David Lorence. He notes that they differ from Mexican collections by having spines that are straight and in groups of four but are similar in other aspects. Both collections were made in the Cordillera de Tilaran, between 600 and 1 200 m elevation. One collection was flowering in May. The species ranges to Mexico.
Randia thurberi S. Watson, Proc. Amer. Acad. Sci. 24: 53. 1889. R. crescentioides Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 88. 1944. Figure 22.
Small trees or shrubs, (2-)3-6 m tall, dioecious, branches usually held horizontally, puberulent in early stages, glabrescent and grayish, spines opposite, 6-20 mm long, 1.2-2.8 mm thick near the base, sharp-tipped; stipules less than 2 mm long. Leaves opposite or 4, on condensed (4 x 1.5 mm) short-shoots, 2 or 4, equal or unequal in size, petioles poorly differentiated, 0-8 mm long, puberulent or glabrous; leaf blades 6-40(-60) mm long, 4-18(-50) mm broad, obovate to obovate-subor- bicular, apex rounded to bluntly obtuse, base gradually narrowed and cuneate, decurrent on petiole, drying membranaceous to stiffly chartaceous, grayish green, gla- brous to densely puberulent with whitish hairs 0.2-0.4 mm long, 2° veins 3-5/side, ascending. Inflorescences of several or solitary flowers terminal on the ends of stems or short-shoots, subtended by a pair of thin brownish bracts 1-2 mm long, sessile. Flowers unisexual, hypan- thium ca. 2 mm long, obconic, glabrous to densely ve- lutinous, calyx lobes 1-2 mm long, acute; corolla fun- nelform, white, tube ca. 15 mm long, 0.9-2 mm diam., to 4 mm at the mouth, puberulent externally distally; lobes 7-10 long, 3-4 mm wide, broadly elliptic. Fruits 2.5-3.5 cm diam., globose, grayish green but turning black when dried, glabrous to densely puberulent, per- sisting calyx absent or small (1-2 mm high), 3-4 mm wide, pericarp ca. 3 mm thick.
Plants of seasonally very dry deciduous forests of the northwestern Pacific lowlands, 0-300 m el- evation (to 1 200 in Mexico). Fruiting in July-Oc- tober. This species ranges from Mexico along the Pacific lowlands of Central America to north- western Costa Rica.
Randia thurberi is recognized by its deciduous forest habitat, small often unequal obovate leaves usually clustered in 4's (2 pairs) at the ends of short-shoots, straight paired spines, and medium- sized flowers and fruits. Plants may differ greatly in the degree or absence of pubescence. At present the species is only known in Costa Rica from Santa Rosa National Park and Finca La Pacifica near Canas.
Randia vazquezii Lorence & Dwyer, Bol. Soc. Bot. Mex. 47: 37^48. 1987.
Vines or lianas, to 20 m high with trunks 1 8 cm diam., probably dioecious, leafy stems pilose to spreading pi- losulous, spines 3-4/node, 5-9(-30) mm long, ca. 2 mm diam., erect or reflexed; stipules 4-1 1 mm long, 3-5 mm broad at the base, thin, externally glabrous, deciduous. Leaves separated along the stem or clustered distally, petioles 7-20(-30) mm long, at first strigillose; leaf blades 8-16(-22) cm long, 4-9(-l 2) cm broad, elliptic to ovate- elliptic or obovate, apex acuminate with a tip 6-14 mm long, base obtuse to cuneate, drying chartaceous, grayish or brownish green, pubescent on both surfaces, 2° veins 6-10/side. Inflorescences terminal, of solitary 9 flowers or 1-4 fasciculate $ flowers, pedicels 5-6 mm long. Flow- ers with hypanthium 3-5 mm long, calyx tube 1-1 8 mm long, lobes 5, 6-1 5 mm long and linear in $, 1 5-20 mm long and oblanceolate in 9; corolla salverform, white or yellowish, strigose to sericeous externally, tube ca. 8 cm long in <?, ca. 5 cm long in 2, lobes 5, 30-35 mm long, 9-15 mm broad; anthers 8-9 mm long. Fruits ca. 9 cm long, 6-7 cm diam., ellipsoid, brown, warty, pericarp 3- 5 mm thick.
In wet forests at 400-1 200 m elevation. Fruiting in May and August. Known from Braulio Carillo N.P. and near San Isidro del General. The species also occurs in southern Mexico and Guatemala.
Randia vazquezii is recognized by its climbing habit, spiny stems, thin-pubescent leaves, and large flowers. Compare R. retroflexa with longer pu- bescence and shorter flowers.
Raritebe Wernham
REFERENCE— J. H. Kirkbride, Jr., Raritebe, an overlooked genus of the Rubiaceae. Brittonia 3 1 : 299-312. 1979.
Small trees or shrubs, without raphides, hairs with incomplete septa; stipules interpetiolar, free, entire, per- sisting. Leaves opposite or 3/node, petiolate; leaf blades entire, pinnately veined, domatia absent. Inflorescences terminal, open paniculate with cymose distal branches, pedunculate, the flowers pedicellate. Flowers bisexual and radially symmetrical, homostylous or heterostylous, hypanthium turbinate, calyx short-tubular, interior of the tube with 2-7 small glands near the base of each sinus, calyx lobes 4-5(-7), small or absent and the calyx entire, persisting; corolla salverform, white, lobes 5, val- vate in bud, distally carinate and the proximal part flat triangular; stamens 5, filaments borne on the tube near or below the middle, anthers free or coherent, 4-thecous, narrow, dorsifixed below the middle, obtuse to cordate at the base, the connective extended and acute, included, disc circular and entire; ovary 2-locular, ovules many on V-shaped to cordiform placentas, style glabrous, stigma bilobate and included. Fruits fleshy and indehiscent, green
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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becoming orange and finally black; seeds many and glob- ular.
A genus of six species (Dwyer, 1980) or only one species (Kirkbride, 1979) ranging from Costa Rica and Panama to Colombia, Ecuador, and Peru. The genus is closely related to Isertia and Am- phidasya according to Kirkbride (cited above, 1979, p. 302), but there are Costa Rican specimens placed in Coussarea that also seem to be close. The genus Dukea Dwyer is a synonym of Raritebe. Kirk- bride's broad interpretation is followed here.
Raritebe palicoureoides Wernham, J. Bot. 55: 337. 1917. Coussarea euryphylla Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 294. 1929. Bertiera panamensis Standl., N. Amer. Fl. 32: 180. 1934. C. colonis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 179. 1940. Dukea euryphylla (Standl.) Dwyer, Ann. Missouri Bot. Gard. 53: 366. 1966. Figure 43.
Shrubs or small trees, 1-7 m tall, leafy stems 1.8-7 mm thick, usually flattened (compressed) on opposing sides in early stages, with thin-ascending hairs 0.2-0.3 mm long; stipules 7-18(-24) mm long, 2.5-7 mm broad at the base, narrowly triangular and acute to long-acu- minate, puberulent on both surfaces. Leaves with peti- oles (5-)10-45 mm long, 1-2.5 mm thick, narrowly can- iculate above, minutely appressed puberulent; leaf blades 1 2-37 cm long, 4-1 4 cm broad, elliptic-oblong to elliptic or narrowly elliptic-oblong, apex acuminate with tip 10- 1 5 mm long, base acute to attenuate, drying thin-char- taceous, glabrous or very sparsely strigose above (the thin hairs difficult to see), with thin appressed-ascending hairs 0.2-0.5 mm long on the veins beneath or subgla- brous, 2° veins 8-13/side, arcuate-ascending and weakly loop-connected in the distal third of the blade. Inflores- cences (2-)6-18 cm long, (3-)5-20 cm broad, peduncle 1.5-7.5 cm long, appressed puberulent, branches op- posite and distant, bracts 1-5 mm long and usually lin- ear, stipule-like structures also present at the lower branching nodes, pedicels 1-8 mm long, slender and appressed strigulose. Flowers homostylous in subspecies dwyeranum, hypanthium 1.6-2.2 mm long, ca. 2 mm diam., calyx tube becoming 1-1.7 mm long, calyx lobes 0.1-0.4 mm long, broadly triangular to obscure; corolla tubular-salverform, white, tube (5-)6-9 mm long, 0.9- 1.4 mm diam., minutely puberulent with hairs 0.1-0.2 mm long, lobes (3-)4-8 mm long, 0.5-1.2 mm broad, linear to narrowly oblong, glabrous to sparsely puberu- lent externally; stamens with filaments 0.3-1 .6 mm long, anthers 2.2-4.4 mm long; styles 2-2.4 or 3.8-6.5 mm long, stigmas 0.9-2.3 mm long. Fruit 4-6 mm diam., globose, yellow-orange; seeds 0.2-0.6 mm diam.
Small trees and shrubs of evergreen rain forest and cloud forest formations of the Caribbean slope, from near sea level to 1400 m elevation. Probably
flowering and fruiting throughout the year (mostly flowering in March-August and fruiting in March- December). This species (considered in the wide sense) ranges from Costa Rica and Panama to Peru. Raritebe palicoureoides is recognized by its ap- pressed puberulence, the terminal compact to widely branched inflorescences, the small tubular flowers with poorly developed calyx lobes, the sta- mens cohering together at anthesis (in our sub- species), and the small fleshy fruit with many round seeds. Only subspecies dwyeranum Kirkbride oc- curs in Costa Rica and Panama; it is distinguished from other members of this species by having homostylous flowers with anthers 2.8-4.4 mm long and styles 4.2-6.5 mm long. There is a great deal of variation in the leaves and inflorescences of different individual plants in this species. We fol- low Kirkbride, who interpreted this species so broadly as to include all six species recognized by Dwyer (1980, p. 452). Specimens in early stages of flowering can be easily mistaken for specimens of some species of Coussarea.
Richardia Linnaeus
REFERENCE— W. H. Lewis and R. L. Oliver, Re- vision of Richardia (Rubiaceae). Brittonia 26: 27 1- 301. 1974.
Herbs, perennial or less often annual, decumbent and forming mats or erect, stems terete, usually densely pu- bescent; stipules interpetiolar, adnate to the petiole base and forming a truncated sheath with long distal setae (awns). Leaves opposite, sessile or short-petiolate; leaf blades linear to ovate, margins entire and often revolute, hirsute to scabrous or glabrous, domatia absent. Inflo- rescences terminal, capitate and sessile, subtended by an involucre of 4 decussate or 2 opposite (rarely 1) invo- lucrate bracts, the bracts similar to foliage leaves but often wider and palmately veined, the flowers closely crowded, pedicels absent or short. Flowers bisexual and monomorphic, radially symmetrical, with hypanthium turbinate to subglobose, calyx lobes 4-8, usually 2-3 times as long as the calyx tube, persisting; corolla fun- nelform or less often rotate to salverform, white to lav- ender (rarely yellow or orange), lobes 3-6, ovate to lan- ceolate, valvate in bud; stamens (3-)4 or 6, filaments short (long in a few species) and borne near the mouth of the corolla tube, anthers elongate or ovoid, dorsifixed, exserted; disc an unlobed ring or cylinder; ovary inferior or half-inferior, usually 3- or 4-locular (rarely 2 or 5-6), with 1 ovule affixed to the septum near the base in each locule, style simple or branched, stigmas capitate or spat- ulate. Fruits breaking up into 3 or 4 (2, 5-6) dry meri- carps (cocci), thin- to thick-walled, the mericarps usually obovoid and smooth to papillose, the adaxial face nar- rowly to broadly sulcate; seeds oval or ellipsoid, smooth, brown.
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A genus of 15 species in the tropics and sub- tropics of the Americas, from southeastern United States to northern Argentina. Most species occur in Brazil but none are found in the Amazon basin. A few species have become weeds in Africa and elsewhere. The genus is characterized by the short
weedy habit, usually narrow leaves attenuate at the base, terminal capitate heads of flowers, three- or four-locular ovaries, and small fruit breaking up into mericarps. This genus is closely related to Crusea and Diodia but differs in the usually three- carpellate ovary and the involucrate heads.
Key to the Species of Richardia
la. Mericarps papillose-muricate, with 1 narrow sulcus on the inner face; leaf surfaces very scabrous with short stiff hairs [a common species] R. scabra
Ib. Mericarps minutely hirsute, with 2 broad depressions and a median keel on the inner face; leaf surfaces smooth or only slightly scabrous above R. brasiliensis
Richardia brasiliensis Gomes, Mem. Ipecacuanha Bras. 31, t. 2. 1801. Richardsonia brasiliensis (Gomes) Hayne, Arzengew. 8: 21, t. 21. 1855.
Prostrate herbs, annual or perennial, often forming mats from a central taproot, stems to 50 cm long (rarely rooting from the nodes), leafy stems 0.3-4 mm thick, with thin spreading hairs 0.3-2 mm long; stipule sheaths 1-3.5 mm long, with 3-5 setae 1-5 mm long. Leaves with petioles to 1 5 mm long but poorly denned because of the decurrent leaf base; leaf blades l-4(-7) cm long, 4- 20(-27) mm broad, elliptic to ovate, apex acute, base gradually narrowed and attenuate, decurrent on the pet- iole, scabrous and with thin hairs on both surfaces. In- florescences 7-12 mm diam., usually with more than 20 flowers, the longer bracts 15-35 mm long, 6-20 mm broad, shorter bracts 10-17 mm long and 4-10 mm broad or sometimes absent. Flowers with hypanthium 1-2 mm long, calyx tube to 1 mm long, calyx lobes 6 (5), 1-1.5 mm long and 0.3-1 mm broad, lanceolate to ovate-triangular, margins with conspicuous cilia; corolla funnelform, white (rose), corolla tube 2-8 mm long, lobes 6 (4), 1-3 mm long and 0.5-1 mm broad; stamens 6 (4); ovary 3-locular, style 3-4 mm long, stigmas 3, 0.2-0.4 mm long. Fruit breaking up into 3 (2) mericarps 2-4 mm long and 1 .4-2 mm broad, oblong-obovoid, adaxial sur- face smooth and broadly concave with a longitudinal keel or rib; seeds ca. 2.5 mm long and 1.8 mm thick.
Weedy plants of open sunny sites in both ev- ergreen and seasonally deciduous areas, from near sea level to 2000 m elevation. Not yet collected in Central America but very likely to be present. The species, originally from Brazil, is now also found in Mexico, the southeastern United States, tropical Africa, southern Asia, and the Pacific.
Richardia scabra L., Sp. PI. 330. 1753. Figure 4.
Annual or perennial herbs, stems sprawling or erect, to 70 cm long and 30 cm high, not usually rooting at the nodes, leafy stems 0.7-3.5 mm thick, with stiff thin
spreading hairs to 1.8 mm long; stipule sheaths 2-4 mm long with 3-7 setae 1-5 mm long, difficult to see among the longer hairs at the nodes. Leaves with petioles 3-12 mm long, poorly differentiated from the lamina base; leaf blades (10-)20-45(-65) mm long, (3-)5-l 5(-23) mm broad, narrowly elliptic to elliptic-oblong or elliptic-ob- lanceolate, apex acute to obtuse, base attenuate and de- current on petiole, scabrous and with short (ca. 0.5 mm) stiff appressed or ascending hairs along the margins and on both surfaces, 2° veins 2-3/side and strongly ascend- ing. Inflorescences terminal and solitary, 6-16 mm diam., capitate, usually with more than 1 0 flowers, usually sub- tended by an opposite pair of ovate bracts 10-27 mm long and 6-15 mm wide and shorter (4-15 x 3-9 mm) bracts. Flowers with hypanthium 2-3 mm long, calyx tube ca. 1 mm long, calyx lobes 6, 1.8-3 mm long, ca. 1 mm wide, hispid or with a ciliate margin; corolla fun- nel-form to salverform, white or pale pink or marked with pink, glabrous, tube (2-)4-7 mm long, lobes 6 (5), 0.8-3 mm long and 1-1.5 mm wide; stamens 6 (5), ex- serted or included, anthers 0.4-0.8 mm long; ovary usu- ally 3-locular, style ca. 7.5 mm long, stigmas 3, ca. 0.7 mm long. Fruits usually with 3 mericarps 1.8-2.7 mm long and 1-1.8 mm broad, plano-convex and oblong to obovoid, inner face with a narrow longitudinal sulcus, muricate to papillate on the abaxial surface with the papillae larger (0.3 mm) distally; seeds 2-2.5 mm long and 1.3 mm thick, purplish brown.
Common weedy plants of open sunny sites in both seasonally deciduous and evergreen forest formations, from near sea level to 1800 m ele- vation. Flowering throughout the year but pri- marily in the wet season (June-December) in de- ciduous formations. The species ranges from the Carolinas, in the eastern United States, southward through Mexico and Central America to northern South America, Cuba, and Jamaica. It has also become naturalized in parts of Africa, southeast Asia, and the Pacific.
Richardia scabra is recognized by the usually procumbent mat-forming pubescent stems, sca- brous leaves attenuate at the base, small terminal
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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capitate inflorescences consistently subtended by a pair of broad leaf-like bracts, flowers with six stamens, and fruit usually with three mericarps. The mericarps, borne on the exterior of the fruit, can be mistaken for muricate-papillate seeds when they fall off. The dried plants have a slightly gray- ish appearance due to the thin transparent hairs on many parts.
Rondeletia Linnaeus
REFERENCES— J. H. Kirkbride, A revision of the Panamanian species of Rondeletia (Rubiaceae). Ann. Missouri Hot. Gard. 55: 372-391. (1968) 1969. D. H. Lorence, New species and combina- tions in Mexican and Central American Ronde- letia (Rubiaceae). No von 1: 135-157. 1991.
Shrubs or small trees, branchlets terete or angular, usually puberulent; stipules interpetiolar, rarely slightly united above the petioles and forming a very short sheath (intrapetiolar), variable in size and form, usually trian- gular (rarely bilobed). Leaves opposite (rarely 3/node), petiolate or rarely sessile; leaf blades entire, membra- naceous to coriaceous, glabrous to densely puberulent, domatia often present. Inflorescences terminal or axil- lary, paniculate with opposite branching, cymose to long- thyrsoid (rarely racemose or spike-like), pedunculate, bracts small to large, flowers sessile to pedicellate. Flow- ers bisexual and radially symmetrical, often dimorphic, hypanthium rotund to oblong, usually puberulent, calyx tube absent or very short, calyx lobes 4-5 (6, 7), equal or often unequal; corolla salverform, pink to rose red, reddish purple, yellow, or white, corolla tube usually narrow, the interior of the throat glabrous with an an- nular thickening at the mouth (in Rondeletia sensu stric- to), yellow-bearded at the mouth (Rogierd) or glabrous without a callosity at the mouth (Arachnothrix), lobes 4-5(-6), usually spreading, imbricate in bud; stamens 4- 5(-6), filaments usually very short and attached in the upper half of the tube, anthers dorsifixed, oblong and introrse, included (rarely exserted); ovary 2-locular, ovules many in each locule on placentas borne on the septum, style slender, stigmas narrowly 2-lobed. Fruit a thin- walled capsule, globose to ovoid, oblong or oblate, 2-locular, dehiscing loculicidally or septicidally, the valves often opening at the top and bipartite from the top; seeds
many, oblong and winged at both ends (in Rondeletia sensu stricto) or rhomboid to triangular and usually with- out wings or ribs (in Arachnothrix and Rogierd).
A genus of about 125 species, ranging from the West Indies and Mexico, through Central America into tropical South America. Important centers of diversity are located in Central America and the West Indies. Rondeletia is recognized by its rela- tively small flowers with narrow tubes, the calyx lobes often slightly enlarged or conspicuously un- equal, the thin-walled capsules, and the narrow winged seeds (in some species). The genus is not represented in Costa Rica's Caribbean lowlands, except near the Panama border.
As observed by Williams (Standley & Williams, 1975, p. 178), Central America has an unusual number of narrowly endemic Rondeletia species. Among our species, R. aspera, R. calycosa, R. cha- conii, R. cooperi, R. monteverdensis, R. povedae, R. stenostachya, R. tayloriae, and R. urophylla ap- pear to have very restricted ranges. Another un- usual feature of some species is the near-identity of form between floral bracts and the expanded calyx lobes (an example of homeosis or hetero- topy). Some of our species are easily mistaken for species of Gonzalagunia and even Buddleja (Lo- ganiaceae).
The circumscription of Rondeletia has not been resolved. Steyermark (1974, p. 296) and Borhidi (Bot. Kozlem. 62: 27. 1975) accepted Arachno- thrix as a separate genus, and Borhidi accepted Rogiera and Javorkea as well (Acta Bot. Hung. 28: 67. 1982). Kirkbride (cited above) suggested that species in the centers of diversity need to be better understood before generic concepts are redefined, and Lorence (cited above) recently argued for a broad interpretation of the genus. We have ben- efitted from the annotations and advice of David Lorence, who is studying this genus in Central America and Mexico; he has contributed the de- scriptions of three new species (see below).
Key to the Species of Rondeletia
la. Leaves with a dense white tomentum with persisting arachnoid hairs beneath; calyx lobes small and equal or subequal 2
Ib. Leaves not persistently whitish tomentulous beneath, lower leaf surface visible between the minor venation in mature leaves, arachnoid hairs only on young shoots if present; calyx lobes equal to
very unequal 3
2a. Inflorescences long-thyrsoid; weedy and widespread, 500-2000 m elevation
. R. buddleoides
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FIELDIANA: BOTANY
2b. Inflorescences subcapitate; endemic, Pacific slope, 600-1200 m elevation R. aspera
3a. Inflorescences long-thyrsoid, much longer than wide 4
3b. Inflorescences paniculate, usually as wide as long 7
4a. Some flowers or flower groups sessile and widely separated along the rachis; fruits fleshy, not
forming a dehiscent capsule Gonzalagunia stenostachys
4b. Flowers and flower groups borne on short lateral branches of the inflorescence rachis; fruits
becoming dehiscent capsules 5
5 a. Flowers borne closely together on the branches of the inflorescence; calyx lobes small and
subequal [corolla tube usually glabrous; usually trees] R. brenesii
5b. Flowers borne slightly distant on the lateral branches of the inflorescence; calyx lobes prom- inent and distinctly unequal 6
6a. Each flower with 1, 2, or 3 calyx lobes distinctly enlarged, corolla tube glabrous or sparsely
puberulent externally; shrubs; Golfo Dulce area R. urophylla
6b. Usually with only 1 calyx lobe enlarged on each flower; corolla tube puberulent; trees of the
Caribbean lowlands R. bertierioides
la. Leaf blades subcoriaceous in texture, becoming bullate in age, minor venation raised beneath and hirsutulous, often broadly ovate; corolla yellow barbate in the throat; stipules usually becoming
reflexed [to 17 mm long; widespread, 1000-2500 m elevation] R. amoena
7b. Leaf blades membranaceous to chartaceous, never bullate in age, minor venation flat beneath, glabrescent to sericeous, blades never broadly ovate; corolla not yellow barbate in the throat;
stipules not becoming reflexed 8
8a. Stipules leaf-like in form, with a narrow petiole-like stalk and rounded distal blade [leaves 2/node,
Caribbean slope, 500-1200 m elevation] R. costaricensis
8b. Stipules not leaf-like in form, broad at the base and without a petiole-like base 9
9a. Leaves usually 3/node 10
9b. Leaves usually 2/node 11
lOa. Corolla tube glabrous on the exterior, 11-15 mm long; petioles 2-10 mm long; western
cordilleras at 1200-1700 m elevation R. monteverdensis
lOb. Corolla tube strigulose on the exterior, 10-11 mm long; petioles 1-2 mm long; ca. 700 m
elevation on the Pacific slope R. povedae
1 la. Calyx lobes subequal or equal 12
1 Ib. Calyx lobes unequal 14
1 2a. Calyx lobes to 6 mm long and lanceolate; leaves to 9 cm long; corolla usually pinkish or rose
red [Central Highlands at 1 100-1700 m elevation] R. calycosa
12b. Calyx lobes to 2.5 mm long and triangular to ovate or elliptic; corolla white or yellowish
13
13a. Calyx lobes to 2.5 mm long, broadly elliptic to ovate or obovate; corolla tube 9-10 m long;
Caribbean slopes of the Talamanca range 800-1400 m elevation R. tayloriae
13b. Calyx lobes to 1.5 mm long, narrowly triangular; corolla tube 5-9 mm long; deciduous
lowlands of Guanacaste R. hamelifolia
1 4a. Plants of the Caribbean lowlands of southern Costa Rica and western Panama [corolla tubes 9-
1 2 mm long] 15
14b. Plants of the central cordilleras, 600-2000 m elevation 16
1 5a. Inflorescences with 2-3 pairs of lateral branches; enlarged calyx lobes puberulent
R. cooperi
1 5b. Inflorescences with 4-6 pairs of lateral branches; enlarged calyx lobes glabrescent
R. bertierioides
16a. Larger leaf blades lanceolate to elliptic, usually 3—4 times longer than wide, petioles to 10 mm long [small calyx lobes 0.5-2 mm long, large calyx lobes 6-12 mm long and bluntly rounded
distally; 0-1000 m elevation] R. salicifolia
1 6b. Large leaf blades elliptic, ca. 2 times as long as wide, petioles to 40 mm long 17
17a. Small calyx lobes 1-3.5 mm long and linear, large calyx lobes 2-6 mm long; 900-2200 m elevation
R. torresii
1 7b. Small calyx lobes 3-5 mm long and subulate, large calyx lobes 6-9 mm long; rarely collected plants of the wet Caribbean slope at ca. 700 m elevation R. chaconii
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 293
Rondeletia amoena (Planch.) Hemsl., Diagn. PI. Nov. Mexic. 26, 1879. Rogiera amoena Planch., Fl. Serres 5: 442. 1849. Rondeletia versicolor Hooker, Bot. Mag. 77: t. 4579. 1851. Rondeletia latifolia Oerst., Vidensk. Meddel. Dansk. Na- turhist. Foren. Kjobenhavn 1852: 43. 1853. R. rugosa Benth. ex Oersted, loc. cit. 43. 1853. R. pittierii K. Schum. & Krause, Bot. Jahrb. Syst. 40: 316. 1908 (photo from B at F). Figure 41.
Shrubs or small trees, 2-7 (-10) m tall, leafy branchlets 2-6 mm thick, with soft erect or ascending yellowish hairs 0.2-0.5 mm long, becoming brown; stipules 4- 13(-17) mm long, 3-10 mm broad, triangular, sericeous externally, usually glabrous within, becoming reflexed, persisting or deciduous. Leaves opposite, petioles 3- 12(-18) mm long, 1.5-3 mm thick, densely yellowish puberulent; leaf blades 6-16(-22) cm long, (3-)4-9(-12) cm broad, broadly ovate, to ovate-elliptic or ovate-ob- long (rarely broadly elliptic), apex acuminate (rarely abruptly acute), base broadly obtuse to rounded and sub- truncate, drying subcoriaceous or stiffly chartaceous, up- per surface with thin strigose hairs 0.3-0.7 mm long (dense over the midvein), lower surface with similar yel- lowish erect hairs, 2° veins 5-8/side, the 3° veins prom- inent beneath, veins often becoming impressed above in older leaves. Inflorescences usually terminal, 5-18 cm long and almost as broad, rounded and open paniculate with 2 or 3 pairs of opposite branches, peduncles 2-10 cm long, first pair of lateral branches much larger than the distal (the inflorescence often somewhat tripartite), branches sometimes subtended by small leaves, reduced leaves or small narrow bracts, densely yellowish seri- ceous-velutinous, pedicels 0-3 mm long. Flowers with hypanthium ca. 1.5 mm long, urceolate or subglobose, densely velutinous with ascending pale yellowish hairs, calyx lobes 4-6, 0.7-1.5 mm long, oblong and slightly unequal, glabrous within; corolla white or pink, minutely puberulent externally, tube 9-14 mm long, ca. 2 mm diam. distally, with dense erect yellowish hairs 0.5 m long at the mouth of the tube, lobes 5-6, 1 .5-3 mm long, ca. 1.2-2 mm broad, rounded distally; stamens with fil- aments 1.5-2.5 mm long, anthers ca. 2 mm long. Fruits 3-5 mm long, 3-6 mm broad, oblate-subglobose with 2 longitudinal sulci on opposite sides, woody, loculicidal, calyx lobes reflexed or deciduous, glabrescent in late stages.
Plants of evergreen montane forest formations, from 1000 to 2000(-2500) m elevation. Flowering is primarily in November-February in Costa Rica; fruiting throughout the year. It has been collected in the Cordillera de Tilaran (Monteverde), on the western and eastern edges of the Meseta Central, and in the Cordillera de Talamanaca. The species ranges from southern Mexico to western Panama.
Rondeletia amoena is recognized by its thick broadly ovate leaves often becoming bullate in age, reflexed stipules, open panicles often with three main branches, pink flowers with yellowish hairs
in the throat, and the small rounded two-locular capsules. This common species is quite different in appearance from our other species of Rondeletia and explains Planchon's using this species as the basis for his genus Rogiera. This species has been used as a garden ornamental and is known as quina, quina falsa, and teresa.
Rondeletia aspera Standl., N. Amer. Fl. 32: 54. 1918. Figure 41.
Shrubs or small trees to 1 2 m tall, trunk to 25 cm dbh, leafy branchlets 2-3.5 mm thick, covered at first with soft white arachnoid (and sericeous) hairs but quickly becoming glabrous and reddish brown to very dark brown; stipules 2-6(-10) mm long, broadly triangular (when short) to oblong-lanceolate, erect, puberulent, deciduous. Leaves often clustered at the end of branches, petioles 3-8(-12) mm long, 0.5-1 mm thick, puberulent; leaf blades 5-10(-12) cm long, 1.5-3.5(-4) cm broad, lan- ceolate to lanceolate-oblong or narrowly ovate-elliptic, apex tapering gradually and acute or acuminate, base acute to obtuse, drying thin-chartaceous, usually dark green above with soft or slightly rough hairs (scabrous in the type), at first densely arachnose tomentulose be- neath and covering the entire surface, the older leaf sur- face remaining white or becoming green with thin hairs 0.3-0.5 mm long beneath, 2° veins 5-8/side and strongly arcuate ascending. Inflorescences terminal, 2.5-7 cm long, compact subcapitate cymose panicles, peduncle 5-30 mm long, white tomentulose, the flowering portion as broad as long and the flowers closely crowded, subsessile. Flow- ers with hypanthium 1.5-2 mm long, ca. 1.3 mm diam., with ascending sericeous and tomentulose white hairs, calyx lobes 0.5-1.5 mm long, narrowly triangular to rounded, equal or subequal; corolla whitish to reddish brown, tube 8-1 1 mm long, 0.7-1.5 mm diam., with soft thin white hairs externally, lobes 4, 2.5-3 mm long, 1.5- 2 mm broad, glabrous within. Fruits 3-6 mm long, 3-6 mm broad, puberulent, closely clustered, at first opening partially septicidally and then each half opening (locu- licidal) at the top.
Plants of partly deciduous forest formations on the Pacific slope, from about 600 to 1200 m ele- vation. Flowering in December-February; fruiting in April, June, and August. The species is only known from the region between Monteverde and Ciudad Colon (San Jose) in central Costa Rica.
Rondeletia aspera is recognized by its smaller leaves crowded at the ends of slender branches, lamina that have a dense white arachnoid tomen- tum beneath and dry dark above, and the small subcapitate inflorescences. The restricted range of this species is another important distinction. This species would have to be transferred to Arach- nothrix, if that genus were accepted.
294
FIELDIANA: BOTANY
Rondeletia bertierioides Standl., Publ. Field Co- lumb. Mus., Hot. Ser. 4: 267. 1929.
Small trees to 6 m tall or rarely shrubs, leafy branchlets 1-3 mm thick, arachnoid-tomentulous and whitish in early stages, soon glabrescent; stipules 3-5(-7) mm long, triangular to subulate with a narrow tip 1-3 mm long, puberulent at the base, glabrous distally and drying dark. Leaves opposite and equal or unequal, petioles 4-1 2(-30) mm long, 0.5-1 mm thick, with thin whitish arachnoid and appressed hairs in early stages but quickly glabres- cent; leaf blades (4-)6-15(-22) cm long, 1.5-5(-8) cm broad, narrowly elliptic to elliptic or narrowly ovate- elliptic, apex tapering gradually and acuminate (acute), base acute to obtuse and slightly decurrent on petiole, drying thin-chartaceous, dark brown above, quickly be- coming glabrous above, appressed-puberulent along the veins beneath or glabrescent, 2° veins 5-7/side, 3° veins subparallel. Inflorescences terminal (rarely axillary), 7- 20 cm long, 3-8 cm broad, narrowly pyramidal with 5- 6 pairs of short helicoid-cymose lateral branches 1-3 cm long, peduncles 2.5-5 cm long, 0.7-1.4 mm thick, ap- pressed-tomentulose, bracts 2-4(-9) mm long, linear to lanceolate, pedicels 0-2(-3) mm long. Flowers with a hypanthium 1-1.5 mm long, ca. 0.7 mm diam., ap- pressed-tomentulose, calyx lobes 4, the 3 small lobes 0.3-2 mm long, triangular to oblong or linear, the long calyx lobes 1.5-4(-12) mm long, l-3(-5) mm wide (usu- ally 3 times longer than the small lobes), white; corolla white tinged with pink, tube 9-12 mm long, ca. 0.8 mm diam., puberulent externally, lobes 4, 2-2.7 mm long, broadly rounded distally; anthers 2-2.7 mm long. Fruits ca. 4 mm long, ellipsoid with longitudinal ribs (costae), glabrous at maturity, septicidal, calyx lobes persisting and perhaps enlarging.
Plants of the lowland Caribbean rain forest for- mations and premontane rain forest formations, from near sea level to 1000 m elevation. Flowering in February, April, July, and September. Two atypical Costa Rican collections are tentatively placed here (see below). Otherwise, this species is only known from the provinces of Bocas del Toro, Code, and Veraguas in western Panama.
Rondeletia bertierioides is recognized by the long narrow or pyramidal inflorescences with slender corolla tubes, the calyx with a single enlarged lobe, and arachnoid hairs on newly developing plant parts. Included here are Barringer et al. 3523 (CR, F, MO) and Burger & Antonio 10896 (CR, F) col- lected near BriBri in southern Limon Province, Costa Rica. These have more broadly (to 10 cm) paniculate inflorescences and conspicuously en- larged calyx lobes to 1 2 mm long and 5 mm broad. The young parts are arachnose tomentulose (in the Barringer collection) and other features conform to the measurements given by Kirkbride. The type (Cooper 12231 us) has narrow (3 cm) inflores- cences and calyx lobes only 1-3 mm long.
Rondeletia brenesii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1370. 1938. Gonzalagunia exaltata Standl., loc. cit. 22: 385. 1940. Figure 21.
Trees, 8-18 m tall, trunks 20-30 cm diam., leafy branchlets 3-4 mm thick, with thin appressed hairs but soon glabrescent; stipules 7-15 mm long, united above the petioles for a short (1-4 mm) distance or overlapping to form a short tube, with a narrow attenuate tip 2-7 mm long, with thin whitish hairs or glabrescent. Leaves often closely clustered at the tips of branches, petioles 6-20 mm long, 1-1.7 mm thick, puberulent with lateral margins continuous with the lamina margins; leaf blades 1 1-22 cm long, 3-6.5 cm broad, narrowly elliptic-oblong to oblong-oblanceolate or elliptic, apex tapering gradu- ally and narrowly acuminate with tip 7-25 mm long, base gradually narrowed and attenuate, decurrent on the petiole, drying thin-chartaceous, brown above, glabres- cent above, with very thin soft whitish hairs 0.5-1 mm long beneath, 2° veins (8-) 11-1 5/side, strongly ascending but not loop-connected near the margin. Inflorescences terminal and solitary, 15-18 cm long, thyrsoid panicles only 2-3 cm broad, peduncles 1-2 cm long and 2 mm thick, the rachis pilose with thin whitish hairs, with 5- 1 5 closely crowded flowers on each lateral branch, sec- ondary peduncles 1-3 mm long, pedicels 0-1 mm long. Flowers with hypanthium 0.5-0.9 mm long, subglobose, calyx lobes 4, 0.5-1 mm long, subequal, triangular to acuminate, sparsely puberulent; corolla white, tube 4-8 mm long, 0.4-0.7 mm diam., glabrous or sparsely pu- berulent, lobes 4, ca. 1.5 mm long and 1.5 mm broad at the base, rounded distally; anthers ca. 1.5 mm long, in- cluded. Fruits dicoccous, mericarps 3 mm diam., subgl- obose, flattened laterally.
Plants of the evergreen premontane rain forest and moist forest formations (primarily on the Ca- ribbean slope), from 300 to 1100 m elevation. Flowering in February-April. This species is only known from the area between Volcan Arenal (Al- ajuela) and the western slopes of the Meseta Cen- tral, and with a single collection from near San Isidro del General (San Jose), in central Costa Rica.
Rondeletia brenesii is recognized by the narrow- ly elliptic leaves with many 2° veins, the unusual stipules, the long narrow thyrse-like inflores- cences, and small almost glabrous four-lobed co- rollas. The larger stature attained by this species is also unusual. Compare R. stachyoidea J. D. Smith of northern Central America, which has narrower leaf blades, fewer secondary veins, sessile flower groups, and a lower-elevation habitat.
Rondeletia buddleoides Benth., PI. Hartw. 69. 1 840. R. affinis Hemsl., Diagn. PI. Nov. Mexic. 28. 1 879. R. buddleoides var. aspera Kirkbride, Ann. Missouri Bot. Gard. 55: 379. 1968. Figure 21.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
295
Shrubs or small trees, 2-8(-12) m tall, trunks to 20 cm diam., leafy branchlets 1.2-6 mm thick, white to- mentulose at first but becoming glabrous and brown; stipules 4-1 1 mm long, 1-5 mm wide, usually narrow near the base and broader distally, oblong to oblong- obovate, often with a strong keel along the back, usually glabrous. Leaves with the petiole 3-20 mm long, white tomentulose when young; leaf blades 4-22 cm long, 1- 8 cm broad, narrowly elliptic to elliptic-oblong (less often somewhat ovate or obovate), apex acute or abruptly acu- minate, base obtuse to acute, drying thin- to stiffly char- taceous, upper surfaces becoming very dark after the thin hairs are lost, lower surfaces whitish on all surfaces with a dense matted tomentum of arachnoid and thin straight hairs, the major veins often darkly outlined against the white tomentum beneath, 2° veins 8-12/side, 3° veins subparallel. Inflorescences usually terminal, solitary, 14- 22(-30) cm long, 2-4 cm broad, elongate thyrsoid pan- icles, peduncles 3-30(-80) mm long, the lateral branches short (2-5 mm) and bearing cymes or helicoid branches of 3-12 flowers, rachis at first white tomentulose, brac- teoles 1-2 mm long, pedicles 0-1 mm long. Flowers with hypanthium ellipsoid to subglobose, 1-1.5 mm long, 0.5- 1.5 mm diam., white tomentulose, calyx lobes 0.3-1 mm long, equal; corolla white, rose or yellowish, tube 6-1 1 mm long, 0.5-1 mm diam., minutely tomentulose, lobes 4, 1.5-2.5 mm long, equally broad; anthers ca. 1.5 mm long. Fruits 3-4 mm long, subglobose, dehiscing septi- cidally.
Common shrubs and trees of evergreen for- mations on both the Pacific and Caribbean slopes, (600-)800-2000 m elevation. Flowering primarily in May-September; fruiting in January-Septem- ber. In Costa Rica, the species occurs along the central highlands from the Cordillera de Guana- caste to the eastern parts of the Cordillera de Tala- manca. Despite this range, the species appears to be absent in some areas, such as around the central volcanoes. The species ranges from southern Mex- ico to western Panama.
Rondeletia buddleoides is recognized by the usu- ally narrow leaves with a dense white tomentum beneath, the long narrow thyrsoid inflorescence with pink, white, or yellowish flowers, the slender corolla tubes, and the small septicidal capsules. This species resembles Buddleia americana (Lo- ganiaceae), in regard to both the leaves whitish beneath and the long narrow inflorescences with tubular flowers. It is not easily confused with any of our other species of Rondeletia.
Rondeletia calycosa J. D. Smith, Bot. Gaz. 56: 69. 1913. Figure 41.
Shrubs or small treelets to 5 m tall, leafy branchlets 1-3 mm thick, at first ascending sericeous with slender hairs ca. 0.3 mm long, quickly becoming glabrous and dark brown; stipules 2-3(-4) mm long, triangular to cus-
pidate with a narrow awn ca. 1 mm long, glabrous, per- sisting with the leaves. Leaves opposite, petioles (2-)3- 9 mm long, 0.5-0.8 mm thick, at first with ascending sericeous hairs; leaf blades 4-9 cm long, 1-3 cm broad, elliptic to elliptic-oblong or narrowly elliptic-obovate, apex acuminate with tip 8-14(-18) mm long, base acute or obtuse, drying stiffly chartaceous, much darker above than below, glabrous or with very thin hairs 0.3-0.8 mm long on the upper surface, with short (0.3 mm) ap- pressed-ascending hairs on the veins beneath, 2° veins 4-6/side, arcuate-ascending. Inflorescences terminal, 3- 7 cm long, corymbose-rounded to pyramidal, peduncles 1-3 cm long, basal branches often subtended by reduced leaves, flowers usually in distal 3-flowered cymes, bracts 2-4(-6) mm long, linear-oblong to narrowly elliptical, pedicels 1-5 mm long, appressed-puberulent. Flowers with hypanthium ca. 2 mm long and 1.3 mm diam., with ascending yellowish hairs, calyx tube ca. 1 mm long, calyx lobes 4, usually unequal with 1 (2) larger lobes 3- 5 mm long and 3 (2) smaller (1-2 mm) lobes, or occa- sionally the 4 lobes subequal and 3-6 mm long, linear- oblong; corolla red to dark rose or pink, tube 1 2-1 5(-l 7) mm long, 0.8-1 mm diam. throughout its length, ap- pressed-puberulent, lobes 4, 2-3 mm long, ca. 2.5 mm broad, rounded and undulate distally; style ca. 9 mm long, stigmas linear, 2.5 mm long. Fruits ca. 5 x 4 mm, ellipsoid, laterally flattened.
Shrubs of lower montane rain forest formations along the Caribbean escarpment, from 1100 to 1 700 m elevation. Flowering in August and Sep- tember-October. The species is only known from near Zarcero, east of Volcan Barva, Cerros de Zur- qui, and the La Palma-La Hondura area, in central Costa Rica.
Rondeletia calycosa is recognized by the small stature and small leaves on slender stems, the small open inflorescences with flowers in distal cymes, the large equal or unequal calyx lobes, and the long narrow reddish corolla tube. Development of the sepal lobes appears to differ significantly in different collections; compare material placed un- der R. torresii. A collection (Zuniga & Jimenez 15 CR, F) from Res. Biol. Carara at 300 m is tenta- tively placed here; it may be an unusual population or a related species.
Rondeletia chaconii Lorence, sp. nov. Figure 4 1 A.
Species Rondeletiae scabrae Hemsley affinis, sed ram- ulis dense velutino-tomentosis, foliis majoribus (6.5-)9- 20 x (3.5_)4-7.8 cm, stipulis grandioribus, 7-1 1 x 3.2- 5.2 mm, pedunculo 5 cm longo, corolla sericea differt.
TYPUS— /. A. Chacon 177 (holotypus CR 92850), from bosque socolado, finca de Bernardo Gomez, 700 m, Magsasay, Heredia. Costa Rica, 21 Jan. 1983. Figure 41A.
Small tree, 5 m tall, leafy twigs 2-3 mm diam., at first densely velutinous, the trichomes 1-1.5 mm long, light
296
FIELDIANA: BOTANY
brown, simple straight, septate, the internodes 1 .5-6 cm long; stipules 7-1 1 x 3.2-5.2 mm, ovate, interpetiolar, brown, externally strigose, internally white-sericeous, also with several dark brown colleters 0.5 mm long. Leaves opposite, those of a pair at a node subequal, petioles 5- 30 x 1-1.5 mm, strigose-hirsute; leaf blades (6. 5-)9-20 cm long, (3.5-)4-7.8 cm broad, elliptic, apex acuminate with tips 1.5-2 cm long, base acute or cuneate, occa- sionally slightly attenuate, stiffly chartaceous, discolor- ous, drying dark brown adaxially, abaxially paler brown, adaxially strigose-hirsute with scattered pale brownish white trichomes 0.5-1 mm long, trichomes denser along the costa, veins and margin, the pubescence slightly more dense abaxially, 2° veins 9-12/side, arcuate-ascending, brochidodromous, 3° veins forming a strong intersec- ondary cross-webbing pattern. Inflorescences terminal, becoming pseudoaxillary, cymose-corymbiform with 10- 1 5 sessile or subsessile flowers, densely hirsute-sericeous with pale brown or white trichomes, the peduncle 5 cm long, 0.6 mm diam., with 3 1° branches, each 1.2-1.4 cm long and bearing 1-3 cymules of 1-3 flowers, the branches, cymules and flowers basally bracteolate, brac- teoles hastate or lanceolate-subulate, 4-6 x 0.6-3 mm, sericeous. Flowers 4-merous, hypanthium ca. 1 x 1 mm, obovoid, densely sericeous, calyx cup 0.4 mm deep, ca- lyx lobes sericeous externally, strigose within, unequal, 1 large (6-9 x 2-3 mm) and foliaceous, venose, oblan- ceolate or elliptic, the smaller 3 subulate, 3-5 x 0.4-0.8 mm, each sinus with several short brown colleters; co- rolla white, salverform, tube 13-16 long, 1.2-1.4 mm wide distally, externally sericeous, internally pilosulous in basal 'A, lobes suborbicular, 2.5-3 x 2.5-3 mm, ex- ternally sericeous basally mixed with shorter hirtellous trichomes above and on margin, internally minutely pa- pillose-puberulent especially around throat, the margins often crisped; stamens in long-styled flowers included, attached 3.5-4 mm below throat, in short-styled flowers attached 2 mm below throat, filaments 0.5 mm long, anthers 2-2.2 mm long, linear-ellipsoid; style glabrous, in short-styled flowers included, 6.5-7 mm long, the stig- mas linear-ellipsoid, 2 mm long, in long-styled flowers 1 3 mm long with ovoid stigmas 0.5 mm long, disc pilose. Fruits and seeds not seen.
Plants of the wet evergreen Caribbean slope at 700-900 m elevation. Flowering in January and known only from the type collection and Bella 2082 CR, MO.
Rondeletia chaconii belongs to section Calyco- sae of subgenus Arachnothryx. It is most closely allied to the groups of large-flowered species with large, unequal calyx lobes and large white corollas including R. macrocalyx Standl. & Steyerm., R. uxpanapensis Lorence & Castillo-Campos, and es- pecially R. scabra Hemsl. (synonyms: R. mexiae Standl. and R. acuminata (Oerst. ex Standl.) Lo- rence & Castillo-Campos). It is easily distin- guished from these species by its long-pedunculate inflorescence. It is most unusual that the holotype, which consists of two twigs, has both short-styled flowers (left twig) and long-styled flowers (right twig). The collector's data state "Arbol de 5 m de
alto, conflores blancas, " but it is unlikely that both floral morphs would occur on the same plant. [Contributed by D. Lorence.]
Rondeletia cooper! Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 267. 1929.
Shrubs or small trees, 2-7 m tall, trunk ca. 5 cm diam., leafy branchlets 1-3 mm thick, puberulent with soft apressed-ascending hairs 0.2-0.5 mm long, with erect hairs ca. 1 mm long at the node (usually covered by the stipules); stipules 2-1 1 mm long, triangular (when short) or cuspidate with a narrow tip to 8 mm long, appressed- puberulent. Leaves with petioles 4-22(-30) mm long, 0.7-1.7 mm thick, puberulent; leaf blades (8-)12-22 cm long, (3.5-)5-9 cm broad, ellipic to elliptic-oblong or slightly obovate, apex acuminate with a narrow tip 5- 1 2(-25) mm long, base obtuse to acute, drying thin-char- taceous, glabrescent above, minutely (0.1-0.3 mm) ap- pressed puberulent on the veins beneath, 2° veins 5-87 side, arcuate ascending distally but not clearly loop-con- nected. Inflorescences terminal, 9-17(-25) cm long, to 1 0 cm broad, open panicles with 2-5 pairs of primary branches to 5 cm long, peduncles 4-16 cm long, ca. 1.5 mm thick, strigulose, flowers on distal cymose or scor- pioid axes, pedicels 0-1 (-3) mm long. Flowers with hy- panthium ca. 2 mm long and 1.3 mm diam., oblong, densely ferrugineous with ascending hairs, calyx tube ca. 0.5 mm long, calyx lobes 4, strongly unequal, the 1 large lobe 3-4 mm long and 1.5-2 mm broad, narrowly ovate to elliptic, the small lobes 0.7-1 mm long, ca. 0.4 mm wide, narrowly oblong; corolla white, tube 9-1 1 mm long, 0.6-1 mm diam., strigillose, lobes 4, 2-3 mm long. Fruits to 4.5 mm long and 3.5 mm diam., costa te, sep- ticidal to the middle, calyx lobes persisting.
Plants of lowland Caribbean rain forest for- mations, from near sea level to 400 m elevation. Flowering in the period of January-March (Cooper 600 F holotype, MO). This species is only known from Bocas del Toro, Panama.
Rondeletia cooperi is recognized by its lowland habitat, single enlarged calyx lobe on each flower, and narrow corolla tube. This is a very poorly known species, rather similar to R. bertieroides but lacking the arachnoid hairs in early stages and hav- ing widely branched inflorescences. However, it may prove to be synonymous with that species.
Rondeletia costaricensis Standl., N. Amer. Fl. 32: 61. 1918. Figure 41.
Shrubs or small trees, 2-10 m tall, leafy branchlets 1- 3 mm thick, glabrous from earliest stages (or with a few hairs at the nodes); stipules (3-)4-7 mm long, to 6 mm broad, leaf-like distally with a short (1 mm) petiole-like stalk and rounded lamina cordate at the base and emar- ginate or retuse at the rounded apex, glabrous or with a
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
297
few hairs at the base. Leaves with petioles 4-10(-l 6) mm long, 0.6-1.7 mm thick, glabrous, drying dark; leaf blades 7_14(_18) cm long, 2.3-5. 5(-8) cm broad, elliptic to el- liptic-oblong or slightly obovate, apex usually short- acuminate (acute), base obtuse to acute and slightly decurrent on the petiole, drying membranaceous or thin- chartaceous, usually dark in color above, glabrous above and below, 2° veins 4-6/side and weakly loop-connected distally. Inflorescences terminal, 4-9 cm long, to 8 cm broad, open panicles with 1 or 2 pairs of major opposite lateral branches, peduncles to 5 cm long and 1 mm thick, minutely (0.1-0.2 mm) appressed-puberulent, bracts 2- 4 mm long, linear, pedicels 0-3 mm long. Flowers with hypanthium 1-1.3 mm long, ca. 1 mm diam.. obconic to urceolate, densely sericeous with yellowish hairs, calyx lobes 4, to 0.7 mm long, equal, glabrous distally; corolla white, tube 6-9 mm long, 0.5-1.5 mm diam., minutely appressed puberulent, lobes 4, 1-3 mm long, rounded and undulate distally. Fruits 4-5 mm long, 3.5^4.5 mm diam., subglobose to turbinate in forms, slightly rounded at the truncated distal surface, persisting calyx lobes ca. 0.5 mm long; seeds ca. 1.2 mm long, broadly ellipsoid- lenticular, with a few hairs along the edge.
Plants of premontane rain forest formations on the Caribbean slope, from 600 to 1 200 m eleva- tion. Flowering in March-May; fruiting in June- October. This species is only known from the Ca- ribbean slope in the region between P. N. Rincon de la Vieja (85°19'W) and above Ciudad Quesada (84°27"W) in north-central Costa Rica.
Rondeletia costaricensis is distinguished by its unique stipules, thin glabrous leaves, small ter- minal inflorescences, small calyx lobes not differ- ing in size, and slender corolla tubes. The stipules are slightly intrapetiolar and united above the base for about 1-2 mm before becoming narrowed to the petiole-like base of the broad distal laminar part. This broad leaf-like distal part of the stipule is glabrous and has pinnate venation.
Rondeletia hamelifolia Dwyer & Hayden, Ann. Missouri Hot. Gard. 54: 144. 1967. Figure 41 A.
Small trees or shrubs, 5-8 m tall, leafy stems 1.5-3 mm thick, strigulose-sericeous and later glabrescent, te- rete; stipules 2-10 mm long, 1.5-5 mm broad at the base, triangular and acute, sericeous, persistent. Leaves with petioles 2-1 5 mm long, densely strigulose; leaf blades 4-19 cm long, 2-6 cm broad, elliptic, apex deltoid-acu- minate to acute, base cuneate, drying chartaceous, sparsely strigillose above and below (more densely on the veins), 2° veins 5-9/side. Inflorescences terminal, 4-13 cm long and 3-9 cm broad, open paniculate with 60-180 flowers in distal cymes, peduncles (l-)2.5-4.5 cm long, densely strigose-sericeous (as are branches, bracts and pedicels), bracts 1-4 mm long, triangular to linear, pedicels 0-^4 mm long. Flowers with subglobose hypanthium 1-1.2 mm long, calyx lobes 5, 0.5-2 mm long, linear; corolla salverform, white, tube 5-9 mm long, ca. 1 mm diam.,
lobes 5, 1.5-2.5 mm long, rounded. Fruits 3-7 mm long and ca. 5 mm wide, ellipsoid to subglobose, papery, dehiscing to half the length; seeds 1.2-1.9 mm long, fusiform with a wing at each end.
Plants of seasonally dry deciduous forest, 1 00- 250 m elevation. Flowers and fruit were collected in January (Liesner 4521 CR). In Costa Rica this species has only been collected from along Rio Guapote in the Santa Rosa National Park, Guan- acaste; it also occurs in Panama.
Rondeletia hamelifolia is recognized by its de- ciduous forest habitat, often larger leaves that ta- per gradually to both apex and base, and the slen- der white corolla tubes. The five-parted flowers, corolla with a fleshy ring around the throat, lo- culicidal capsules, and seed characters place this species in subgenus Rondeletia.
Rondeletia monteverdensis Lorence, No von 1: 147- 148. 1991. Figure 41.
Shrubs or small trees, 2-6(-16) m tall, trunk to 1 5 cm dbh, leafy branchlets 1-3 mm thick, with a few thin whitish hairs in the earliest stages and quickly glabrescent or glabrous throughout (but with erect tufted hairs at the node within the stipules), becoming dark brown; stipules 1-3 mm long, 1-2 mm wide with a short (0.5 mm) sheath, triangular or subulate, with a few hairs along the distal margins, sericeous within. Leaves usually 3/node, subequal to unequal (50%) at a node, petioles 2-6(-10) mm long, 0.5-1 mm thick, glabrescent; leaf blades (2-)3- 8.5 cm long, 1-2.5 cm broad, elliptic to narrowly elliptic- obovate or oblanceolate, apex abruptly narrowed to a long-acuminate with tip 7-15 mm long, base gradually narrowed and acute, drying stiffly chartaceous, dark above and pale grayish green beneath, glabrous or with scat- tered very thin whitish hairs to 1 mm long above, with few hairs or glabrescent beneath, 2° veins 3-6/side, as- cending. Inflorescences terminal, 3-7 (-12) cm long, 3-5 cm broad, pyramidal with 2 or 3 ternate nodes and 1 5- 30 flowers, peduncles 5-40(-60) mm long, 0.6-1 mm thick, glabrous or sparsely puberulent, primary bracts 5- 17 mm long, 1-3 mm broad, linear to narrowly oblong, floral bracts 4-5 mm long, pale brown and resembling the enlarged calyx lobes, essentially glabrous, palmately 3-veined, deciduous, pedicels 0-3 mm long. Flowers with hypanthium 1.5-2 mm long, obo void-ellipsoid, with thin white hairs, calyx lobes 4, unequal but all enlarged, (3-)5- 1 1 mm long, 1-2.5 mm broad, lanceolate, greenish white but drying pale brown, glabrous; corolla salverform, white, glabrous externally, tube 11-15 mm long, 0.5-1.2 mm diam., lobes 4, 2-4 mm long, 2-2.5 mm broad, broadly rounded distally; stamens sessile, anthers 2.5-3 mm long. Fruits 4-5 mm long, 2.5^4 mm broad, obovoid-ellip- soid, bisulcate, with 8 pale longitudinal ribs, apparently at first septicidally dehiscent and the 2 valves then open- ing at the apex.
Plants of lower montane and premontane rain
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forest formations in the northwestern cordilleras, from ( 1 200-) 1 400 to 1 700 m elevation. Flowering in February-August; fruiting in May and July- November. This species is only known from the Cordillera de Guanacaste and Cordillera de Ti- laran in northwestern Costa Rica.
Rondeletia monteverdensis is recognized by the small ternate leaves with prominent drip tips, the compact inflorescences with large bracts and near- ly identical large calyx lobes, and the glabrous white corollas. The similarity in form, texture, and color of the inflorescence bracts and enlarged sepal lobes is a striking example of homeosis or heterotopy: it is also seen in the closely related R. calycosa, where the bracts and calyx lobes are not as large and distinctive. A collection by G. Herrera (657 CR, F, MO) from 1200 m on Volcan Santa Maria differs from the Monteverde material in having narrower leaves, smaller bracts, and smaller calyx lobes.
Rondeletia povedae Lorence, sp. nov. Figure 41 A.
Species Rondeletiae monteverdensi Lorence affinis, sed inflorescentia ramulis petiolisque dense strigulosis, fo- liorum petiolis 1-2 mm longis, laminae pagina abaxiali tomentosa, inflorescentia subsessili, lobis calycinis mi- noribus ligulato-subulatis 1-2 x 0.3-0.5 mm differt.
TYPUS— L. J. Poveda 441 (holotypus CR 51260, iso- typus MO 2211249), from Santa Rosa de Puriscal, San Jose, Costa Rica, 17 Jan. 1973. Figure 41 A.
Shrubs, ca. 4 m tall, leafy twigs 1.2-1.5 mm thick, at first densely strigulose, the trichomes brownish white, simple, straight or curved, to 0.5 mm long, the internodes 0.3-3.5 cm long, bark brown; stipules 2-3.5 mm long, 1-2 mm broad, narrowly triangular, acuminate, intra- petiolar sheath 0.5 mm long, externally densely strigu- lose, internally densely white-villous, the margins and internal surface with several blackish digitate colleters. Leaves usually in whorls of 3, occasionally opposite, those at a node unequal, the smallest usually 50% the size of the largest, subsessile, petioles 1-2 mm long, 0.5- 0.8 mm thick, densely strigulose; leaf blades (3-)4-9.5 cm long, (0.9-)1.5-2.7 cm broad, oblanceolate or nar- rowly elliptic-lanceolate, apex long-acuminate and often falcate, tip 5-1 5 mm long, base acute or usually abruptly rounded or obtuse, chartaceous, strongly discolorous, drying dark brown adaxially and pale brown abaxially, adaxially sparsely arachnoid-pubescent with scattered curling trichomes, glabrescent, abaxially moderately arachnoid-tomentose with persistent white trichomes, strigulose along the costa, 2° veins 7-8/side, camptodro- mus, arcuate-ascending, a distinct marginal vein present, tertiary veins forming a strong intersecondary cross-web- bing pattern. Inflorescence terminal, often inclined, cy- mose-corymbiform, 12-16-flowered, densely strigulose with whitish trichomes, the slender peduncle 2-3.5 cm long, the rachis with l(-2) whorls of 3 1° branches 8-12 mm long, each ending in a 1-3-flowered cymule, the axes
bracteolate, the bracteoles subulate, 2-3 mm long, ped- icels 0.3-4 mm long, 0.2-0.3 mm thick (central flowers sessile). Flowers 4-merous, hypanthium 1.2-1.4 x 1.2 mm, ovoid compressed, densely strigulose, calyx cup 0.3 mm deep, lobes 4 (rarely 5), subequal to unequal, 1 usually larger than the other 3, linear-subulate to lan- ceolate, 1-2.5 x 0.2-0.5 mm, densely strigulose exter- nally, recurved, each sinus with 1-3 small brown col- leters; corolla salverform, white and pink, tube 10-1 1 mm long, 1.5 mm wide distally, externally strigillose, internally hirtellous in basal Vs, lobes irregularly subor- bicular, 2-3 x 2-3 mm, externally sparsely strigulose, internally glabrous, the throat glabrous, the margin un- dulate; stamens in long-styled flowers included, sessile, attached 5 mm below throat, anthers linear, 2.5 mm long; style in long-styled flowers 1 1 mm long, glabrous, the stigmas 2 mm long, narrowly ellipsoid, slightly ex- serted, the disc pilose without. Fruits broadly ovoid- subglobose capsules, 2-3 x 2. 5-4 mm, smooth, blackish, strigulose, bisulcate, dehiscence loculicidal then septi- cidal; seeds 0.7-0.9 x 0.4-0.5 mm, ellipsoid, light brown, narrowly winged, the wing broader at each end, the testa cells elongate, shallowly reticulate.
Plants of the seasonally dry Pacific slope at ca. 400 m elevation in central Costa Rica. This species is only known from the type collection.
Rondeletia povedae is recognized by the smaller often lanceolate leaves, usually three at each node, and the Pacific slope habitat. In terms of floral and capsule morphology, R. povedae is referable to subgenus Arachnothryx, section Calycosae, al- though the elongate seeds with a wing at each end and elongate testa cells with a shallow reticulum are suggestive of subgenus Rondeletia. This new species is most closely allied to R. monteverdensis Lorence, which also has ternate leaves but differs by the characters given in the diagnosis. [Contrib- uted by D. Lorence.]
Rondeletia salicifolia Dwyer & Hay den, Phyto- logia 15: 58. 1967.
Trees, 4-8 m tall, leafy branchlets with slender yel- lowish hairs; stipules to 4 mm long, triangular from a broad and thickened ring-like base, with a short curved awn, puberulent. Leaves with petioles 5-10 mm long; leaf blades 3-20 cm long, 2-9 cm broad, lanceolate to elliptic, the larger blades 3-4 times longer than wide, apex obtuse to slightly acuminate with narrow (1.5-2 mm) tip to 20 mm long, base obtuse to acute, drying stiffly chartaceous, dark brown, with minute yellowish hairs on the veins beneath, 2° veins 4-6/side. Inflores- cences terminal or axillary, paniculate with 2 or 3 pairs of opposite branches, peduncles to 6 cm long, lateral branches up to 12 cm long, tomentulose, flowers sessile in cymose or scorpioid arrangements. Flowers with hy- panthium ca. 2 mm long and tomentulose, calyx tube ca. 0.5 mm long, calyx lobes 4, strikingly unequal with
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
299
the single enlarged lobe 3-4 times longer than the short lobes, small lobes 0.7-3 mm long, the large calyx lobe 6-12 mm long and up to 7 mm broad, elliptic to ovate, bluntly rounded; corolla white, corolla tube 8-15 mm long, glabrate externally, lobes rounded; filaments to 2 mm long, anthers ca. 3 mm long. Fruit to 4 mm diam., subglobose, costate, the hairs deciduous.
Plants of premontane rain forest and cloud for- est formations in Panama, from near sea level to ca. 1000 m elevation. This species is reported to occur at the Monteverde Reserve. The species ranges into central Panama.
Rondeletia salicifolia is recognized by the nar- row leaves, and calyx with 1 enlarged lobe. Pho- tographs distributed as R. salicifolia H.B.K. do not represent a published name; they are R. leuco- phylla H.B.K. of Mexico (fide Lorence in herb.).
Rondeletia tayloriae Lorence, sp. no v. Figure 4 1 A.
Species Rondeletiae torresii affinis, sed lobis calycinis aequalibus vel subaequalibus foliaceis ellipticis ovatis vel obovatis 1.5-2.5 mm longis 0.8-1.5 mm latis, api- cibus obtusis vel rotundatis et corolla minore 2-3 mm longa, 1.8-2.4 mm lata differt.
TYPUS— Herrera & Chacon 2656 (holotypus CR, iso- typi MO, PTGB), from Fila Matama, Valle de la Estrella, 1350 m, Canton de Limon, Limon, Costa Rica, 19 Apr. 1989.
Small trees, 6-8 m tall, slender twigs 1 .2-3 mm diam., densely strigillose when young, trichomes 0.2-0.3 mm long, stiff and acicular, the internodes 1.5-12 cm long; stipules erect, rigid, 2-3.5 x 2.5-5 mm, deltoid to broad- ly deltoid, dark brown, densely strigillose externally. Leaves opposite, those of a node equal or subequal, pet- ioles 1-3.5 cm long, 1-1.2 mm diam., narrowly winged distally, densely strigillose; leaf blades (5.5-)6.5-19.5 cm long, (2-)3-8.2 cm broad, elliptic to ovate, apex acu- minate with an abrupt, straight or falcate acumen 1-2 cm long, base acute, usually attenuate and decurrent along the petiole, thinly to thickly chartaceous, drying brown- ish green, adaxially with scattered strigillose hairs on surfaces, denser on the veins and costa, glabrescent, abaxially densely strigillose, on veins and costa, hairs persistent, 2° veins 7-9/side, arcuate, brochidodromus. Inflorescences terminal, 5.5-1 6 cm long, 5-1 1 .5 cm broad, corymbiform-cymose with 20-30(-60) flowers, sessile or the peduncles 4-8 cm long, axes trichotomous and dense- ly fulvous strigillose-velutinous, branching to the 3°, with 2 pairs of 1 ° branches, these 1 -7 cm long, branching again l(-2) times dichasially, ending in cymules of 2-3 flowers, branches and cymules subtended by subulate bracteoles 1 mm long, pedicels 0-2 mm long, densely strigillose- velutinous. Flowers 4-merous, hypanthium 1.5-2 x 1.2- 1.5 mm, obovoid, calyx cup 0.6-0.8 mm deep, calyx cup and hypanthium densely strigillose-velutinous, calyx lobes subequal, 1.5-2.5 x 0.8-1.5 mm, broadly elliptic, ovate or obovate, apex obtuse or rounded, both surfaces strigillose, green when fresh; corolla white, salverform,
tube 9-10 mm long, 1.2-1.4 mm diam. medially, exter- nally densely strigillose-velutinous, internally hirtellous in basal 'A, lobes 2-3 x 2-3 mm, subcircular or obovate, spreading, densely strigillose basally, internally minutely papillose-puberulent, basally with larger white bulbous trichomes; stamens in long-styled flowers sessile, at- tached 3 mm below throat, anthers 2-2.5 mm long, el- lipsoid, included; style 10-11 mm long in long-styled flowers, tip exserted, glabrous except for a few hairs to- ward base, stigmas 0.8 mm long, ellipsoid. Fruits (im- mature) 6-7 x 4.5-5 mm, ovoid, bisulcate and crowned by the persistent calyx, strigillose; seeds not seen.
Plants of the premontane and montane tropical wet forest on the Caribbean slope of the Cordillera de Talamanca, 850-1350 m elevation. Flowers and immature fruits were collected in April. Only two collections have been seen, the type and Robles & Chacon 2664 (CR, MO, PTGB), also from the Fila Matama (Cerro Muchilla) at 850 m elevation, Li- mon Province. Endemic.
Rondeletia tayloriae belongs to subgenus Arach- nothryx, section Calycosae. This species is most closely related to R. torresii, which differs by its strongly unequal calyx lobes— the three small ones 1.5-3 mm long and linear to subulate with acute apices and the large one 4-5 mm long, obovate- elliptic to oblanceolate with an acute to obtuse apex— and larger corolla lobes 3-4 x 4-5 mm. This new species is also related to R. calycosa, a species that differs in having a pubescence of lon- ger silky trichomes, linear-lanceolate calyx lobes 3-7 mm long with acute or acuminate apices, and dark pink or rose red corollas. I take pleasure in naming this species for Charlotte M. Taylor. [De- scription and discussion contributed by D. Lo- rence.]
Rondeletia torresii Standl., J. Wash. Acad. Sci. 1 8: 163. 1928. Figure 41.
Shrubs or small trees, 1.5-5 m tall, leafy branchlets 1-3 mm thick, minutely puberulent with thin whitish ascending hairs 0.2-0.5 mm long, glabrescent; stipules 2-5 mm long, triangular, strigulose, deciduous. Leaves opposite, petioles 6-40 mm long, 0.5-1 mm thick, strigu- lose; leaf blades 5-12(-18) cm long, 2-5.5(-7) cm broad, elliptic to narrowly ovate-elliptic or elliptic-lanceolate, apex acuminate with tip 5-1 1 mm long, base acute to obtuse, drying thin-chartaceous, usually much darker above than below, glabrescent or minutely puberulent on the veins above, with short (0.2-0.4 mm) hairs along the veins beneath and occasionally with tufted hairs 0.5 mm long in some of the vein axils (domatia), 2° veins 5-7/side. Inflorescences terminal, 3.5-10 cm long, 2-8 cm broad, open pyramidal with spreading or compact opposite branches, peduncles 0.5-7 cm long, 0.5-1 .5 mm thick, appressed-puberulent, bracts 3-12 mm long and
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linear (or basal bracts larger and leaf-like), flowers usually borne in terminal cymes, pedicels 0-2 mm long. Flowers with hypanthium 2-3 mm long, 1.5-2 mm diam., tur- binate, densely ascending sericeous, calyx tube ca. 1 mm long, calyx lobes 4 and strongly dimorphic, the 3 smaller lobes 1 .5-3.5 mm long and linear, the single larger lobes 4-6 mm long and 1-3 mm broad; corolla white, densely sericeous with ascending whitish hairs externally, tube (6-)ll-15 mm long, 1-2 mm diam., lobes 4, 3-4 mm long, 3-5 mm broad, rounded-truncate at the apex; an- thers included. Fruits 6-8 mm long, 4-6 mm broad, narrowly ovoid to rounded-oblong, minutely puberulent and with prominent longitudinal ribs, the calyx persist- ing until dehiscence.
Plants of evergreen montane rain forest for- mations along the Caribbean escarpment and con- tinental divide, from 900 to 2200 m elevation. Flowering in January-May and July; fruiting in May-September. As presently known, this species is endemic to the area ranging from the Cordillera de Tilaran, Guanacaste, to Tapanti, Cartago.
Rondeletia torresii is recognized by the thin of- ten long-petiolate leaves, calyx with a single en- larged lobe, slender corolla tubes, and montane cloud forest habitat. There seems to be consider- able variation in the form of the inflorescences and the development of the calyx lobes in different collections, but this variation does not appear to be correlated with other attributes.
Rondeletia urophylla Standl. & L. O. Williams, Phytologia 26: 129. 1973. Figure 21.
Shrubs or slender treelets, 4-9 m tall, leafy stems 2-
4 mm thick, densely strigillose with appressed-ascending hairs 0.5-1 mm long, glabrescent in age; stipules 6-10 mm long, to 4 mm broad at the base, subulate with a long-linear tip from a short (2 mm) broadly triangular base. Leaves opposite and often unequal at each node, petioles 4-18 mm long, 1.2-1.8 mm thick, densely ap- pressed-puberulent; leaf blades 10-22 cm long, 3-7 cm broad, elliptic to elliptic-oblong or elliptic-lanceolate, apex tapering gradually and long-acuminate with tip 1 5- 30 mm long, base acute or obtuse and slightly decurrent on petiole, drying thin-chartaceous, dark brown above, glabrescent and often lustrous above, with short (0.3- 0.5 mm) thin appressed hairs on the veins beneath, 2° veins 7-9/side. Inflorescences terminal, solitary, ca. 16 cm long, 2.5-5.5 cm broad, narrow thyrsoid panicles, peduncles 4-6 cm long, 1.1-2.2 mm thick, appressed- puberulent, secondary peduncles of the lateral branches 4-14 mm long, bracts 3-10 mm long, linear, flowers in short distal scorpioid dichasia, pedicels 0-1.5 mm long. Flowers with hypanthium 1.5-2 mm long, ca. 1 mm diam., appressed-puberulent, calyx lobes 4, clearly un- equal with the 2 smaller 1-3 mm long and 0.5-1 mm broad, the larger calyx lobes usually 2 and opposite, ca.
5 mm long and 1.5-2 mm broad, with 3 palmate veins, lanceolate, white, sparsely puberulent; corolla white, gla-
brous externally, tube 7-10 mm long, 0.7-1.3 mm diam., lobes 4, 2-3.5 mm long, 2-3 mm broad, broadly oblong or ovate and rounded distally; style branches linear, stig- mas ca. 2.5 mm long. Fruits ca. 3 mm long, 4-5 mm wide after opening.
Plants of the lowland Pacific rain forest for- mations around the Golfo Dulce area, 30-300 m elevation. Flowering in July-August (Allen 6291 holotype F; EAP), with older fruits in February. Endemic to southern Costa Rica.
Rondeletia urophylla is recognized by the larger long-acuminate leaves, the long narrow inflores- cences, the glabrous corolla tubes, the four unequal calyx lobes usually in two opposite pairs, and the restricted range. There seems to be considerable variation in the development of the calyx lobes. This species is similar in general appearance to R. bertierioides.
Rudgea Salisbury
Shrubs or small trees, stems usually terete, glabrous or sparsely puberulent; stipules interpetiolarly, coria- ceous, sometimes also united intrapetiolar to form a cap over the shoot apex, with cartilaginous appendages along the distal edge or on abaxial surface, the appendages digitate (and thickened) to dentate or aristate, persisting or deciduous. Leaves opposite (rarely 3 or 4/node), short- petiolate or sessile; leaf blades entire and pinnately veined, chartaceous to coriaceous, often with thin vein-like tis- sue along the edge, domatia present or absent. Inflores- cences terminal and solitary (in Central America), pa- niculate to umbellate or capitate (reduced to a single flower in R. monofructus), pedunculate, flowers often in cymose groups. Flowers bisexual, monomorphic or dis- tylous, radially symmetrical, 5-parted (rarely 4- to 8-parted), hypanthium generally ovoid to obconic, calyx tube short or long, calyx lobes 4—6 or absent, usually small; corolla salverform to funnelform or tubular, white, the throat glabrous or puberulent, corolla lobes 5 (rarely 4 or 6-8), valvate in bud; stamens borne in the throat of the tube or on the lower half, anthers dorsifixed, in- cluded or exserted; ovary 2-locular with 1 erect ovule in each locule from the base of the septum. Fruits baccate or dry, rounded and made up of 2 pyrenes (rarely 1), the pyrenes plano-convex, 1 -seeded; seeds ellipsoid, with an incurved ventral (adaxial) surface.
A genus of ca. 1 60 species, ranging from Mexico and the West Indies through tropical South Amer- ica. A great majority of species are South Amer- ican. This genus is characterized by the two-loc- ular ovary with only two ovules, the baccate fruit without prominent longitudinal ribs, the seeds with incurved ventral surface, and the unusual stipular developments. While the stipules with their in- durated teeth may be distinctive, they are often
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301
caducous and may be lacking in some species. The inflorescence branches often have a succulent rath- er jointed appearance and they usually lack bracts. Specimens of this genus may be difficult to differ-
entiate from species of Psychotria and Coussarea, and some may resemble species of Faramea and Raritebe as well.
Key to the Species of Rudgea
la. Hypanthium and calyx 1.5-3 mm long; stipules with the teeth usually borne along the broadly rounded distal margin or hidden within the distal stipule margin; domatia often present on undersides
of leaf blades 2
Ib. Hypanthium and calyx 3.5-9 mm long; stipules rounded distally with the thickened teeth only in the center of the distal edge and with broad rounded distal margins; domatia absent on undersides
of leaf blades 5
2a. Leaf blades 14-25 cm long, 4-12 cm broad, usually narrowly obovate [drying darker greenish]; domatia absent; inflorescences with 3-4 branches at the first node [Caribbean slopes, mostly
at 600-900 m elevation] R. reducticalyx
2b. Leaf blades 4.5-15 cm long, 2-7 cm broad, elliptic to oblanceolate; pit domatia often present;
inflorescences with 2 lateral branches at the first node 3
3a. Leaf blades subsessile, oblanceolate to narrowly obovate-oblong; corolla tubes 12-14 mm long
R. amplexicaulis
3b. Leaf blades rarely subsessile oblanceolate or narrowly obovate-oblong; corolla tubes 3-8 mm
long 4
4a. Leaf blades drying pale grayish green, petioles 0-5 mm long; 50-400(-700) m elevation ....
R. cornifolia
4b. Leaf blades drying dark greenish or yellowish brown, petioles 5-16 mm long; ca. 1500 m
elevation Coussarea chiriquiensis
5a. Flowers usually solitary [corolla lobes ca. 1 1 mm long and with recurved tips 3-4 mm long; leaf
blades usually acute at the base]; Caribbean slope of northern Costa Rica R. monofructus
5b. Rowers in open panicles of (3-)5-15 flowers; plants of central and southern Costa Rica 6
6a. Leaf blades often slightly auriculate at the base [drying chartaceous and abruptly acuminate]; corolla lobes 3-7 mm long, with or without distal appendages to 2 mm high [calyx and hypanthium 4-6
mm long; southern Costa Rica to Panama] R. skutchii
6b. Leaf blades acute to obtuse at the base; corolla lobes ca. 6-8 mm long with erect tips 1.5-6 mm
long 7
7a. Calyx and hypanthium 5-8 mm long; leaf blades to 1 6 cm long and stiffly chartaceous; Caribbean
slopes and lowlands of the Talamanca range R. trifurcata
7b. Calyx and hypanthium 4-5 mm long; leaf blades to 27 cm long and subcoriaceous; evergreen Pacific lowlands . . R. raveniana
Rudgea amplexicaulis Dwyer, Ann. Missouri Bot. Gard. 67: 475. 1980.
Sprawling or clambering shrubs to 2 m long, stems glabrous; stipules to 1.5(-3) mm long, usually entire and cupulate or with a few stiffcolleters within. Leaves sessile or the petioles to 3 mm long; leaf blades 5-16 cm long, 2-5 cm broad, oblanceolate to narrowly obovate-oblong, apex acuminate, base tapering gradually and auriculate or slightly rounded, drying chartaceous, green, 2° veins 7-1 I/side, weakly loop-connected distally. Inflores- cences solitary, terminal, 2-4 cm long, cymose-panicu- late with ca. 3 branches and 7-9 flowers, peduncles to 8-35 mm long, pedicels 0-6 mm long. Flowers with
hypanthium 1-2 mm long (continuous with the pedi- cels), calyx tube 0.5-1 mm long, lobes poorly developed; corolla white, funnelform, tube 11-14 mm long, lobes 5, 3-6 mm long. Fruits ca. 10 mm long and 6 mm diam., ovoid.
Rudgea amplexicaulis is distinguished by its clambering habit, small inflorescences, and rela- tively large corollas. The type (Croat 33133 MO) from 1200 to 1500 m in Chiriqui, Panama, was flowering in March. Costa Rican material ascribed to this species is more likely to be R. cornifolia,
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where occasional individuals can have similarly narrow subsessile leaves.
Rudgea cornifolia (Humb. & Bonpl. ex Roem. & Schult.) Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 432. 1931. Psychotria cornifolia Humb. & Bonpl. ex Roem. & Schult., Syst. Veg. 191. 1819. P.fimbriata Benth., J. Bot. 3: 226. 1841. R.fimbriata (Benth.) Standl. in Standl. & Calde- ron, Lista PI. Salv. 274. 1925. R. ceratopetala J. D. Smith, Bot. Gaz. 35: 3. 1903. Figure 46.
Shrubs or small treelets, 1 .5-6(-10) m tall, leafy stems 1.2-3 mm thick, glabrous, terete, drying grayish to yel- lowish green, the nodes conspicuously thickened; stip- ules 2-7 mm long, 2.5-5 mm broad, oblong or broader than long, with a truncated apical margin with (l-)3-5 short (0.5-1 mm) stiff teeth. Leaves subsessile or peti- olate, petioles 1.5-6 mm long, to 2 mm thick, glabrous; leaf blades 4.5-1 5(-l 8) cm long, 2-7(-9) cm broad, broadly elliptic to narrowly elliptic, elliptic-oblong, el- liptic-obovate or ovate, apex acuminate with tip 7-15 mm long, base obtuse to narrowed and often slightly rounded (subcordate), drying chartaceous, pale grayish or yellowish green, glabrous above and beneath, 2° veins (3-)5-9/side, usually with a very slender veinlet along the margin, often with slit-like or rounded pit domatia along the midvein beneath. Inflorescences solitary or rarely 2, 3-7 cm long, 2-5 cm broad, few-branched open pyramidal, the first node with 2 lateral branches, pe- duncles 1-5 cm long, 0.5-1 mm thick, glabrous, bracts vestigial, distal branches cymose with 3-5 flowers and white in life, flowers sessile or with pedicels to 5 mm long. Flowers glabrous, distylous, hypanthium 0.6-1.3 mm long, oblong to obconic, calyx tube 0.6-1.2(-2) mm long, broadly cupulate to almost tubular, 1.5-2 mm in diameter distally, calyx lobes very short (0-0.5 mm) and broadly triangular or obsolete; corolla, salverform, white, tube (3-)4-5(-7) mm long and 1 mm diam., lobes 3-4.5 mm long, 0.8-1.5 mm broad, oblong. Fruits 6-9 mm long, 5-9 mm diam., oblong-ellipsoid or subglobose, gla- brous, becoming white and fleshy, often with a promi- nent coriaceous calyx tube 0.5-3 mm long and ca. 2 mm diam. at the apex.
Plants of evergreen rain forest and wet forest formations of the Caribbean slope and southern Pacific slope in Costa Rica, from 50 to 800 m elevation. Flowering in February-September (mostly in June-July); fruiting in January-March and July-November. The species ranges from eastern Mexico through Central America to Brazil and Bolivia.
Rudgea cornifolia is recognized by the unusual stipules with little digitate projections on the distal edge, lack of puberulence on all parts, leaves often subsessile and tapering to a slightly auriculate base, small inflorescences with few opposite branches,
small flowers with poorly developed calyx lobes, and smooth rounded fruit (often with a persistent calyx tube). The leaves usually dry grayish green beneath and can differ greatly in form on different plants. Some specimens may resemble material of Coussarea. Breeding biology was studied by Bawa and Beach (1983).
Rudgea monofructus Gomez-Laurito & Dwyer, Novon 1: 50. 1991.
Shrubs or small treelets, 2-4.5 m tall, to 13 cm dbh, leafy branchlets 1.2-3.5 mm thick, glabrous and drying yellowish or grayish, soon becoming terete; stipules 3- 10 mm long, 3-5 mm broad, oblong to broadly obovate, rounded distally with small (ca. 0.5 mm) fimbriate teeth clustered in the center of the distal margin, succulent and glabrous, caducous. Leaves clustered distally or distant, decussate, petioles 3-12 mm long, 0.8-2 mm thick, gla- brous; leaf blades 7-1 5(-18) cm long, 2.5-7(-9) cm broad, elliptic-oblong to elliptic-obovate or obovate, apex abruptly narrowed and acuminate, tip 8-16 mm long, base acute, drying thin-chartaceous to chartaceous, dark grayish green above (distinctly paler or yellowish green beneath), glabrous above and below, 2° veins 4-8/side, edge of the leaf often with vein-like tissue, domatia ab- sent. Inflorescences of solitary (rarely 2-3) terminal flow- ers, peduncles/pedicels 12-32(-40) mm long, 1-1.5 mm thick, glabrous, articulate but lacking bracts along their length and emerging from the terminal stipule pair, ped- icels ca. 2 mm long on lateral flowers when more than 1 flower is present. Flowers glabrous, hypanthium and calyx not differentiated, 8-10 mm long, 4-5 mm diam., rounded at the base and tubular distally (slightly ex- panded distally), coriaceous, calyx lobes 3 or 4, 1.5-4 mm long and ca. 3 mm broad at the base, rounded- obtuse at the apex or irregular; corolla salverform-rotate, white, carnose, tube 5-9 mm long and equal to or slightly longer than the calyx tube, ca. 5 mm diam., lobes 6 (5, 7), apparently held horizontally (to 3 cm diam.), 8-12 mm long with an additional recurved or vertical tip 3- 5 mm long; stamens included, filaments ca. 1 mm long, anthers ca. 3.5 mm long with a small (0.4 mm) append- age. Fruits 12-20 mm long (including calyx), 10-14 mm diam., ovoid-oblong and crowned with the persisting calyx tube 2-6 mm long and 4-6 mm diam., becoming greenish white.
Plants of evergreen rain forest formations on the Caribbean slope, from 600 to 1200 m elevation. Flowering in July-November; fruiting in Febru- ary, June, August, and October-December. The species is only known from the Caribbean slopes of northern Costa Rica (Upala area to Ciudad Que- sada) in Alajuela Province.
Rudgea monofructus is characterized by its usu- ally solitary flowers (rarely with two to three flow- ers from a terminal node), large cylindrical hy-
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panthium/calyx tube, and the long corolla lobes with distal recurved or erect tips. Many similari- ties (especially the flowers and stipules) suggest a close relationship with R. skutchii and R. trifur- cata. The flowers are very unusual in being solitary and having the corolla tube largely hidden within the calyx tube. In addition, the usually six hori- zontal petals have conspicuous erect distal ap- pendages.
Rudgea raveniana W. Burger, sp. nov.
Frutices vel arbores ad 6 m altae, ramulis glabris; stip- ulis 0.5-3 mm longis, integris. Foliae lamina 10-21 cm longa, 3.5-8 cm lata, anguste vel late elliptica, glabra, venis lateralibus 6-11. Inflorescentiae terminales, soli- tariae, 5-7 cm longae, flores 5-15 gerentes. Flores extus glabri; corolla hypocrateriformi, tubo 10-14 mm longo, lobis 5, ca. 6 mm longis. Fructus maturus non visus.
TYPUS— Burger & Liesner 7251 (holotypus CR, isotyp- us F), from Rincon de Osa, Costa Rica.
Shrubs or small trees, 1.5-6 m tall, leafy twigs 1.2-5 mm thick, glabrous, flattened in early stages, becoming terete and grayish; stipules united to form a very short (0.5-3 mm) entire collar or cup (rarely with ca. 8 apical closely crowded teeth 1-2 mm long and these caducous). Leaves with petioles 4-20 mm long, 1.5-3.5 mm thick, glabrous and drying dark; leaf blades (10-) 14-21 (-2 7) cm long, (3.5-)4.5-8(-l 5) cm broad, broadly to narrowly elliptic, elliptic-oblong or slightly obovate, apex short- acuminate or acute (acuminate in the smaller leaves), base acute to obtuse, laminae drying subcoriaceous, dark olive green above (paler grayish green or yellowish green beneath), glabrous above and below, 2° veins 6-1 I/side (largest leaves with 10-11). Inflorescences terminal and solitary, 5-7 cm long, paniculate-cymose, with 1 or 2 pairs of lateral branches and a distal branch (each usually terminating with 3 flowers for a total of 9-1 5), peduncles 14-25 mm long (to 35 mm in fruit), 1.2-1.8 mm thick and glabrous, bracts rudimentary (< 1 mm), pedicels 2- 9 mm long. Flowers glabrous externally, white, hypan- thium ca. 2 mm long and 1.3 mm thick, calyx limb 2- 3 mm long, 3-4 mm broad, calyx lobes 5-7, minute; corolla salverform with recurved lobes, tube 10-14 mm long, ca. 1.7 mm diam. in the center and 3 mm near the mouth, lobes 5, ca. 6 mm long and 1.6 mm broad, with a distal reflexed appendage 1.5 mm long; stamens 5, anthers ca. 5 mm long and 0.4 mm broad, exserted. Fruits ca. 20 mm long and 8 mm diam. (not fully ma- ture?), with a 2-4 mm high persisting calyx tube), ellip- soid, smooth and drying black.
Plants of evergreen forest formations in the Pa- cific lowlands of southern Costa Rica, 50-300 m elevation. Flowering in January (the type), late flowers with fallen corollas in March (Croat & Grayum 59753 CR, MO), immature fruits in Feb- ruary (Burger et al. 12242 CR, F, MO, Hammel et al. 18081 CR, MO), and more mature fruits in May
(Hammel 1521 1 CR, MO). The species is only known from the hills north of Quepos and near Rincon de Osa.
Rudgea raveniana is recognized by its larger stiff glabrous leaves, few-flowered paniculate inflores- cences with distal triads of flowers, petal lobes with distal reflexed appendages, and ellipsoid fruit. The teeth on the stipules appear to be early caducous or undeveloped at many nodes, with the stipular tube entire. This species is related to R. trifurcata and R. skutchii but differs in leaf size and texture and in the flower and inflorescence. The ovary of the flower appears to have two well-developed loc- ules (each with an ovule), but only one seed de- velops in the ellipsoid fruits, which are similar to other species of Rudgea. The fruiting collection from near Quepos (12242) has unusually large leaves whose measurements are given in paren- theses (not included in the Latin description). The species is named in honor of Peter Raven, who made important collections in southern Costa Rica and whose leadership of the Missouri Botanical Garden has supported floristic research in many parts of the world.
Rudgea reducticalyx Dwyer, sp. nov. Figure 46.
Frutices 1.5-3 m alti, ramulis glabris. Folia anguste obovata vel anguste elliptico-oblonga, 1 5-27 cm longa, 5-12 cm longa, 5-12 cm lata, chartacea, glabra; petiolis ad 20 mm longis; stipulis 3-6 mm longis. Inflorescentiae terminales paniculatae vel umbellatae, 5-12 cm longae, 4-7 cm latae, pedunculo ad 8 cm longo. Flores hypanthio glabro 0.5-1 mm longo, dentibus calycinis nullis vel minutis; corolla alba, tubo ad 6 mm longo, lobis 4 vel 5, 2-3 mm longis. Fructus 7-8 mm longus, glaber.
TYPUS— Croat 36330 (holotypus CR, isotypus MO), from near Bijagua, Alajuela, Costa Rica.
Shrubs, 1.5-2.5(-3) m tall, with spreading branches, leafy stems 2-8 mm thick, glabrous and terete; stipules 3-6 mm long, 4-6 mm broad, united to form a short (1- 2 mm) tube, often ovate and rounded distally, thick and drying dark, indurated teeth along the distal edge or hidden within, caducous and leaving a circular scar above the petiole. Leaves subsessile or with petioles 3-8(-20) mm long, 1.5-2.5 mm thick, glabrous; leaf blades (13-)15-20(-27) cm long, (4-)5-9(-12) cm broad, nar- rowly obovate to narrowly elliptic-oblong or oblanceo- late (rarely broadly elliptic), apex gradually or abruptly short-acuminate with tip 5-9 mm long, base gradually narrowed and abruptly rounded or auriculate at the pet- iole, drying chartaceous, olive green above, glabrous on both surfaces, 2° veins 8-12/side, arcuate-ascending dis- tally but not loop-connected, the leaf edge with thin vein- like tissue. Inflorescences 5-12 cm long, 4-7 cm broad, open paniculate or subumbellate with 3—4 branches at the first node, white, peduncles becoming 3-8 cm long in fruit, 1-2.5 mm thick, glabrous or minutely papilla te-
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puberulent, bracts and floral bracts minute (0.5 mm) or reduced to ridges, flowers mostly in distal 3-flowered cymes, pedicels 0-5 mm long. Flowers glabrous, with hypanthium 0.5-1 mm long, ca. 0.7 mm diam., cylin- drical to obconic, calyx limb 0.5-1 mm long, ca. 1 mm diam. distally, calyx lobes absent or small and broadly triangular, essentially glabrous; corolla tubular-slaver- form, white, minutely papillate-puberulent, tube 4-6 mm long, 0.7 mm diam. at base and 1.3 mm distally, lobes 4-5, 2.3-3 mm long. Fruits 7-8 mm long, 6-7 mm diam., globose-oblong, with smooth surfaces and bisulcate at maturity, persisting calyx 0.5 mm long and usually en- tire.
Plants of the very wet forests of the Caribbean slopes of the Meseta Central and Cordillera de Tilaran, (100-)400-800(-1 100) m elevation. Flowering in December-July; fruiting December- February. This species is endemic to northern and central Costa Rica.
Rudgea reducticalyx is recognized by its longer obovate subsessile (or short-petiolate) leaf blades with short-acuminate apices and leaf edges with thin vein-like tissue. The apparently ebracteate in- florescences with three to four branches at the first node and small flowers with subentire calyx tube and very minute puberulence (or the puberulence not apparent) are also distinctive. The stipules do not always display the indurated teeth that char- acterize the genus. These teeth may be hidden within (adaxially) and are caducous with the stip- ules. This species is very similar to Coussarea im- petiolaris, but that species has more elliptic leaves, smaller inflorescences, and much larger lenticel- late single-seeded fruit. Material of this species collected in the 1920s was misidentified as R.fim- briata (Benth.) Standl. by Standley (in herb, us); that name is a synonym of R. cornifolia according to Steyermark (1972). The following collections are placed here: Burger & Antonio 11103, Grayum & Herrera 7819, Hammel et al. 12812, INBio 19, A. Jimenez 1339, Standley & Valeria 45382 & 44593, and Utley & Utley 3908.
Rudgea reducticalyx may be represented by an atypical population or a closely related and un- described species on the Osa Peninsula. The leaves are subsessile and up to 38 x 17 cm, and the rounded fruits are 2 cm long (Burger & Liesner 7242 CR, F, Liesner 3225 MO).
Rudgea skutchii Standl., Publ. Field Mus. Nat. Hist., Hot. Ser. 18: 1374. 1938. Figure 46.
Shrubs or small trees, 1-6 m tall, leafy stems 1-3.5 mm thick, glabrous, at first somewhat flattened but be- coming terete, nodes thickened; stipules (3-)4-8 mm long,
2-4 mm broad, ovate-oblong or obovate, at first united and forming a 4-angled cap over the apex, coriaceous and glabrous, the thickened apical teeth becoming 1 mm long and borne in a small (0.5-1.5 mm) tight group at the center of the rounded apical margin, soon turning brown and deciduous. Leaves subsessile or petiolate (on different plants), petioles 0-5(-10) mm long, ca. 1.5 mm thick, glabrous; leaf blades 6-14(-17) cm long, 2-5.5(-7) cm broad, elliptic to elliptic-oblong or obovate, apex gradually or abruptly narrowed and acuminate, tip 6- 17(-25) mm long, base quite variable (acute to obtuse or more often slightly auriculate at the petiole), drying chartaceous, grayish green to dark brownish, glabrous on both surfaces, 2° veins 5-1 I/side, with vein-like tissue at the leaf edge, pit domatia rarely present. Inflorescences 4-14 cm long, 3-10 cm broad, open-branched panicles with relatively few (5-1 5) flowers, with 2 lateral branches at the first node, peduncles 2-6 cm long, 0.7-1.6 mm thick, glabrous, bracts to 1 mm long or absent, flowers sessile or on pedicels to 7(-12) mm long. Flowers gla- brous, hypanthium difficult to distinguish from the calyx tube, ca. 2 mm long, calyx tube 2-4 mm long, 2-3 mm diam. distally, entire or with 5 short lobes; corolla white, tube 3-7 mm long, 1.3-2 mm diam., lobes 5, 3-7 mm long, with a distal small (0.5-1 mm) thickened area or recurved appendage 1-3 mm long (as in the type, Skutch 2836 us); stamens 5, anthers 2.5-3 mm long. Fruits 6- 8 mm long (not including calyx), 6-8 mm diam., subglo- bose with a persisting tubular calyx 1-2 mm long.
Plants of evergreen rain forest and wet forest formations on both the Caribbean and Pacific slopes, from 50 to 1 300 m elevation (but not below 600 m on the Pacific slope). Flowering in Febru- ary-November; fruiting in July-August and Oc- tober-January. The species ranges from Monte- verde and the Cordillera de Talamanca southward to the Choco area of Colombia.
Rudgea skutchii is recognized by the unusual stipules, lack of pubescence, subsessile leaves, few- flowered inflorescences, prominent calyx tube, and the appendaged corolla lobes. This species is re- lated to R. trifurcata and R. monofructus, but those species have larger flowers and inflorescences with little or no branching. It differs from R. cornifolia in the much more open inflorescences and larger flowers. Material from Bocas del Toro often has simpler inflorescences, resembling those of/?, tri- furcata (see below).
Rudgea trifurcata Gomez-Laurito, Brenesia 33: 139. (1990) 1991. Figure 46.
Small trees, 3-6 m tall, leafy stems 0.7-3 mm thick, glabrous, becoming pale grayish; stipules 3-7 mm long, 4-6 mm broad, ovate to oblong, thick, rounded distally but with a group of thickened teeth at the apex 1-2 mm broad and 1 mm long (rarely absent), deciduous. Leaves with petioles 3-10 mm long, 0.7-1 .8 mm thick, glabrous,
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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sulcate above, drying yellowish or brown; leaf blades 7- 16 cm, 2.5-8(-10) cm broad, elliptic to elliptic-obovate or obovate, apex acuminate to caudate-acuminate, tip ca. 10 mm long, base obtuse to acute, drying stiffly char- taceous, dark green above, glabrous above and below, 2° veins 5-7/side. Inflorescences solitary and terminal, 4.5-10 cm long, 4-5 cm broad, paniculate with 3 major branches from the apex of the peduncle (each usually cymose with 3 flowers), usually with 9 flowers, peduncle 2.5-6 cm long, 0.7-1.7 mm thick, glabrous and drying yellowish green or yellowish brown, bracteoles minute and caducous, pedicels 3-10 mm long. Flowers glabrous externally, hypanthium and calyx difficult to distinguish, 5-8 mm long, 2.5-4 mm diam. distally, calyx lobes 0.5 mm high or poorly developed; corolla white, salverform with rotate lobes, 2 cm diam., tube 7-1 1 mm long and 2 mm diam. distally, lobes 5, 6-8 mm long (not including the 2-5 mm long distal appendages held vertically at 90°); stamens 4, subsessile, anthers 4-4.5 mm long. Fruits unknown.
Plants known only from the Caribbean slopes and lowlands below the Cordillera de Talamanca, 50-1 300 m elevation. Flowering in January (Leon 345 usj) and November (Baker & Burger 66 CR, F). Gomez-Laurito & Vargas 11957 CR is the ho- lotype. This species ranges from Moravia, Carta- go, on the Caribbean slope of the Talamanca range southward to the mountains of Bocas del Toro Province, Panama.
Rudgea trifurcata is distinguished by the few- flowered inflorescences, relatively large flowers with long calyx and short corolla tube, and the long corolla lobes with distal vertical appendages. This species is closely related to both R. skutchii and R. monofructus. The higher-elevation collections from Moravia (Leon 345 usj) and San Ramon
(Loiselle 149 MO) have smaller flowers and may be intermediate with R. skutchii.
Rustia Klotzsch
REFERENCE— D. Simpson, Studies in Neotrop- ical Rubiaceae. I. Rustia. Phytologia 33: 4-8. 1976.
Trees, branchlets glabrous; stipules interpetiolar and free (rarely intrapetiolar), entire or bifid, large and de- ciduous. Leaves opposite, petiolate; leaf blades medium to large in size, entire and pinnately veined, often with punctate oil glands, domatia absent. Inflorescences ter- minal and solitary, long-paniculate with decussate op- posite branching, the lateral branches usually monocha- sial. Flowers with hypanthium turbinate or cupulate, calyx tube short and cupular, truncate or with small calyx teeth; corolla campanulate to funnelform, corolla tube glabrous or puberulent within, lobes 4-5, valvate in bud; stamens borne on the throat of the tube, filaments very short, anthers basifixed, opening by terminal slits or 2 pores beneath the apex, included or exserted; ovary 2-locular, placentation axile with many horizontal ovules in each locule. Fruits capsules, coriaceous, narrowly obovate to subglobose-obovate, dehiscing loculicidally into 2 valves; seeds small, pointed at the ends, angular or with minute marginal wings.
A genus of about 1 2 species, found in Mexico, the West Indies, southern Central America, and tropical South America. The genus is easy to rec- ognize because of the anthers usually opening by terminal pores and the leaves with pellucid dots. Standley's genus Stomandra is a synonym of Rus- tia (D. Lorence, pers. comm.).
Key to the Species of Rustia
la. Rowers pink, corolla tubes 6-9 mm long; larger leaf blades 4-1 1 cm broad; fruit 8-12 mm long;
shrubs 2-5 m tall R. occidentalis
Ib. Flowers white, corolla tubes 1-2 mm long; larger leaf blades 9-19 cm broad; fruit ca. 5 mm long;
trees 5-10 m tall . . .R. costaricencis
Rustia costaricensis (Standl.) Lorence, comb. nov. Stomandra costaricensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 247. 1947. R. panamensis Dwyer, Ann. Missouri Bot. Gard. 67: 483. 1980. Figure 38.
Trees to 1 2 m tall, leafy stems ca. 4 mm thick, terete, glabrous and grayish; stipules not seen, caducous. Leaves opposite, petioles (l-)2-6 cm long, ca. 2.5 mm thick, glabrous, with 2 slightly elevated adaxial ridges, thick- ened at the base; leaf blades 15-45 cm long, 7-19 cm broad, oblong to narrowly oblong-obovate, apex obtuse
to short-acuminate, base obtuse, drying stiffly charta- ceous and brown, glabrous above and below, 2° veins 12-17/side, lower surfaces with small (0.2-0.3 mm) cir- cular projections. Inflorescences 1-3 at the ends of stems, 20-26 cm long, to 20 cm broad, open paniculate with opposite branching from the central rachis, peduncles 2- 4.5 cm long, 2-3.5 mm thick, glabrous, dark brown, distal bracts 1-2 mm long and ovate, bracteoles sub- tending the flowers 1-1.5 mm long, flowers alternate on the distal branches or in cymes, pedicels 0-2 mm long. Flowers glabrous externally, hypanthium 1-1.5 mm long, ca. 1 mm diam., turbinate, calyx limb 0.5 mm long, calyx lobes poorly developed (distal margin undulate); corolla white, tube 1-2 mm long, ca. 1.5 mm diam., lobes 4-6,
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1.5-2 mm long, narrowly ovate to triangular, thick; sta- mens 5, filaments ca. 1 mm long, anthers 1 .5-2 mm long, partly exserted; style to 4 mm long. Fruits to 5 mm long, broadly oblong to subglobose, subsessile; seeds ca. 0.8 mm long.
Plants of evergreen forest formations of the Pa- cific slope at 200-800 m elevation. Flowering in February-March and August. The species is only known from southeastern Costa Rica and western Panama.
Rustia costaricemis is recognized by its long- petiolate narrow or oblong leaves with pellucid dots, glabrous parts, large inflorescences, small flowers, anthers with distal pores, and small cap- sular fruit. This species has been collected from the General Valley (Skutch 4777 F the holotype) and in the hills northeast of Quepos (Burger et al. 12314 CR, F).
Plants of evergreen rain forest formations, from 5 to 300 m elevation. Flowers and fruits have been collected in February- April. This species seems to have a disjunct distribution of small populations. It is found near Bluefields, Nicaragua, near Moin (Limon), in Bocas del Toro, Panama (all along the Caribbean coast), and Cocos Island. This species is also reported from the lowlands of Choco, Co- lombia (Dwyer, 1980).
Rustia occidental^ is recognized by its narrow and long-acuminate leaf blades with pellucid dots, glabrous parts, narrow inflorescences, short pink corollas, thick anthers opening by pores, and the capsule breaking into two valves. The species ap- pears to be restricted to wet or swampy sites at very low elevations.
Sabicea Aublet
Rustia occidentalis (Benth.) Hemsl., Biol. centr. amer. Bot. 2: 14. 1881. Exostemma occidentale Benth., Bot. voy. Sulph. 104. 1844. Figure 38.
Much-branched shrubs or small trees, 2-6(-12) m tall, leafy stems 3-6 mm thick, glabrous and smooth, terete, often pale grayish; stipules 16-24 mm long, 2-4 mm broad, tapering gradually to the acute apex, glabrous and drying brown, caducous. Leaves opposite or suboppo- site, petioles 13-35 mm long, 1.2-2.5 mm thick, gla- brous, usually thickened at the base; leaf blades 13-33 cm long, 4-1 1 cm broad, oblanceolate to narrowly ob- long-oblanceolate or narrowly obovate, apex tapering gradually and long-acuminate, tip to 35 mm long and often curved, base tapering gradually, drying stiffly char- taceous and brownish, glabrous above and below, with small (0.1-0.3 mm) rounded scale-like projections on upper and lower leaf surfaces, 2° veins 9-12/side. Inflo- rescences 4-15 cm long, to 8 cm broad in fruit, paniculate or racemiform, peduncles 5-30 mm long, 1.2-2.8 cm thick, glabrous, lateral branches to 3.5 cm long, with 1- 5 flowers, bracts 1-2 mm long, triangular, pedicels 3-12 mm long. Flowers glabrous externally, hypanthium 3-5 mm long, narrowly turbinate, calyx tube ca. 1 mm long, calyx lobes not well developed (to 0.3 mm high); corolla pink or violet, tube 6-9 mm long, 2.5-3.5 mm diam., constricted above the base, lobes 4 or 5, 5-6 mm long and 3 mm broad near the base; stamens 5, ca. 6 mm long and 1.8 mm broad. Fruits 8-12 mm long, 6-8 mm diam., broadly obovoid to subglobose, drying very dark brown, truncated at the apex, the persisting calyx ca. 0.5 mm high, 3.5-5 mm diam., open valves with a con- spicuous median septum.
Vines, shrubs with clambering branches, or erect shrubs, stems usually pubescent, terete; stipules interpetiolar, erect or recurved, ovate to Ungulate, persisting. Leaves opposite or verticillate, petiolate, entire, venation pin- nate, domatia absent. Inflorescences axillary, sessile or pedunculate, corymbose to glomerulate or capitate, with bracts and bracteoles, the flowers often in distal cymes, pedicellate or sessile. Flowers bisexual, monomorphic or distylous, radially symmetrical, small or medium, hy- panthium turbinate to urceolate, calyx lobes 3-6, per- sisting; corolla funnelform or salverform, tube pubescent at the mouth, lobes 4-6, valvate in bud; stamens 4-6, anthers dorsifixed, included or partly exserted; ovary (2-)3-5-locular, ovules numerous and horizontal on ax- ile placentas, style with 3-5 stigmatic branches. Fruits baccate, with persisting sepal lobes; seeds small, ovoid or angulate, reticulate to foveolate.
A genus of about 130 species in tropical Amer- ica, Africa, and Madagascar. Our species of this genus are recognized by the climbing habit (or with clambering branches), prominent pubescence on most parts, small axillary inflorescences, small flowers, and baccate puberulent fruit with persist- ing calyx lobes and many small seeds. Our two species are very similar in appearance; some col- lections may be difficult to place with certainty and may represent hybrids. Alternatively, it is pos- sible that Costa Rican material does not represent two distinct species.
Key to the Species of Sabicea
la. Inflorescences to 4 cm broad, often on short peduncles, pedicels to 5 mm long; corolla tube 6-8 mm long; leaf blades obtuse to acute at the base but not usually decurrent on the petiole; hairs to 1 mm long and appressed 5". panamensis
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Ib. Inflorescences to 2 cm broad, sessile and the flowers sessile; corolla tube 4-5 mm long; leaf blades usually decurrent on the petioles; hairs to 2 mm long, spreading or appressed S. villosa
Sabicea panamensis Wernham, Monogr. Sabicea 30. 1914. S. costaricensis Wernham, Monogr. Sabicea 31. 1914. Figure 35.
Vines or shrubs with clambering branches, to 5 m high, leafy stems 1-4 mm thick, with appressed -ascending straight, yellowish hairs 0.7-1 .4 mm long, terete; stipules 5-10 mm long, 4-5 mm broad, ovate with an acute apex, strigose externally, glabrous and dark brown within, per- sisting and often recurved. Leaves opposite, petioles 5- 1 5(-20) mm long, 0.7-1 .6 mm thick, strigose; leaf blades 5-16 cm long, 2-7 cm broad, elliptic to elliptic-oblong or lanceolate, apex acuminate (acute), base acute to ob- tuse, drying chartaceous, often dark brown, with thin appressed-ascending hairs to 1 mm long on the upper surface, more densely strigose on the veins beneath, and with shorter hairs between the hairs beneath, 2° veins 7- 11 /side. Inflorescences 1.5-3 cm long, to ca. 2 cm broad, corymbose, peduncles 0-7(-15) mm long, ca. 1 mm thick, with ascending strigose hairs, basal bracts broadly involucrate but tearing (stipule-like) and to 8 mm long, distal bracts ca. 3 mm long and narrow, ped- icels 2-7 mm long. Flowers monomorphic, with slender appressed-ascending hairs ca. 1 mm long externally, hy- panthium and calyx tube ca. 2 mm long and 1.3 mm diam., calyx lobes 5, 2.3-3(-4) mm long, ca. 1.2 mm broad, glabrous on the inner surface; corolla white, se- riceous, tube 6-8 mm long, ca. 1 mm diam. near the base and 2 mm diam. at the apex, lobes 5, 2-4 mm long, narrowly triangular; stamens 5, anthers ca. 2 mm long. Fruits 5-9 mm diam., globose, appressed-strigulose, pink to red then purple-black at maturity; seeds ca. 0.5 mm long.
Plants of evergreen forest formations through- out Costa Rica, from 20 to 1200 m elevation. Flowering throughout the year; probably fruiting throughout the year. The species ranges from Be- lize to Colombia.
Sabicea panamensis is recognized by the vining habit, appressed pubescence, axillary inflores- cences with evident peduncle or prominent ped- icels, small white flowers, and fleshy red fruit. This species can become a high-canopy liana (Stevens 24620 MO). The differences between this species and its local congener are minor; it would be in- teresting to know if there are naturally occurring intermediates.
Field Columb. Mus., Bot. Ser. 7: 52. 1930. Fig- ure 35.
Vines or shrubby vines to 4 m high, leafy stems 0.7- 3 mm thick, with thin spreading hairs 1-2 mm long (in variety villosa) or with appressed hairs ca. 1 .2 mm long (in variety adpressa), terete; stipules 3-8 mm long, 4-8 mm broad, broadly ovate, glabrous on the inner side, becoming reflexed, persisting. Leaves opposite, petioles 6-20 mm long, 0.5-1.2 mm thick, densely spreading- villose or appressed-strigose; leaf blades (4-)5-14 cm long, (2-)2.5-6 cm broad, elliptic to ovate-elliptic, ellip- tic-rhomboid, or elliptic-obovate, apex bluntly acute to short-acuminate, base obtuse to acute and slightly de- current on the petiole, drying thin-chartaceous, dark brown or dark green above, smooth or slightly scabrous to the touch, with thin spreading hairs ca. 1.5 mm long (in variety villosa) or with appressed hairs 0.3-1.2 mm long (in variety adpressa) above, more densely puberu- lent below (especially on the secondary veins), 2° veins 7-1 1(-1 3)/side. Inflorescences 5-1 5 mm long, present at almost all distal internodes, capitate, peduncles not ap- parent among the closely crowded stipules, bracts and sessile flowers. Flowers monomorphic, hypanthium and calyx tube covered with thin straight ascending yellowish white hairs, calyx lobes 5, 2.5—4 mm long, 1 .2 mm broad, usually glabrous within; corolla white, tube 4-6 mm long and 1 mm diam., with stiff ascending hairs, lobes 5, 1- 3 mm long. Fruits sessile, 5-10 mm diam., globose to oblong, with thin yellowish hairs and becoming red or pink to lavender, calyx lobes persisting and usually re- flexed; seeds ca. 0.3 mm long.
Plants of evergreen forest formations, from near sea level to 600 m elevation. Probably flowering and fruiting throughout the year. The species is found primarily along the Caribbean coast in Cen- tral America, but there are a few collections from Nicoya to the Golfo Dulce region along the Pacific coast. The species ranges from southern Mexico and the West Indies to Brazil and Peru.
Sabicea villosa is recognized by the climbing habit, very small compact axillary inflorescences, small white flowers, and fleshy reddish or pink fruit. This species is not as common in Costa Rica as S. panamensis. Variation within this species can make separation from S. panamensis difficult; compare material of that species.
Sabicea villosa Willd. ex Roem. & Schult., Syst. Veg. 5: 265. 1819. 5. hirsuta H.B.K., Nov. gen. sp. 3: 417 (quarto). 1820. S. hirsuta var. ad- pressa Wernham, Monogr. Sabicea 55. 1914. S. villosa var. adpressa (Wernham) Standl., Publ.
Schradera Vahl
Epiphytic shrubs, or lianas with rooting branches, stems glabrous; stipules interpetiolar and intrapetiolar, united and forming a terminal cap over the nonflowering shoot apices, caducous. Leaves opposite, petiolate; leaf blades
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entire and pinnately veined, usually coriaceous, domatia absent. Inflorescences terminal and solitary or 3, pedun- culate (rarely sessile), capitate and subtended by a broad involucre or with each flower with a small involucre and in an umbellate arrangement, the flowers sessile. Flowers bisexual and radially symmetrical, hypanthium turbi- nate, calyx tube short-cupulate, truncated and without lobes or with small teeth; corolla salverform to tubular, white and thick, villous within the throat, lobes 5-8 (- 1 0), narrow, valvate in bud, spreading or reflexed at anthesis; stamens 5-8(-10), filaments very short and adnate to the tube, anthers dorsifixed, linear, included or exserted; ovary 2-4-locular, with many ovules in each locule on axile
placentas. Fruits closely clustered or united into a syn- carp, fleshy and baccate; seeds numerous, small, hori- zontal, compressed, suborbicular.
A genus of about 40 species, mostly South American and with a few in the West Indies; two species are found in Costa Rica and adjacent Pan- ama. The epiphytic habit, pedunculate capitula with a small involucre, and semisucculent flowers that dry black make this a very distinctive genus. Compare Appunia.
Key to the Species of Schradera
la. Leaf blades rounded at the apex; capitula always solitary; corolla tubes to 8 mm long; Costa Rica 5". costaricensis
Ib. Leaf blades acuminate at the apex; capitula solitary or 3; corolla tubes to 30 mm long; western Panama . . S. blumii
Schradera blumii Dwyer and Hayden, Phytologia 15:59. 1967.
lyx tubes without calyx lobes are distinctive fea- tures of this species.
Scandent or epiphytic shrubs, leafy stems 2-5 mm thick, glabrous, at first quadrangular; stipules not seen, united at the base and leaving a conspicuous scar be- tween the leaf bases and a narrower scar above the pet- ioles. Leaves opposite and in a single plane, petioles 5- 14 mm long, 0.6-2 mm broad, glabrous, with lateral or adaxial margins continuous with the lamina margins; leaf blades 4-1 1(- 15) cm long, 2-5 cm broad, oblong to elliptic-oblong or ovate-oblong, apex acuminate with tip 8-12 mm long, base acute to obtuse or rounded, drying stiffly chartaceous to subcoriaceous, very dark brown, glabrous above and below, 2° veins 1 0-1 2/side and weakly loop-connected near the margin (a thinner 2° vein usually present and parallel between two major 2° veins). Inflo- rescences solitary or 3 at the end of a stem, 4-7 cm long, 2-4 cm broad, peduncles 12-25 mm long, 1-3.3 mm thick, glabrous and drying black, involucre ca. 1 cm long and to 2.5 cm broad, glabrous and drying black, flowers sessile or subsessile. Flowers glabrous externally and dry- ing black, hypanthium 4-6 mm long, calyx tube 4-6 mm long, entire distally (calyx teeth absent); corolla not seen at anthesis, white, tube to 30 mm long, slender, lobes 4, ca. 30 mm long, thick, villose within near the base; sta- mens 5, filaments ca. 1 mm long, anthers ca. 5 mm long. Fruits united near the base and borne tightly together (not truly connate?) on the involucrate apex of the pe- duncle, body of the fruit subglobose and ca. 8 mm diam. (dried), the persisting calyx forming a distal tube 6-8 mm long and ca. 4 mm diam.
Schradera blumii occurs in cloud forest on the Pacific slope of Panama in Code and Chiriqui provinces at about 500-1500 m elevation. The epiphytic/climbing habit, glabrous parts, acumi- nate leaves, involucrate heads, and prominent ca-
Schradera costaricensis Dwyer, sp. nov. Figure 1 9.
Frutices epiphytici. Folia oblonga vel anguste oblonga, coriacea, glabra, 4-8 cm longa, 1 .4-3 cm lata, venis la- teralibus ca. 10; petiolis ad 1.6 cm longis; stipulis ter- minalibus ad 1.8 cm longis. Inflorescentiae terminales solitariae capitatae involucratae, pedunculo 2.5-4 cm longo. Flores tubis calycinis 3-4 mm longis; corolla alba, tubo ad 8 mm longo, 4 mm lato, lobis ad 8 mm longis. Fructus non visus.
TYPUS— G. Herrera 2004 (holotypus CR, isotypi F, MO), from Bijagua, Alajuela, Costa Rica.
Epiphytic shrubs ca. 1-2 m tall, leafy stems 2-5 mm thick, glabrous, with 4 prominent ridges (apparently semisucculent and becoming distorted on drying), pale grayish; stipules 7-10 mm long and 3-8 mm broad (to 1 8 mm long beneath the inflorescences), oblong, rounded at the apex, glabrous and drying black, caducous. Leaves loosely clustered at the ends of branches, petioles 6-16 mm long, ca. 1.3 mm thick, glabrous and drying black; leaf blades 4-8 cm long, 1 .4-3 cm broad, oblong to nar- rowly oblong, apex rounded, base acute to obtuse and slightly decurrent on petiole, drying stiffly chartaceous or subcoriaceous, gray to blackish above, the margin often revolute when dry, glabrous above and below, 2° veins 8-1 2/side and loop-connected near the margin (but difficult to see). Inflorescences terminal and solitary, 4- 7 cm long, drying black, the capitulum 1 5-25 mm diam. (not including the corollas), hemispherical or subglobose, peduncles 25-40 mm long, 2-3.5 mm thick, glabrous, drying black, involucre ca. 1 5 mm broad (4-6 mm from edge to peduncle), flowers sessile or subsessile and closely compressed, with hairs below the flower bases. Flowers
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drying black, glabrous externally, hypanthium ca. 3 mm diam., calyx tube 3-4 mm long, 3-4 mm diam., entire or subentire; corolla white, tube 3-8 mm long, 2-3 mm diam., lobes 5, 4-10 mm long, 1-2 mm broad, recurved; anthers 3.5 mm long, yellow. Fruits apparently united basally in a syncarpous capitulum with the distal part of the individual fruits separate, calyx tube persisting and succulent.
Plants of evergreen cloud forest formations on the Caribbean slope, at 7 00-9 5 0(- 11 00?) m ele- vation. Flowering in January-March, May, July, and October. This species has been collected near Bijagua (Alajuela), at the Monteverde reserve, and the Reserva Forestal de San Ramon. A collection has also been made at Alto Uren near the border with Panama.
Schradera costaricensis is recognized by its epi- phytic habit, small stiff oblong leaves rounded at the apex, solitary capitulum with flowers subtend- ed by a small circular involucre, and the small flowers with long calyx tube. The restricted alti- tudinal range is also distinctive. The following col- lections are placed here: Burger et al. 12139 & 12465, Gomez-Laurito 11147, Haber & Bella 8077 & 8192, and Herrera 452, 2004 & 3325.
Serissa Commerson ex Jussieu
Shrubs, glabrous or puberulent, tissues giving off a fetid odor when crushed; stipules united to form a broad sheath between the petiole bases and with stiff distal awns, persisting. Leaves opposite, small, subsessile, without domatia. Inflorescences axillary or terminal, of solitary or few fascicled flowers. Flowers bisexual and radially symmetrical, small, hypanthium obconic, calyx lobes 4-6; corolla funnelform, white, tube and throat pilose within, lobes 4-6; stamens 4-6; ovary 2-locular, with 1 ovule in each locule, stigmas 2. Fruits subglobose drupes, usually 2-seeded.
A genus of one (or three) species of southeast and east Asia. A cultivar is widely planted.
Serissa japonica (Thunb.) Thunb., Nov. gen. pi. 9: 132. 1798. Lycium japonicum Thunb., Nov. Act. Roy. Soc. Sci. Upsal. 3: 207. 1780. Lycium foetidum L.f., Suppl. PI. 150. 1781.
Shrubs to ca. 0.8 m tall, leafy stems 0.4-1 .7 mm thick, with short (0. 1-0.3 mm) hairs in 2 opposite longitudinal lines, grayish and glabrescent in age; stipules with a sheath 0.5-1 mm long and 3 awns (6/node) to 3 mm long. Leaves opposite or pseudoverticillate by reduction of internodes, petioles 0-2 mm long, poorly differentiated; leaf blades 4-10(-20) mm long, 1.5^(-12) mm broad,
elliptic to lanceolate or obovate, apex usually acute, base cuneate and decurrent on petiole, stiff, drying grayish green and with margins enrolled, 2° veins 2-3/side. In- florescences of solitary or few flowers mostly terminal on short-shoots, subsessile and subtended by the distal leaves. Flowers glabrous externally, calyx lobes ca. 1 mm long, triangular; corolla pink in bud but becoming white, tube 4-5 mm long, lobes 2—4 mm long. Fruits 1-2 mm diam.
Ornamental small shrubs sometimes used as hedges. Some cultivars have double flowers and variegated leaves. They have been planted in the Meseta Central. Leafy stems resemble those of Diodia polymorpha or Arcytophyllum lavarum (see fig. 1).
Sherardia Linnaeus
Annual herbs, with straight stiff trichomes; stipules most apparent (transformed to become leaf-like). Leaves 4-6/node and subsessile or up to 10 and united at the base to form an involucre at the ends of flowering shoots, domatia absent. Inflorescences of subsessile flowers in terminal involucrate capitula. Flowers bisexual and ra- dially symmetrical, calyx with 4-6 lobes; corolla fun- nelform, pink to blue, lobes 4-6, valvate in bud; stamens 4-6; ovary with 2 locules and 1 ovule in each locule, style with 2 unequal branches, stigmas subcapitate. Fruits splitting into 2 dry 1 -seeded mericarps.
A genus with only a single Eurasian species, now found as a weed in many parts of the world.
Sherardia arvensis L., Sp. PI. 102. 1753.
Procumbent herbs, leafy stems 0.3-0.9 mm thick (dried), with 4 longitudinal ridges, hairs erect and whit- ish, 0.4-0.7 mm long; stipules leaf-like. Leaves 4-6(-10)/ node, subsessile or the terminal involucral leaves united for 1-2 mm; leaf blades 4-9(-17) mm long, 0.8-3(-5) mm broad, lanceolate to elliptic or ovate, apex acute, pale grayish green when dried and with revolute margins, with scabrid hairs along the margins and scattered thin straight hairs ca. 0.5 mm long. Inflorescences 1-2 cm broad, usually on the ends of short-shoots, with 4-10 flowers. Flowers with corolla 4-5 mm long, lilac to pink or blue. Fruits 2-5 mm long (including the sepal lobes), splitting into 2 mericarps 1.2-1.5 mm diam., with min- ute stiff hairs.
Sherardia arvensis is a small weed resembling Galium and some Spermacoce. The stiff whorled leaves, lack of apparent stipules, and involucrate inflorescences are distinctive. It has only been col- lected near the cities of Turrialba and San Jose in Costa Rica.
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Simira Aublet
Trees or shrubs, the wood often turning red or rose- purple after being cut (the color then fading with expo- sure to light); stipules interpetiolar, often large, glandular at the base within, caducous or persisting. Leaves op- posite, petiolate; leaf blades entire or slightly pinnately lobed distally, pinnately veined, usually large, a few spe- cies with domatia. Inflorescences terminal or axillary to distal leaves, paniculate with decussate branching or con- gested and capitate, pedunculate, usually with many flowers, the flowers sessile or pedicellate, bibracteolate. Flowers bisexual and monomorphic, radially symmet- rical, small to medium in size, fragrant, calyx tube cu- pular or campanulate, 4-6-dentate or entire; corolla cam- panulate to funnelform, white to yellowish or greenish white, tube with hairs at the stamen bases, lobes 4-5(-6), imbricate or open in bud; stamens 4-6, filaments borne on the lower half of the tube, anthers dorsifixed in the middle, sagittate at the base, conspicuously exserted; ovary 2-locular, ovules many in each locule, horizontally bi- seriate on axile placentas. Fruits capsules, usually glo- bose and woody, 2-locular and 2-valvate with loculicidal dehiscence; seeds large, numerous, lunate to semioblong, with a membranous marginal wing.
A genus of about 35 species, ranging from Mex- ico and Central America into tropical South America. This genus is now circumscribed to in- clude the genus Sickingia. The small flowers, sta- mens with hairs along the base of the filaments, locules with many seeds, capsular or rounded fruits with flat horizontal seeds, and reddish wood (in some species) characterize this genus.
Simira maxonii (Standl.) Steyerm., Mem. New York Hot. Gard. 23: 306. 1972. Genipa maxonii Standl., J. Wash. Acad. Sci. 8: 642. 1918. Sic- kingia maxonii (Standl.) Standl., Trop. Woods 14: 30. 1928. Figure 38.
Trees, to 20 m tall, trunks to 30 cm dbh, cut wood often remaining pinkish, stems glabrous or puberulent at first but soon glabrescent, quadrangular but soon te- rete; stipules 2.5-4 cm long flattened and 6 mm broad, glabrous or minutely appressed-puberulent at the base. Leaves large, subsessile with petioles 3-9(-18) mm long, 3-6 mm thick, glabrous; leaf blades 22-48(-60) cm long, 1 0-26(-40) cm broad, broadly elliptic to elliptic-obovate or rhombic, sometimes lobed distally along the edge (with lobes 3-10 mm long), apex acuminate with a narrowed tip 8-20 mm long, base gradually narrowed and slightly rounded (subauriculate) at petiole, drying stiffly char- taceous, dark green or brown above, slightly paler or reddish beneath, glabrous above, glabrous or with short (0.1-0.2 mm) stiff hairs on the veins beneath, 2° veins 12-18/side, 3° veins subparallel and perpendicular to 2° veins. Inflorescences 6-15(-22) cm long, to 14(-18) cm broad, a compact panicle with many closely spaced
branches, peduncles 0-3(-5) cm long, 5-6 mm thick, minutely puberulent or glabrous, flowers sessile or on short (1-3 mm) pedicels. Flowers with hypanthium 3-5 mm long, obconic, sparsely and minutely puberulent or glabrous, calyx tube 1-2 mm long, broadly cupulate, calyx lobes 5, 0.5-1 mm long and 2 mm broad, minutely ciliolate along the edge; corolla broadly funnelform or campanulate, greenish yellow to cream or orange, tube 4-5 mm long, ca. 4 mm diam. distally, lobes 5, 2-3 mm long; stamens 5, anthers 3-4 mm long, purplish, exserted. Fruits hard, 5-9 cm long, 3-7 cm diam., globose to rounded-oblong, smooth and grayish externally, pericarp 4-5 mm thick; seeds 12-25 mm long and 2-3 mm thick.
Trees of evergreen rain forest formations along the Caribbean lowlands and on the Osa Peninsula, from near sea level to 300 m elevation. Flowering in January-February, June-July, and September; fruiting in February-March, June-August, and November. The species ranges from southernmost Nicaragua to central Panama.
Simira maxonii is recognized by the large op- posite leaves on very short petioles, the compact densely flowered inflorescences, the hard round fruit with large flat horizontal seeds, and the red- dish coloring of wood, anthers, and undersides of young leaves (when present). This very distinctive species is often found in poorly drained sites and along stream edges. The large leaves of some col- lections have small distal lobes, a rarity in Rubi- aceae. The fruits are reported to require 2 years to mature. The distal stems sometimes harbor ants (Frankie 19 Ic from La Selva). Common names are "inkwood," guaiatil, guayatil Colorado, jagua Colorado, and jagua de montana.
Sipanea Aublet
Herbs, erect to clambering or decumbent, usually pu- berulent; stipules interpetiolar, small and inconspicuous to well developed, broadly triangular, acute or with a long narrow awn. Leaves opposite, sessile, subsessile or petiolate; leaf blades entire and pinnately veined, doma- tia absent. Inflorescences terminal or axillary, of 1-3 flowers or branched and paniculate to corymbose or cy- mose, branches often monochasial, bracts narrow, brac- teoles absent, flowers sessile to short-pedicellate. Flowers bisexual and radially symmetrical, calyx lobes 4-5, linear to lanceolate; corolla salverform to funnelform, pink to rose or white, tube glabrous to villous at the mouth, lobes 5, convolute in bud; stamens 5, inserted in the middle or lower part (rarely near the apex) of the tube, anthers dorsifixed and linear; ovary 2-locular, with many ovules on axile placentas. Fruits thin-walled capsules, 2-locular and dehiscing loculicidally; seeds numerous, small, re- ticulate.
A South American genus of about 17 species; a
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single species reaches Costa Rica and Panama. The delicate herbaceous habit, narrow-tubular flowers with corolla lobes contorted in bud, many-seeded capsules, and nonsagittate stamens characterize this genus. The following species has recently been col- lected in Costa Rica. Compare Oldenlandia.
Sipanea biflora (L.f.) Cham. & Schlechtend., Lin- naea 4: 168. 1829. Virecta biflora L.f., Suppl. 1 34. 1 78 1 . Rondeletia biflora (L.f.) Rottb., Descr. rar. pi. surin. 8, pi. 2. 1 798. Manettia hydrophila Dwyer, Ann. Missouri Bot. Card. 67: 280. 1980.
Low creeping or erect herbs to 20(-50) cm tall, leafy stems 0.3-1.7 mm thick, somewhat succulent in life, minutely puberulent with thin curved hairs 0. 1-0.3 mm long; stipules 0.3-1 mm long, bifid or entire distally, minutely ciliate. Leaves evenly spaced and distant along the stems, petioles 4-18 mm long, ca. 0.5 mm broad, minutely appressed-puberulent; leaf blades 12-42(-55) mm long, 6-22(-25) mm broad, ovate, apex obtuse to subacute, base obtuse to subtruncate and acute-decurrent on petiole, drying membranaceous, dark above, glabrous or with sparse short (ca. 0.3 mm) appressed -ascending hairs on the upper surface, usually appressed-puberulent on the veins beneath, 2° veins 3-5/side, ascending. In- florescences usually terminal and spicate, to 6 cm long with 2-5 alternate subsessile flowers (or paniculate with lateral branches bearing 1-2 flowers), peduncles to 15 mm long, ca. 0.4 mm thick, glabrous, pedicels of the distal flowers 5-15 mm long, bracts 0.5-1.5 mm long, linear. Flowers with hypanthium and calyx tube 1-2 mm long, covered with straight stiff ascending hairs, calyx lobes 5, l-3(-4.5) mm long, linear to narrowly triangular, sparsely minutely puberulent; corolla pink or pale purple (white around the throat), glabrous or with a few small hairs externally, tube 5-14 mm long, ca. 1 mm diam., lobes 5, 4-8 mm long, 2-5 mm broad, obovate-oblong; anthers 2.5 mm long, included. Fruits globose capsules, 3-4 mm long (not including the persisting calyx lobes) and 4 mm diam., bisulcate and splitting from the top, with stiff appressed ascending hairs 0.2-0.4 mm long; seeds ca. 0.5 mm thick.
Plants of poorly drained moist sites in the ev- ergreen Caribbean coastal plain, 2-20 m elevation. Flowering and fruiting in March-April and Sep- tember-November. This species ranges from cen- tral Costa Rica southward to Venezuela and north- ern Brazil.
Sipanea biflora is recognized by the semiaquatic or poorly drained lowland habitat, the small her- baceous habit, slender stems with minute curved hairs, thin ovate leaves, few-flowered inflores- cences, hispidulous ovaries, and subglobose cap- sules with persisting linear calyx lobes. The above
description is based, in part, on Steyermark (1974) and Venezuelan material. Compare Pentodonpen- tandrus.
Sommera Schlechtendal
REFERENCE— L. O. Williams, Sommera (Rubi- aceae) in North America. Phytologia 26: 1 2 1-1 26. 1973.
Shrubs or small trees, branchlets usually terete and pubescent; stipules interpetiolar, large, triangular, ca- ducous. Leaves opposite, petiolate; leaf blades usually large, entire and pinnately veined, the minor venation (between the 3° and 4° veins) conspicuously parallel (lin- eolate), domatia present or absent. Inflorescences soli- tary in the distal leaf axils (2/node), corymbose to race- mose, pedunculate, the flowers usually small and often in distal cymes, bracts and bracteoles present. Flowers usually small, hypanthium turbinate, calyx lobes 5, equal or unequal, usually broad and conspicuous, persisting; corolla funnelform to subcampanulate, sericeous exter- nally, somewhat fleshy, corolla tube villous on the throat within, lobes 5, valvate in bud; stamens 5, filaments short and barbate, borne at the throat of the tube, anthers dorsifixed, oblong, obtuse at each end; ovary 2-locular, ovules many on placentas borne on the septum, style short with 2 long style branches, glabrous or hirsute. Fruits baccate, globose to ovoid, 2-locular; seeds nu- merous and minute, subglobose, testa thin and foveolate.
A genus of about 1 5 species, ranging from Mex- ico and Central America into Andean South America. The genus is recognized by the pu- bescence on all parts, larger leaves with unusual parallel (lineolate) minor venation, small flow- ers with large calyx lobes, and fleshy two-locular fruit with many small seeds.
Sommera donnell-smithii Standl., Contr. U.S. Natl. Herb. 17: 436. 1914. S. mesochroa Standl., loc. cit. 439. 1914. 5. dunlapii L. O. Williams, Phy- tologia 26: 122. 1973. S. rivularis L. O. Wil- liams, loc. cit. 125. 1973. Figure 35.
Shrubs or small trees, 1.5-8(-15) m tall, leafy stems 2-7 mm thick, young stems densely strigulose with yel- lowish ascending hairs 0.3-0.9 mm long, becoming terete and dark brown; stipules 14-35 mm long and 3.5-8 mm broad, lanceolate, glabrous or puberulent on the midrib and base abaxially, glabrous on the inside, drying reddish brown, deciduous. Leaves with petioles 1 2—44 mm long, 1-2.4 mm thick, with thin ascending hairs 0.3-0.9 mm long; leaf blades 10-28(-32) cm long, 5-1 2(-14) cm broad,
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FIELDIANA: BOTANY
obovate to elliptic-obovate, oblong or oblong-obovate, apex acuminate with tip 5-15 mm long, base tapering gradually and acute or obtuse, slightly decurrent on pet- iole, drying stiffly chartaceous, brown or yellowish brown, with small (0.5 mm) scattered hairs on the upper surface, appressed-puberlent on the veins beneath with yellowish hairs 0.3-1 .2 mm long, 2° veins 1 1-1 5/side, tufts of hairs or depressions (domatia) often present in the vein axils beneath, smallest veins parallel. Inflorescences 3-7 cm long, to 5 cm broad, paniculate with opposite branching or with 3 major branches, peduncles 1-3 cm long, ca. 1 mm thick and densely puberulent, flowers in corymbs, cymules or racemes, bracts to 10 mm long, linear-lan- ceolate, pedicels 0-4 mm long. Flowers 5-parted, hy- panthium ca. 2 mm long, with dense appressed ascending yellowish hairs, calyx lobes unequal, (l-)2-4(-5) mm long, 1-3 mm broad, usually glabrous on the surfaces and drying pale brown, with parallel or reticulate ve- nation; corolla white, funnelform, tube 3-4 mm long, ca. 2 mm diam., densely appressed-puberulent, lobes 2-3 mm long; anthers 0.8 mm long. Fruits becoming 6-10 mm diam., globose to oblong, becoming dark red or maroon, with ascending hairs and persisting sepal lobes, the infructescence often with elongated branches; seeds angular, 0.9-1.5 mm long, smooth.
Plants often found along rivers and in stream valleys in evergreen rain forest formations, from (100-)400 to 1400 m on the Caribbean slope and from (100-)500 to 1600 m on the Pacific slope. Flowering and fruiting throughout the year. The species occurs in Costa Rica and Panama.
Sommera donnell-smithii is recognized by its large obovate leaves with parallel (lineolate) minor venation, the dense yellowish puberulence, axil- lary inflorescences, small flowers with large calyx lobes and short corolla tube, and fleshy many- seeded fruit. The flowers and inflorescences are quite variable and give little support for delimiting additional species in our area. Williams based two new species on unusual lowland collections: S. dunlapii from Bocas del Toro, Panama (Dunlap 351 F) and S. rivularis from southern Puntarenas (Allen 5431 EAP, F, us). It is noteworthy that col- lections from below 200 m are quite rare (3 of 3 1 collections at F). Sommera grandis (Bartl.) Standl., listed by Standley (1938), is a species of Mexico.
Spermacoce Linnaeus
REFERENCES— C. D. Adams (manuscript sub- mitted to Novon, 1992). W. T. Gillis, The con- fused Spermacoce. Phytologia 29: 185-187. 1974.
to prostrate, stems quadrangular, glabrous or pubescent; stipules united with the petiole bases to form a sheath, with 3-15 stiff erect awns arising from the truncated (transverse) distal margin, persisting. Leaves opposite or pseudoverticillate (with pairs of smaller axillary leaves), petiolate or sessile; leaf blades usually narrow with strongly ascending secondary veins, entire, often sca- brous, without domatia. Inflorescences axillary or ter- minal, peduncles absent (in ours), the flowers fasciculate and verticillate or congested into heads, terminal heads subtended by 2 or more leaves (these sometimes invo- lucrate), bracteoles filiform when present, flowers sessile or subsessile. Flowers bisexual and monomorphic or di- stylous, small, hypanthium obovoid to turbinate, calyx tube short or absent, calyx lobes usually 4 (2-8), lobes acute to dentate and green; corolla funnelform or sal- verform, white to lavender, lobes 4 (3), valvate in bud; stamens 4 (3), filaments short, anthers linear to oblong, included or slightly exserted, often bluish; ovary 2-loc- ular with 1 ovule borne on the middle of the septum in each cell, style slender, stigma capitate or bilobed. Fruits capsular, 2-lobed and resulting in 2 cocci (both dehiscing adaxially or 1 dehiscing adaxially and the other inde- hiscent) or splitting down the center to expose both seeds, the calyx often persisting; seeds oblong to ellipsoid, with a narrow longitudinal ventral sulcus and lustrous mi- nutely pitted dorsal (abaxial) surface.
A pantropical genus of over 1 50 species, with many species in the American tropics. This genus is now interpreted to include Borreria. Spermacoce is difficult to separate from Crusea and Diodia, in which the cocci do not open. The often narrow subsessile leaves, small capitulate or axillary ses- sile inflorescences, four-lobed corolla, single-seed- ed cocci, and herbaceous weedy growth form dis- tinguish these plants and their close relatives: Crusea, Diodia, Mitracarpus, and Richardia. All these plants are characterized by a stipular sheath united to the leaf bases and truncated distally (straight or convex) with slender stiff awns or setae; all have strongly ascending secondary veins, and many have scabrous leaves. A number of our spe- cies appear to be recently introduced weeds, hav- ing been collected only a few times near the larger cities. As in the case of many other weedy plants, different individuals of the same Spermacoce spe- cies can differ greatly in overall size, form of the leaves, and development of inflorescences. This can make their identification difficult. Conse- quently, seeds are often necessary to confirm an identification.
We gratefully acknowledge the suggestions, de- terminations, and measurements provided by C. D. Adams, who has clarified the nomenclature and many of the species problems in this group; he is treating the Spermacoceae for the Flora Meso-
Annual or perennial herbs or small subshrubs, erect americana.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
313
Key to the Species of Spermacoce
1 a. Mature fruits splitting into 2 cocci, one of which does not open and the other of which splits longitudinally along the inner face, or with the mature fruits opening at the base and occasionally
opening slightly at the apex 2
Ib. Mature fruit splitting into 2 cocci, both of which open along the inner face, or open broadly at the top to release both seeds; seeds 0.6-3 mm long and usually easily removed from the cocci at
maturity by crushing between the fingers (species formerly placed in Borrerid) 4
2a. Fruits splitting into 2 and often opening slightly at the bottom; seeds 2-3 mm long [commonly
collected and wide-ranging] S. ocymifolia
2b. Mature fruits splitting into 2 cocci, one of which does not open and the other of which splits
longitudinally along the inner face; seeds 1.4-2.2 mm long (species of Spermacoce sensu
strictd) 3
3a. Leaves scabrous above; calyx lobes ciliate along the margins; fruits puberulent; rarely collected
(in Costa Rica) in open weedy sites S. confusa
3b. Leaves smooth above; calyx lobes glabrous along the margins; fruits glabrous; usually growing
along river edges and wet depressions S. tenuior
4a. Leaves often linear to lanceolate and pseudoverticillate; stems with larger rounded terminal inflo- rescences and only 1 (2, 3) additional nodes with axillary inflorescences (except S. densiflord) 5 4b. Leaves lanceolate to ovate, opposite or less often pseudoverticillate; stems usually with inflores- cences in the axils of many nodes along the stem, but with larger (> 1 cm) rounded terminal
inflorescences only in S. assurgens and S. vegeta 8
5a. Plants growing in water or wet mud; calyx lobes 4; fruits 6-7 mm long (including calyx lobes);
rarely collected in Costa Rica S. scabiosoides
5b. Plants not semiaquatic, not restricted to shallow water and wet mud; calyx lobes 2-4; fruits
1.5-5 mm long; commonly collected species 6
6a. Terminal capitulae usually exceeding 16 mm diam., often with well-developed capitulae at
the distal 3 nodes; calyx lobes 2; body of the fruits 2.5-5 mm long (seeds 1.4-2.2 mm long);
widespread 51. densiflora
6b. Terminal capitulae not usually exceeding 1 6 mm diam., capitulae usually only well developed
at the distal node; calyx lobes 2-4; body of the fruits 1.5-3 mm long; plants mostly confined
to the seasonally dry Pacific slope in Costa Rica 7
7a. Calyx lobes 2(-3) or with 2 larger and 2 very small lobes, narrowly clavate to acute, often
lacking a slender central pigmented midvein; seeds 1.3-1.6 mm long S. verticillata
7b. Calyx lobes 4 (rarely 3), usually pigmented along the slender midvein, narrowly acute; seeds
1 .6-2.2 mm long S. suaveolens
8a. Seeds with transverse sulci or depressions on the abaxial surface and easily seen with a hand lens ( x 10); plants often with hemispheric terminal heads 1-2 cm diam.; calyx lobes 4, but often unequal
9
8a. Seeds without transverse sulci easily seen with a hand lens (minute transverse pits present in some species but seen clearly only at x 40); plants rarely with hemispheric terminal heads; calyx lobes
2-4 10
9a. Calyx lobes ovate to broadly triangular, often rounded distally; seeds 2-2.2 mm long; upper
leaf surface usually glabrous or scabrous; rarely collected in southern Central America ....
S. vegeta
9b. Calyx lobes narrowly triangular, never rounded distally; seeds 1.1-2.1 mm long; upper leaf
surfaces glabrous or with thin hairs; plants of wide distribution S. assurgens
lOa. Corolla tubes 2-5 mm long, calyx lobes 4, 0.4—1.2 mm long; leaves often becoming more than 15
mm broad 12
lOb. Corolla tubes 0.5-2 mm long, calyx lobes 2-4, 0.4-2 mm long; leaves rarely becoming more than
1 3 mm broad 12
lla. Seeds 1.7-2.9 mm long, smooth or minutely reticulate; stems often with thin green longi- tudinal wings to 0.4 mm high; stipule sheath not beginning below the leaf bases; flowers
white, lavender, or blue S. latifolia
1 Ib. Seeds 1 .2-2. 1 mm long, reticulate or transversely sulcate; stems with longitudinal ridges only
3 1 4 FIELDI ANA: BOTANY
0.1-0.2 mm high; stipule sheath beginning well below the leaf bases; flowers white or tinged with pink S1. assurgens
12a. Calyx lobes 2 (often on only 1 side of the capsule); stems often with ciliate hairs along the longitudinal ribs beneath the nodes, leaves not more than 25 mm long; capsule wall thin and fragmenting, seeds 0.7-0.9 mm long, often yellowish, surface reticulate (x40) S. exilis
1 2b. Calyx lobes 4 (rarely 3 or 2); stems rarely with ciliate hairs along the ribs beneath the nodes, leaves often exceeding 25 mm length; capsule wall firmly cartilaginous, seeds 0.7-2.2 mm long, usually dark reddish brown, smooth or clearly reticulate 13
13a. Calyx lobes 4 or fewer, often unequal with the 2 smaller deciduous; seeds 0.7-1.2 mm long, with clearly reticulate surface and larger (0.05-0. 1 mm) pits; plants erect to prostrate or decumbent, usually glabrous 5". prostrata
13b. Calyx lobes 4 and equal, persisting; seeds 1.4-1.8 mm long, with finely reticulate (minutely pitted) surface; plants usually erect with slender stiff stems S. ovalifolia
Spermacoce assurgens Ruiz & Pav., Fl. Peruv. 1:
60, t. 92. 1798. Spermacoce laevis sensu auctores non Lam., Tabl. Encycl. 1: 231. Figure 5.
ams cited above). This is one of the most common species of Spermacoce in Central America. Com- pare S. vegeta, a closely related species.
Herbs, 20-80 cm tall, erect or spreading, leafy stems 0.8-3 mm diam., with thin whitish hairs 0.1-0.4 mm long, glabresent, the hairs often in narrow longitudinal ridges; stipule sheaths 1.5-5 mm high, 2-4 mm broad (to 8 mm beneath inflorescences), often obconic and aris- ing from beneath the leaf bases at sterile nodes, with 3- 9 awns 1-6 mm long, glabrous or sparsely pubescent. Leaves opposite or rarely pseudoverticillate, petioles 0- 6 mm long, slender; leaf blades l-5(-7) cm long, 0.5- 2. 5 (-3) cm broad, lanceolate to ovate-lanceolate or ovate- elliptic, apex acute to obtuse, base obtuse to cuneate or rounded, drying stiffly chartaceous, darker grayish green above, usually glabrous and scabrous above, glabrous or with stiff scabrous or thin hairs to 0.5 mm long beneath, 2° veins 4-6/side. Inflorescences terminal or axillary, globose to verticillate, 3-13 mm long, 5-15 mm diam., subtended by 2-4 leaf-like bracts 1-5 cm long, flowers sessile or subsessile and crowded. Flowers monomor- phic, hypanthium ca. 1.5 mm long, calyx lobes 4, 0.3- 0.7(-1) mm long, acute or blunt; corolla white, tube ca. 2 mm long, lobes ca. 1 mm long, triangular. Fruits 2- 2.5 mm long (not including the calyx lobes), ca. 2 mm broad; seeds 1.2-2.2 mm long, 0.6-1.1 mm broad, dark reddish brown, minutely reticulate and usually with nar- row transverse sulci abaxially.
Weedy plants of open sites in evergreen and partly deciduous forest areas, 0-1700(-2000) m elevation in Costa Rica. Flowering and fruiting throughout the year. This species ranges widely over the tropics and subtropics of the world.
Spermacoce assurgens is recognized by the stip- ular sheath arising from well beneath the leaf bases (especially noticeable at sterile nodes), having both terminal and axillary capitula, four small sepal lobes, small corollas, and seeds with transverse sulci (not always easily seen). Rare individual plants may occur with thin hairs over many surfaces and long-hirsute on ridges of the stems, but interme- diates can be found in Central America (see Ad-
Spermacoce confusa Rendle ex Gillis, Phytologia 29: 185. 1974 (Rendle's description in J. Bot. 74: 12. 1936, lacked a Latin diagnosis). S. ten- uoir sensu auctores non L. Figure 5.
Annual herbs or subshrubs to 80 cm tall, with few or many branches, leafy stems 0.5-2.5 mm thick, with 4 prominent longitudinal ridges and with scabrid hairs along the ridges; stipule sheath ca. 1 mm long, 1-3 mm broad, with 3-7 filiform awns 1-4 mm long, glabrous. Leaves usually opposite and subsessile, petioles 0-3(-6) mm long; leaf blades 2-6 cm long, 2-10(-l 3) mm broad, lanceolate to oblanceolate or linera-lanceolate, apex acute to acu- minate, base narrowed, with minute (0.1-0.2 mm) sca- bridulous hairs above, glabrous beneath, 2° veins 2-37 side, usually impressed above. Inflorescences small and axillary, few-flowered, ca. 4-6 mm long, of 6-15 flowers, the flowers sessile or subsessile. Flowers with hypanthi- um and calyx tube 1-1.5 mm long, calyx lobes 4, 0.4-1 mm long, triangular-lanceolate to linear-lanceolate, con- spicuously ciliolate along the edges; corolla white, rose, lavender, or blue, tube ca. 1.5 mm long and 1 mm diam., lobes 0.5-1 mm long, 0.4-0.5 mm broad, ovate-oblong, obtuse at the apex, minutely puberulent distally; fila- ments ca. 0.2 mm long, borne at the base of the tube, anthers ca. 0.3 mm long. Fruits 1.5-3 mm long, obovoid to ellipsoid or subglobose, with short ascending hispidu- lous hairs to 0.5 mm long, with persisting calyx lobes 0.5-1 mm long; seeds 1.1-2 mm long, 0.5-1 mm broad, ellipsoid-oblong to obovoid-ellipsoid, bluntly rounded at both ends, yellowish to dark brown, lustrous, with minute shallow pits ( x 40), sinus less than 'A the width of the seed and constricted in the middle.
Plants of open weedy sites, from near sea level to 1000 m elevation (to 1600 m in Guatemala). Flowering probably primarily in the wet season: July-December. The species ranges from Mexico through Central America and the West Indies into South America.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
315
Spermacoce confusa is recognized by the very narrow scabrous leaves, small compact inflores- cences, and the asymmetric unevenly setose cap- sule that breaks into two parts, with one failing to open. Rarely collected in Costa Rica (all from the Pacific slope and lowlands: Guanacaste, Puntar- enas, and San Jose provinces).
Spermacoce densiflora (DC.) Liogier, Phytologia 54: 113. 1983. Borreria densiflora DC., Prodr. 4: 542. 1830. 5. spinosa L., Sp. PI. ed. 2, 148. 1762, nom. illeg. (cf. Steyermark, 1972, p. 814). B. spinosa (L.) Cham. & Schlechtend., Linnaea 3: 340. 1828. Figure 4.
Herbs from a woody rootstock, 0.3-1 m tall, with few distal branches, leafy stems 1-4 mm thick, with 4 lon- gitudinal ridges, usually glabrous but scabrous; stipule sheaths 2-5 mm long, to 5(-l 1) mm broad, awns 3-9, 2.5-7 mm long, usually glabrous and pale yellowish gray. Leaves opposite or more often pseudoverticillate with 2 larger and 2-6 smaller leaves at the same node, petioles absent or poorly differentiated; leaf blades l-6(-7) cm long, l-8(-15) mm broad, linear to linear-lanceolate or linear-oblong, apex acute, base narrowly cuneate, drying grayish with incurved scabrid margins, stiffly charta- ceous to subcoriaceous, glabrous but with scabrous sur- faces, 2° veins 3-6/side. Inflorescences terminal and ax- illary to the distal 2(-3) nodes, 8-18 mm long and 14- 28 mm broad, capitate and hemispheric or verticillate, subtended by leaf-like bracts 1-5 cm long, flowers sessile and tightly congested. Flowers with hypanthium 2.5 mm long, pilose distally, calyx lobes 2, 0.9-2.5 mm long, narrow, ciliolate or glabrous; corolla white, campanulate, tube ca. 1 mm long, lobes ca. 1 mm long; anthers ca. 0.4 mm long. Fruits 3-5 mm long not including the calyx lobes; seeds 1.5-2.1 mm long and 0.4-0.6 mm broad, narrowly oblong, dark brown, surface weakly reticulate (x 10) with minute pits (x40).
Weedy plants of open sites in both evergreen and deciduous forest areas, from near sea level to 800 m (to 1200 m in Guatemala). Flowering in October-February. This species ranges from Mex- ico and the West Indies to northern South Amer- ica.
Spermacoce densiflora is recognized by the larg- er terminal hemispheric or subglobose heads, very narrow leaves, stipular sheaths often drying pale grayish, and narrow seeds.
Scheele. S. mauritiana Gideon, Kew Bull. 37: 547. 1983, nom. nov. for Borreria repens DC. non Spermacoce repens Cham. & Schlechtend. Figure 5.
•
Annual herbs, prostrate or clambering, to 30 cm long with few branches, leafy stems 0.6-1.8 mm thick, gla- brous or with cilliate hairs 0.2-0.5 mm long (especially along the raised longitudinal ribs below the node); stipule sheath 0.5-2 mm long, 1.5-6 mm broad, with 5-9 awns 1-2 mm long. Leaves usually opposite, less often verti- cillate with short axillary leaves, petioles 0-3 mm long, lateral margins continuous with the lamina margins; leaf blades 7-24 mm long, 3-1 1 mm broad, ovate-elliptic to ovate-oblong, ovate or elliptic, apex acute to obtuse, base cuneate and decurrent, drying grayish or yellowish green above, membranaceous to thin-chartaceous, glabrous or with thin hairs 0.1-0.4 mm long, scabrous above and along the margins, 2° veins 2-4/side. Inflorescences mostly axillary and verticillate with few sessile flowers, ca. 2 mm high and 6 mm broad, subtended by the stipules and leaves. Flowers with hypanthium 0.5-1 mm long, calyx lobes 2, ca. 0.5 mm long, linear and blunt or acute at the apex; corolla white or purplish white, tube ca. 0.5 mm long. Fruits 0.8-1 mm long, 1-2 mm broad when open, persisting calyx often recurved; seeds 0.7-0.9 mm long, 0.3-0.5 mm thick, oblong, sometimes appearing to have longitudinal ridges ( x 10) but with minute trans- verse pits (x40), yellowish to dark brown.
Weedy plants of open sites mostly in evergreen areas, 50-600 m elevation (higher elsewhere). Probably flowering throughout the year but pri- marily in the wet season: May-December. The species ranges from southern Mexico to South America and the West Indies. It occurs on Cocos Island and may be widely distributed in the Old World.
Spermacoce exilis is recognized by its small stat- ure, little leaves, axillary inflorescences, stems of- ten with ciliolate ridges below the node, minute flowers with two small calyx lobes, and small seeds. The seeds appear to be smooth but are character- ized by transversely elongated pits in vertical rows on the abaxial surface ( x 40) with neither row ori- entation dominant (C. D. Adams, pers. comm., June 1991); see illustration, upper right. The type of S. exilis, from Cocos Island, lacks the ciliolate ridges seen in most other collections but has the characteristic seed. This may be S. gracilis Ruiz & Pav. as used in the Flora of La Selva (Taylor 1991).
Spermacoce exilis (L. O. Williams) C. D. Adams, Novon (in press). Boreria exilis L. O. Williams, Phytologia 28: 227. 1974, nom. nov. for Bor- reria gracilis L. O. Williams, Phytologia 26: 487. 1973, not B. gracilis Miq. in Hook, f., nor
Spermacoce latifolia Aubl., Hist. pi. Guiane 1: 55, pi. 19, f. 1. 1775. Borreria latifolia (Aubl.) K. Schum. in Mart. Fl. Bras. 6(6): 61, pi. 80. 1888. Figure 5.
316
FIELDIANA: BOTANY
Herbs, prostrate, clambering or erect to 1 m high, branched mostly near the base, leafy stems 1-4.5 mm thick, quadrangular and usually with thin greenish lon- gitudinal ridges ca. 0.3 mm high, glabrous or with thin hairs to 0.7 mm long; stipule sheath 1-3 mm long, 2-7 mm broad, with 5-7 slender awns 1-6 mm long, glabrous or with thin hairs to 2 mm long. Leaves opposite and well separated along the stem, petioles 0-8 mm long, poorly differentiated from the lamina; leaf blades 10- 45(-80) mm long, 4-25(-40) mm broad, ovate-elliptic to obovate, elliptic or broadly elliptic, apex acute, base obtuse to cuneate and decurrent on petiole, drying mem- branaceous to stiffly chartaceous, usually greenish above, subglabrous with minute (0. 1 mm) hairs or with hairs to 0.9 mm long, 2° veins 3-6/side. Inflorescences axil- lary, 3-8 mm long, 7-15 mm broad, few-flowered, sessile and glomerulate, separate or verticillate, subtended by the inconspicuous stipule sheath and leaves, flowers ses- sile. Flowers with hypanthium 1-1.5 mm long, turbinate, sparsely puberulent with ascending hairs, sepal lobes 4, ca. 1 mm long, triangular; corolla salverform, white to lavender or blue, 2.5-5 mm long, tube ca. 3 mm long, 0.5-1 mm diam., anthers 0.5-1 mm long. Fruits 2.5-4 mm long, 3-4 mm broad when open, minutely pilose; seeds 1.7-2.9 mm long, 1-1.7 mm broad, oblong, mi- nutely reticulate or smooth, pale yellowish brown to dark brown.
Weedy plants of evergreen forest formations, from near sea level to ca. 1 100 m elevation. Prob- ably flowering and fruiting throughout the year. The species ranges from Mexico and the West In- dies to South America and is found in the Old World as well.
Spermacoce latifolia is distinguished by the broader (usually thin) opposite leaves, stems with conspicuous (often winged) longitudinal ridges, small axillary inflorescences, and larger seeds with very minutely reticulate surface. The leaves and stems are glabrous or only sparsely puberulent. The axillary inflorescences often have only a few (3-7) flowers and the leaves usually dry yellowish green. Mielcillo and chiritillo have been reported as common names.
Spermacoce ocymifolia Willd. ex Roem. & Schult., Syst. Veg. 3: 530. 1818. Hemidiodia ocimifolia (Willd. ex Roem. & Schult.) K. Schum. in Mart., Fl. Brasil. 6(6): 30, pi. 72. 1889. Diodia ocy- mifolia (Willd. ex Roem. & Schult.) Bremek., Rec. Trav. Bot. Neerl. 31: 305. 1934. Figure 6.
Herbs or subshrubs, 30-90(-150) cm tall, branches erect or spreading and decumbent, distal branchlets sim- ple or less often branched, leafy stems 0.5-3 mm thick, with 4 obscure longitudinal ribs and becoming terete, usually sparsely pubescent with thin whitish hairs 0.1- 0.3 mm long; stipule sheath 2-6 mm long, 2-6 mm broad, conical (often difficult to see beneath the inflorescences),
pubescent, with (3-)5-l 1 unequal erect or spreading awns on each side, 2-8 mm long. Leaves opposite, petioles 0- 12 mm long, 0.4-2 mm wide, sometimes with winged margins continuous with the leaf margins; leaf blades 2- 8(-l 1) cm long, 0.6-2(-3) cm broad, narrowly elliptic, narrowly elliptic-oblong or elliptic-lanceolate, apex ta- pering gradually and acute or acuminate, base attenuate and decurrent on petiole, the margins becoming revolute, leaves drying stiffly chartaceous, slightly scabrous above and glabrous or sparsely puberulent, glabrous or sca- brous beneath with short (0.1-0.2 mm) stiff hairs. 2° veins 3-6/side, strongly ascending. Inflorescences axil- lary and verticillate, 6-14 mm broad and 4-8 mm high, subtended by the petiolar sheath and its linear awns, the flowers densely crowded and sessile or subsessile. Flow- ers with hypanthium 1-2 mm long, narrowly oblongoid, with short stiff hairs, calyx lobes 0.3-0.5 mm long and ca. 0.4 mm broad, acute; corolla white, funnelform, tube 1.3-2.5 mm long, ca. 0.6 mm diam. at the base and 1 mm at the mouth, glabrous externally, with short hairs at the mouth within, lobes 3—4, 1-2 mm long and ca. 0.5 mm broad, oblong-lanceolate; stamens 3-4, fila- ments 0.6-2 mm long, anthers 0.6-0.9 mm long, oblong; style ca. 2.5 mm long. Fruits 2.5-4 mm long, 1.5-2 mm broad, oblong-turbinate to obovoid-oblong, glabrous near the base and with thin erect hairs 0.2-0.3 mm long dis- tally, persisting calyx 0.3 mm long, splitting into 2 mer- icarps; seeds 2-3 mm long, 0.6-1.3 mm thick, narrowly oblong, smooth or with obscure transverse depressions abaxially ( x 10) and minutely pitted surface ( x 40), dark reddish brown.
Weedy plants of open sites in evergreen or partly deciduous areas, from near sea level to 1200 m elevation in Costa Rica. Collected on the Carib- bean slope and through the length of the cordil- leras. This species has not been collected in the drier deciduous or semideciduous Pacific slope (0- 500 m) of Central America. Probably flowering and fruiting throughout the year. The species rang- es from Mexico, Central America, and the West Indies to Paraguay. It is found on Cocos Island and in the Old World.
Spermacoce ocymifolia is recognized by the short weedy habit, erect stems, the narrow opposite leaves with strongly ascending veins, sheathing stipules with long narrow awns, the densely crowd- ed small verticillate flowers, and the obovoid cap- sular fruit with short thin erect hairs and short (0.5 mm) calyx lobes. The capsule splits into two mer- icarps, which tend to open near the base adaxially but occasionally the capsules open slightly at the top before separating from the base. These plants closely resemble species of Crusea and Diodia.
Spermacoce ovalifolia (M. Martens & Galeotti) Hemsl., Biol. centr. amer. Bot. 2: 59. 1881. Bor- reria ovalifolia M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 1 1: 129. 1844. S.prin-
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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glei S. Watson, Proc. Amer. Acad. Sci. 25: 152. 1 890. S. ernstii R. Fosberg & D. Powell, Smiths. Contr. Bot. 45: 29. 1980. Figure 5.
Annual herbs to 40 cm tall, often branched near the base and with few distal branches, leafy stems 0.5-2 mm thick, with 4 longitudinal ribs ca. 0.2 mm high, glabrous or minutely puberulent at the nodes; stipule sheath 0.5- 3 mm high, often obscured by the inflorescences, with 3-11 awns 1-5 mm long. Leaves opposite (occasionally pseudoverticillate) with 2—4 smaller axillary leaves, pet- ioles 0-5 mm long, with lateral margins continuous with the leaf margins; leaf blades 6-48 mm long, 2-16 mm broad, elliptic to narrowly elliptic, elliptic-lanceolate or ovate-elliptic, apex acute, base acute to cuneate, drying membranaceous, brownish or dark green above, gla- brous or with a few scabrous hairs above, subglabrous or with minute (0. 1-0.2 mm) stiffhairs beneath, scabrous on the edges, 2° veins 2-5/side. Inflorescences axillary and verticillate or terminal and subtended by 2-6 leaves, 2-4 mm high, 3-6 mm broad, with few (5-12) flowers. Flowers with 4 calyx lobes 0.6-1.1 mm long, 0.1-0.2 mm broad, linear-aculeate, persisting; corolla minute. Fruits ca. 2 mm long and 2 mm broad when opened, body of the capsule ca. 1.5 mm long, glabrous externally or minutely (0.05-0.1 mm) hispid distally; seeds (1.3-)1.5-1.8 mm long, 0.8-0.9 mm broad, very finely reticulate ( x 1 0) and minutely pitted ( x 40), dark reddish brown.
Weedy plants of open sites in evergreen and partly deciduous areas of the Meseta Central, 900- 1400 m elevation. Flowering in the wet season: July-December. The species ranges widely, from the southern United States to South America, the West Indies, and the Old World.
Spermacoce ovalifolia is recognized by its short stature, mostly opposite (not verticillate) leaves, small axillary inflorescences, and slightly larger seeds with minutely pitted surface. Many of our specimens were earlier placed under the name Bor- reria ocymoides (Burm.) DC., an Asian species. These plants are very similar to S. exilis and S. prostrata, with smaller seeds. This species is rare in Costa Rica and perhaps a recent introduction.
Spermacoce prostrata Aubl., Hist. pi. Guiane 1: 58, pi. 20. f. 3. 1 775. Borreria repens DC., Prodr., 4: 533. 1830. B. parviflora G. F. W. Meyer, Prim, fl. Esseq. 83. 1818. Figure 5.
Prostrate or decumbent annual herbs to 40 cm tall, branching mostly near the bases, leafy stems 0.4-2 mm thick, glabrous or rarely with thin hairs to 0.8 mm long, with 4 longitudinal ribs 0. 1-0.3 mm high; stipule sheaths 1-3 mm high, 1.5-6 mm broad, with 3-9 awns 1-2 mm long. Leaves opposite or pseudoverticillate with 2-4 smaller axillary leaves at each node, sessile or subsessile; leaf blades 10-35 mm long, 2-18 mm broad, elliptic to
narrowly elliptic or elliptic-lanceolate, apex acute (ob- tuse), base cuneate and decurrent on petiole, drying membranaceous to chartaceous, greenish or yellowish brown, glabrous or with minute (0.2 mm) stiffhairs above and below, scabrous on the margin, 2° veins 2-4/side. Inflorescences mostly axillary (sometimes terminal), 2- 4 mm high and 3-8 mm broad, flowers few (3-8) to many (ca. 20) and sessile. Flowers with hypanthium 0.5- 1 mm long, calyx lobes usually 2 larger and 2 smaller, 0.6-1 mm long; corolla white, less than 3 mm long. Fruits ca. 2 mm long, body of the fruit 0.8-1.2 mm long; seeds 0.7-1.2 mm long, 0.3-0.5 mm broad, reticulate, often with longitudinal ridges (xlO) and distinctive trans- versely elongated pits ( x 40), dark reddish brown.
Common weedy plants of open sites in ever- green and partly deciduous forest regions (only rarely collected in deciduous forest areas), from near sea level to 1 900 m elevation. Probably flow- ering and fruiting throughout the year. The species ranges from the southern United States to South America and the West Indies.
Spermacoce prostrata is recognized by the small weedy habit, usual lack of puberulence, often ses- sile leaves, minute flowers, and small seeds with reticulated surface of distinctive pits. These pits are larger (0.05-0.1 mm) than in S1. exilis or S. ovalifolia and they are transversely oblong ( x 40); see Figure 5, upper center. The fruits usually ap- pear to have only two calyx lobes because the two smaller lobes fall off. Many of these specimens were earlier placed under the name Borreria ocy- moides (Burm.) DC., an Asian species. Compare with S. exilis.
Spermacoce scabiosoides (Cham. & Schlechtend.) O. Kuntze, Rev. gen. pi. 3: 123. 1898. Borreria scabiosoides Cham. & Schlechtend., Linnaea 3: 318. 1828.
Semiaquatic herbs, floating, procumbent or erect, to 1 m tall, distal stems with long (3-10 cm) internodes and few or no lateral branches, leafy stems 1-5 mm thick, glabrous or rarely with slender hairs to 1.5 mm long at the nodes, terete; stipule sheaths 3-9 mm long, beginning well below the petiole base, 3-12 mm broad, with 3-7 awns 2-7 mm long. Leaves opposite or occasionally pseudoverticillate, petioles 3-10 mm long, with winged lateral margins continuous with the lamina margins; leaf blades 2-10 cm long, 3-16 mm broad, linear-lanceolate to narrowly lanceolate or linear-oblong, apex tapering gradually and narrowly acuminate, base tapering grad- ually and cuneate-decurrent, drying chartaceous, dark above, glabrous or with scabrid hairs, 2° veins 3-4/side. Inflorscences terminal or axillary, 3-12 mm long, 8-16 mm broad, capitate or verticillate, subtended by 1-2 pairs of narrow leaves (rarely without leaves and ap- pearing pedunculate), flowers tightly congested and ses- sile, often few-flowered. Flowers ca. 8 mm long, calyx
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puberulent distally, lobes 4, 1-2 mm long, 0.2-0.3 mm wide, narrow and acute; corolla white, funnelform, pu- berulent distally on the interior and exterior, tube 3-4 mm long, 1.5-3 mm diam. distally, lobes 1.3-2 mm long, triangular; anthers 1-1.5 mm long. Fruits ca. 5 mm long (not including the calyx lobes), ca. 2.5 mm broad distally, obovoid, glabrous in the lower half, splitting from above; seeds 2.6-3 mm long, narrow.
Aquatic plants of evergreen or deciduous low- land vegetation. Collected in Santa Rosa National Park (Liesner 5261 CR, MO), flowering in February, and growing in persisting water of a lagoon. The species ranges from Costa Rica and Panama to Brazil and Argentina.
Spermacoce scabiosoides is distinguished by its growth in shallow water or wet mud, stipular sheath beginning well below the petiole bases, glabrous vegetative parts (but with slender longer hairs oc- casionally at the nodes), narrow leaves to 10 cm long, congested flower, and larger corolla and fruits.
Spermacoce suaveolens (G. F. W. Meyer) O. Kuntze, Rev. gen. pi. 3: 124. 1898. Borreria suaveolens G. F. W. Meyer, Prim. Fl. Esseq. 8 1 , pi. 1. 1818. Figure 4.
Herbs from a woody base, usually erect 15-40(-60) cm tall, branched from the base and often forming dense tufts, distal branches few, leafy stems 0.7-4 mm thick, with 2 or 4 well- or poorly denned longitudinal ridges, puberulent with minute (0.1-0.3 mm) hairs or subgla- brous with a few hairs at the nodes; stipule sheaths 1-3 mm high, 1-4 mm broad, with 3-7 stiff yellowish awns to 6 mm long. Leaves opposite or more often pseudover- ticillate, 1 pair of larger leaves and 2-4 additional pairs of smaller leaves at a node, petioles usually absent; leaf blades 6-45(-60) mm long, l-3(-4) mm wide, linear- lanceolate to linear or linear-oblong, apex tapering grad- ually and acute, base tapering gradually and narrowly cuneate, usually drying stiffly chartaceous, grayish, gla- brous above, minutely puberulent beneath, 2° veins 2- 3/side. Inflorescences terminal (or sometimes axillary and verticillate at the next-to-last node), 4-10 mm high, 8-16 mm broad, dense hemispheric (subglobose) capit- ula, usually subtended by 4 leaves 1-4 cm long and 1- 3 mm broad, often with slender awns among the sessile flowers. Flowers with a hypanthium 0.8-1.7 mm long, with thin whitish hairs near the apex, calyx lobes 4 (rarely 3 or 2), 1.4-2(-2.5) mm long, narrowly triangular and acute at the apex; corolla white to pale pink, tube 1.1- 2.3 mm long, 0.4-1.1 mm diam., lobes 4, 1-1.5 mm long; anthers 0.8-1.2 mm long. Fruits ca. 3 mm long and 1.5 mm diam; seeds 1.7-2.2 mm long, 0.6-0.7 mm thick, dark reddish brown with minutely reticulate sur- faces.
Plants of savannas and open sites in deciduous and partly deciduous areas of the Pacific slope
(rarely in evergreen formations of the central high- lands), 20-1300 m elevation in Costa Rica. It is found around 2000 m elevation on Volcan Baru, Chiriqui, Panama. Flowering is largely in the wet season: May-December. The species ranges from Mexico and the West Indies to southern South America.
Spermacoce suaveolens is recognized by its stiff erect stems usually with four to eight unequal leaves at a node, the rounded terminal inflorescences, the four narrow sepal lobes, and the seeds with retic- ulate surfaces. Dried specimens may be virtually identical to those of S. verticillata; flowers need to be carefully examined to distinguish the two spe- cies. Compare this material with S1. capitata Ruiz & Pav. and Borreria capitata (Ruiz & Pav.) DC. of South America.
Spermacoce tenuior L., Sp. PL 102. 1753. S. glabra auct. non Michx., S. riparia Cham. & Schlech- tend., Linnaea 3: 355. 1828. Figure 6.
Perennial herbs to 80 cm tall, woody at the base, twigs ascending and with few branches, leafy stems 0.2-2.8 mm thick, with 4 longitudinal smooth or scabrous ridges, glabrous or with a few hairs; stipule sheath 0.5-2.5 mm long, with 3-7 awns 1.5-4 mm long. Leaves with petioles 3-7 mm long, glabrous; leaf blades 3-6 cm long, 3- 1 7(-2 1 ) mm broad, lanceolate to linear-lanceolate or ob- long-lanceolate, apex acute to acuminate, base acute, glabrous on both sides but scabrous, 2° veins 3-5/side. Inflorescences small and axillary, usually verticillate, 5- 1 0 mm broad, with 1 0-20 flowers, bracts 1 .5-3 mm long, filiform. Flowers with hypanthium ca. 2.2 mm long and 1 mm diam., calyx lobes 4, 0.5-0. 9(-l. 2) mm long, lan- ceolate to linear-lanceolate, ciliolate only at the base; corolla white, rotate, lobes 0.8-1.2 mm long, 0.4 mm broad, longer than the tube; filaments ca. 0.2-0.3 mm long, borne in the middle of the tube, anthers ca. 0.5 mm long, included. Fruits 2.5-3 mm long, obovoid or oblong-pyriform, usually glabrous; seeds 1 .4-2 mm long, 0.7-1 mm broad, dark reddish brown to purplish brown, rounded at both ends, surface, minutely foveolate ( x 40) with circular pits at the broader apical end and slightly vertically elongated pits at the basal end, in 10-16 ver- tical rows and 25-28 transverse rows, sinus more than 'A the width of the seed.
Herbs of open sites, especially along streams and wet depressions, 0-1200 m elevation. Flowering in January-August in Central America. The spe- cies ranges from the southeastern United States, the West Indies, Mexico, and Central America through most of South America.
Spermacoce tenuior is recognized by its prefer- ence for wet habitats, narrow glabrous leaves (sca- brous on the edges), small sessile inflorescences,
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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little white flowers, and minutely pitted seeds. The fruit splits from the top into an open half releasing the seed and an indehiscent half tightly enclosing the other seed.
Spermacoce vegeta (Standl. & Steyerm.) C. D. Ad- ams, Novon (in press). Borreria vegeta Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 21. 1943.
Annual herbs, erect, 10-30 cm tall, leafy stems 1-5 mm diam., longtudinal ribs obscure in early growth, sparsely to densely puberulent with thin crooked whitish hairs 0.2-0.6 mm long; stipule sheath 3-8 mm long, 6- 23 mm broad, often marked with reddish spots, with 7- 1 3 subequal awns to 9 mm long. Leaves opposite or more often pseudoverticillate with smaller axillary leaves, pet- ioles 3-10 mm long, with broad lateral margins; leaf blades 25-90 mm long, 6-40 mm broad, elliptic-lan- ceolate to ovate-elliptic or lanceolate, apex tapering grad- ually and attenuate-acuminate, base abruptly narrowed and obtuse or cuneate, decurrent on the winged petiole, drying chartaceous, grayish green or yellowish green above (paler grayish beneath), glabrous or with short scabrous hairs above, sparsely to densely puberulent with thin whitish hairs 0.2-0.5 mm long beneath, 2° veins 4-77 side. Inflorescences terminal or axillary, hemispheric or verticillate, to 1 5 mm high and 25 mm broad, subtended by the leaves and enlarged stipular sheath, flowers dense- ly congested. Flowers with 4 calyx lobes, 0.5-1.3 mm long, ca. 1 mm broad at the base, ovate or suborbicular to broadly triangular, rounded to obtuse at the apex, ciliolate; corolla white, ca. 2.5 mm long, anthers ca. 0.4 mm long. Fruits ca. 3 mm long, villose to glabrous; seeds 2-2.2 mm long, 0.8-0.9 mm broad, narrowly oblong, with prominent transverse sulci, dark reddish brown.
Plants of open sites in partially deciduous or evergreen areas, from 600 to 900 m elevation in Guatemala and Honduras (1600 m in Chiriqui). Flowering and fruiting occur in the latter part of the wet season: September-January. This species is native to northern Central America and appears to have been introduced in Costa Rica and Pan- ama.
Spermacoce vegeta is distinguished by its erect stature, larger elliptic-lanceolate leaves, larger ter- minal and axillary inflorescences that are sub- tended by leaves and expanded stipular sheaths, broad calyx lobes, and seeds with clearly de- marked transverse depressions. It is closely related to S. assurgens, but that species is smaller and prostrate to decumbent.
Erect herbs, 15-60(-100) cm tall, perennial from a woody base, branching from the base and with opposite branches along the stems, leafy stems 0.6-4 mm thick, usually with 4 slightly elevated longitudinal ridges, gla- brous or minutely (0.1-0.2 mm) puberulent near the nodes; stipule sheaths ca. 1 mm long, to 3 mm broad, with 3-5 yellowish awns 1-5 mm long. Leaves opposite or pseudoverticillate (4, 6, 8 per node), separated by 2- 5 cm along the stem, a well-defined petiole absent; leaf blades 1-4 cm long, 0.7-5(-8) mm wide, linear to linear- lanceolate, lanceolate or narrowly oblong, apex tapering gradually and acute, base narrowly cuneate, drying stiffly chartaceous to subcoriaceous, grayish with the outer margins becoming involute, glabrous above and below, 2° veins 2-3/side. Inflorescences mostly terminal, a few axillary to distal nodes (rarely more than 2 on a distal stem), 8-14 mm diam., globose to hemispheric capitula, subtended by 2-4 slender leaf-like bracts 1-2 cm long, flowers sessile and densely congested. Flowers 2.5-4.5 mm long, hypanthium 0.5-1 mm long, glabrous or sparsely puberulent, calyx lobes 2(-3-4), ca. 1 mm long and 0.2-0.3 mm broad, narrowly clavate and blunt at the apex or acute, often more darkly pigmented distally, lobes often connected at the base by hyaline tissue 0.4 mm high; corolla white, salverform, tube 1-3 mm long, ca. 1 mm diam., lobes 0.8-1.4 mm long; anthers 0.5- 0.8 mm long. Fruits ca. 1.8 mm long (not including the sepal lobes); seeds 1.3-1.6 mm long, 0.4-0.6 mm thick, narrowly oblong, dark brown and minutely reticulate.
Weedy plants of the seasonally very dry and deciduous Pacific lowlands, 0-300 m elevation (to 900 m in Honduras). Flowering in the wet season: May-November. The species ranges from the southern United States to southern South Amer- ica, the West Indies, and the Old World.
Spermacoce verticillata is distinguished by its erect stems with opposite branches and short nar- row subsessile verticillate leaves, dense terminal heads, two-lobed calyx, minute corolla, and small reticulate seeds. There may be two minute calyx lobes between the two well-developed lobes. Dried material of this species can be virtually identical to that of the more commonly collected Sper- macoce suaveolens.
Tocoyena Aublet
REFERENCES—!. D. Dwyer, Borojoa and Toco- yena (Rubiaceae) in Panama. Phytologia 17: 445- 449. 1969. I. Silberbauer-Gottsberger, G. Gotts- berger and F. Ehrendorfer, Hybrid speciation and radiation in the Neotropical woody genus Tocoye- na (Rubiaceae). PL Syst. Evol. 181: 143-169. 1992.
Spermacoce verticillata L., Sp. PI. 102. 1753. Bor- reria verticillata (L.) G. F. W. Meyer, Prim. Fl. Esseg. 83. 1818. Figure 4.
Trees or shrubs, stems terete, mostly glabrous; stipules interpetiolar, free or connate (intrapetiolar) at the base, narrowly triangular to ovate, usually persisting. Leaves opposite or 3/node, petiolate; leaf blades entire and pin-
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nately veined, drying black or dark brown, stiffly char- taceous to coriaceous. Inflorescences terminal or sub- terminal, corymbose, capitate or cymose, without bracts, flowers sessile to pedicellate. Flowers bisexual, radially symmetrical (or curved), large, very fragrant, hypanthi- um/ovary cylindrical-turbinate to obovoid, calyx lobes 4-6, persisting; corolla long, white or cream, salverform to tubular, corolla tube usually with a puberulent mouth, lobes 4-6, convolute in bud; stamens 4-6, inserted at the corolla mouth, anthers dorsifixed, 2-lobed at the base, exserted; ovary 2-locular, with many horizontal ovules on axile placentation. Fruits baccate or hard, globose to ovoid or oblong, pericarp thick and usually coriaceous, 2-locular; seeds many, horizontal and somewhat flat- tened, imbedded in a pulp.
A genus of ca. 1 8 species, nearly all South Amer- ican. The very long narrow corolla tubes, flowers closely clustered on a terminal inflorescence, and large solitary fruit with horizontally oriented seeds in a fleshy pulp characterize our species. Despite the numerous flowers within each inflorescence, apparently only one large fruit develops at the tips of fruiting branches.
Tocoyena pittieri (Standl.) Standl., Contr. Arnold Arbor. 5: 151. 1933. Posoqueria pittieri Standl., J. Wash. Acad. Sci. 18: 167. 1928. Tocoyena obliquinervia (Standl.) Standl., Contr. Arnold Arbor. 5: 152. 1933. Posoqueria obliquinervia Standl., J. Wash. Acad. Sci. 18: 167. 1928. T. cuatrecasasiiSteyerm., Acta Biol. Venez. 4: 113. 1964. Figure 30.
Trees, 10-18 m tall, trunk 30-60 cm diam., leafy branchlets 4-12 mm thick, glabrous and drying grayish; stipules 7-10 mm long, triangular, acute to acuminate, glabrous, persisting. Leaves with petioles (l-)2-4.5 cm long, 1.5-3.5 mm broad, glabrous; leaf blades 20-35 cm long, 8-20 cm broad, obovate to obovate-elliptic or broadly elliptic, apex abruptly narrowed or rounded and short-acuminate, tip 5-10 mm long, base gradually nar- rowed and obtuse or cuneate, drying chartaceous, usually blackish, glabrous above and below or with few minute hairs on the major veins beneath, sometimes with tufted hairs (domatia) in the vein axils beneath, 2° veins 9- 137 side. Inflorescence terminal and solitary or 3, 5-17 cm long (including corollas), to 1 5 cm broad, a dense corymb or panicle with crowded cymose branches, peduncles very short (1-5 mm) beneath the primary branches, gla- brous, bracts and bracteoles minute (1-2 mm long) and triangular-acuminate, pedicels 0-5 m long, merging with the hypanthium. Flowers glabrous externally and drying blackish, hypanthium 4-5 mm long, oblong-turbinate, calyx tube 2-3 mm long, calyx lobes minute, broadly triangular or rounded and apiculate; corolla long-tubular, carnose, yellow, tube 5-8 cm long, 1.5-3 mm diam., lobes 6-12 mm long, 5-7 mm broad, rounded distally and spatulate, imbricate, minutely papillate within. Fruits apparently solitary and sessile, ca. 1 0 cm long and 9 cm
diam., subglobose, the wall thick (8-10 mm dried) and warty lenticellate; seeds to 20 mm long, 13-16 mm wide, and 3-4 mm thick, imbedded in a fleshy matrix.
Trees of lowland rain forest or partly deciduous forest formations, from near sea level to 300 m elevation. Immature flowers were collected in March (Jimenez & Zuniga 814 CR, Tonduz 9528); mature flowers were collected in April (Hammel & Bozzoli 16628 CR, F, MO, and Q. Jimenez et al. 579 CR, F). Fruiting in April. The type of T. ob- liquinervia (Tonduz 9528 us) was collected from 200 to 250 m elevation along Rio Naranjo, San Jose; the other collections are from the Reserva Biologica Carara and the Golfo Dulce region. The species ranges southward to Colombia.
Tocoyena pittieri is distinguished by the large leaves, long-tubed flowers in crowded inflores- cences, and large round fruit with many horizontal seeds in a fleshy pulp. Nearly all parts of the plant (excepting the stems) turn black on drying. At first, it appeared that Standley was correct in separating T. obliquinervia as a separate species. However, recently collected material suggests that all our collections are part of the variation to be expected within T. pittieri. The fruit requires a year to ma- ture, according to Croat (1978).
Uncaria Schreber
REFERENCE— C. E. Ridsdale, A revision of Mi- tragyna and Uncaria (Rubiaceae). Blumea 24: 43- 100. 1978.
Woody lianas or clambering shrubs, climbing with the aid of recurved spines (developed from partly formed peduncles), young stems quadrangular or terete, glabrous or puberulent; stipules interpetiolar, entire to bifid, usu- ally with colleters at the base within. Leaves opposite, decussate, petiolate; leaf blades entire and pinnately veined, domatia often present in the vein axils beneath. Inflorescences axillary or terminal, flowers in globose capitula, solitary and pedunculate or on open opposite- branched panicles of 3-5 heads, flowers closely con- gested, sessile or pedicellate and bracteolate. Flowers bisexual and monomorphic, radially symmetrical, green- ish white to yellowish or orange, hypanthium turbinate, glabrous to densely puberulent, calyx lobes 5; corolla salverform or slightly funnelform, puberulent to glabrous externally, lobes 5, valvate in bud; stamens 5, filaments short, inserted in the throat of the tube, anthers oblong and sagittate at the base (or sterile and not sagittate), exserted; ovary 2-locular, ovules numerous ascending, placentation from the upper part of the septum, stigma capitate. Fruits thin-walled capsules, 2-locular, septici- dally dehiscent into 2 valves that usually remain united at the base, each valve slightly 2-parted from the top;
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seeds many, imbricate, winged at both ends with the lower end deeply bifid.
A genus of 34 species: 2 species are American, 3 species occur in Africa and Madagascar, and 29 species are found in Asia, Australia, and nearby areas. The woody vining habit, paired recurved spines at many nodes, flowers in globose heads and capsular fruits with many imbricate winged seeds distinguish this genus. Only one species is found in Central America. The flowers and capit- ula are similar to those of Cephalanthus occiden- talis L., which ranges from southeastern Canada to Guatemala.
Uncaria tomentosa (Willd. ex Roem. & Schult.) DC., Prodr. 4: 349. 1830. Nauclea tomentosa Willd. ex Roem. & Schult., Syst. Veg. 5: 221. 1819. N. aculeata H.B.K., Nov. gen. sp. 3: 382 (quarto). 1819, non Willd. 1790. Figure 37.
Vines or woody lianas to 30 m high, 8-25 cm diam. near the ground, leafy stems 2-6 mm thick, minutely appressed-puberulent with pale brownish hairs 0. 1-0.4 mm long, quadrangular and becoming terete, hooked thorns to 15(-20) mm long and 6-10 mm broad at the base, recurved; stipules 6-14 mm long, to 9 mm broad at the base, ovate-triangular, minutely puberulent, per- sisting or deciduous. Leaves usually distichous in a single plane, petioles 8-1 8 mm long, 1 .2-2 mm thick, minutely puberulent; leaf blades 7-15 cm long, 4-9 cm broad, ovate to broadly oblong, apex obtuse to subacuminate, base obtuse to subtruncate or rounded and subcordate, drying stiffly chartaceous, much darker above than be- low, glabrous and often lustrous above, pale grayish be- neath and glabrous or with longer (0.3-0.7 mm) thin ascending hairs, 2° veins 5-8/side, pit domatia some- times present in the vein axils beneath. Inflorescences axillary, 6-15 cm long, of 3 or 5 spherical heads on an open opposite-branched panicle, heads 1 5-25 mm diam. at an thesis, peduncles 1.2-4(-7) cm long, to 5 mm thick at the base (and resembling the recurved spines), mi- nutely appressed-puberulent, lateral branches subtended by broadly ovate stipule-like bracts, flowers sessile and tightly clustered, bracteoles obscure. Flowers yellowish white to golden yellow, hypanthium 1-1.5 mm long, obconic, calyx tube ca. 0.5 mm long, calyx lobes 0.2- 0.3 mm long and 0.4 mm broad at the base, rounded distally; corolla tubular-salverform, tube 4.5-6 mm long and 0.5-0.9 mm diam., with minute whitish hairs, lobes 1-1.8 mm long, ca. 0.8 mm broad, oblong, glabrous within; anthers sessile on the mouth of the tube, partly exserted, ca. 1.3 mm long; the ellipsoid stigma extended to 4 mm beyond the mouth of the tube. Fruits 7-9 mm long and 4 mm broad, ellipsoid with a short crown of persisting calyx, longitudinally ribbed and puberulent; seeds 3^4 mm long and 0.4 mm broad (including the unequal wings), body of the seed ca. 0.5 mm long.
Plants of evergreen rain forest formations of the
Caribbean lowlands, from 10 to 300 m elevation. Probably flowering primarily in January-April; fruiting in April-September. The species ranges from Guatemala and Belize through Central America to Colombia, Ecuador, and Venezuela.
Uncaria tomentosa is recognized by the climb- ing habit and thick recurved thorns, the opposite- branched inflorescences of closely compacted ses- sile flowers in globose heads, the slender corolla tubes, and the small capsules with narrowly winged seeds. These lianas can grow into the canopy to 30 m high (Croat, 1 978). The leaves are often whit- ish below in life. The sharp thorns can easily cause injury and make these plants troublesome weeds (called rangayo in Costa Rica).
Vangueria Jussieu
Shrubs or small trees, glabrous or pubescent, usually without spines; stipules interpetiolar, often broad at the base, villous within. Leaves opposite, short-petiolate, blades entire, glabrous or pubescent, domatia absent. Inflorescences axillary, usually arising from older leafless nodes, paniculate or racemose, flowers in distal cymose groups, bracteate. Flowers bisexual, calyx conic to rounded, 5-lobed, usually persisting, glabrous or pubes- cent; corolla campanulate or salverform, villous in the throat, lobes 5, valvate in bud, often as long as the tube; stamens 5, borne near the apex of the tube, filaments short, anthers oblong to ovate; ovary 5-locular, 1 ovule per locule pendulous from the apex, stigma thick, usually 5-lobed. Fruits drupaceous, usually crowned by the per- sisting sepal lobes, usually 5-seeded; seeds oblong to subglobose, with bony testa.
A genus of 27 species of tropical Africa and Madagascar. One species is occasionally planted or escaped in the New World tropics.
Vangueria madagascariensis J. F. Gmelin, Syst. Nat. ed. 13, 2: 367. 1791. V. edulis Vahl, Skr. Naturh.-Selskr. Kjobenhavn 2: 208, t. 7. 1792.
Shrubs or small trees, 5-15 m tall, leafy stems 1.5-5 mm thick, glabrous; stipules 4-7 mm long, ca. 3 mm broad at the base, triangular with narrow acute apex, united for 0.5-1 mm above the petiole. Leaves with pet- ioles 5-16 mm long, ca. 1 mm thick, glabrous; leaf blades 7-20(-28) cm long, 4-10(-15) cm broad, ovate to ovate- elliptic or elliptic, minutely puberulent on the veins above, 2° veins 7-9/side. Inflorescences 3-6 cm long, equally broad, branched near the base, peduncles 2-5 cm long, ca. 1 mm thick, with ascending or curled hairs 0.2-0.5 mm long, pedicels to 4 mm long, minutely pubescent. Flowers with hypanthium 0.5-0.7 mm long, conic, calyx lobes 0.5-1.5 mm long, triangular; corolla pale yellow, usually glabrous, tube 3.5-4 mm long, ca. 2.2 mm diam..
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lobes ca. 2 mm long, becoming reflexed; stamens bluish lilac. Fruits 2.5-5 cm diam., green to brownish, with 4- 5 thick woody pyrenes.
Known from only a single collection in Costa Rica: Chacon & Chacon 2258 CR, flowering in Sep- tember near Volcon Cacao at 1 000 m elevation in Guanacaste Province. The species, native of Af- rica, is grown in gardens and called "tamarind of the Indies."
Warszewiczia Klotzsch
Trees or shrubs; stipules interpetiolar, with glandular colleters at the base within, persisting. Leaves opposite and decussate or verticillate, petiolate; leaf blades large, entire and pinnately veined, domatia rarely present. In- florescences terminal or in the axils of distal leaves, made up of small trichotomous cymes along an elongate ra- cemiform axis, with or without expanded colorful calyx lobes. Flowers bisexual and monomorphic, small, hy- panthium and calyx tube campanulate to cupulate, calyx lobes 5, 1 of the lobes often expanded into a large colorful leaf-like blade; corolla funnelform to salverform, corolla tube villous within, lobes 5, implicate in bud; stamens 5, filaments borne in the throat of the tube, anthers dor- sifixed and versatile, sagittate at the base; ovary 2-loc- ular, ovules many and horizontal, placenta borne on the septum. Fruits woody capsules with septicidal basi petal dehiscence, the 2 valves entire or slightly bifid at the top; seeds many, horizontal, with a narrow margin around the edge.
A genus of about six species, ranging from Cen- tral America into South America. The genus is represented by a single very distinctive species in Costa Rica. Warszewiczia coccinea is perhaps our most colorful native species of Rubiaceae, having long inflorescences with small opposite flower clusters, each with a single brilliant calyx lobe.
Warszewiczia coccinea (Vahl) Klotzsch, Monats- ber. Konigl. Preuss. Akad. Wiss. Berlin 1853: 497. 1853. Macrocnemum coccineum Vahl, Symb. Bot. 2: 38. 1791. Mussaenda coccinea (Vahl) Poir. in Lam., Encycl. 4: 394. 1797. Ca- lycophyllum coccineum (Vahl) DC., Prodr. 4: 367. 1838. Figure 16.
Shrubs or small trees, 2-10(-l 5) m tall, leafy branches
3.5-12 mm thick, minutely (0.2 mm) appressed-puber- ulent and glabrescent, flattened at first but becoming terete; stipules 1-4 cm long, 6-12 mm broad at the base, narrowly triangular and acute to acuminate, minutely appressed-puberulent or glabrous distally, slightly (1 mm) united (intrapetiolar) above the petioles. Leaves 2 or 3/node, petioles (5-)12-25(-50) mm long, 2-4.5 mm thick, minutely appressed-puberulent; leaf blades (15-)20-36(-60) cm long, 7-15(-23) cm broad, elliptic to elliptic-oblong or elliptic-obovate, apex obtuse to acu- minate, base gradually narrowed and obtuse or acute, slightly decurrent on the petiole, drying chartaceous, dark brown, glabrous and slightly lustrous above, with short (0.3 mm) thin appressed hairs beneath, 2° veins 13-207 side. Inflorescences terminal or axillary, 20-80 cm long, racemiform, primary peduncles to 10 cm long and ca. 4 mm thick, appressed-puberulent, the broad cymose flower clusters 1-5 cm distant and opposite or subopposite along the central rachis, to 2 cm long on short (4-15 mm) lateral peduncles, subtended by broad-based bracts to 10 mm long, mostly glabrous and drying black, each flower cluster with 5-30 flowers and 1 greatly enlarged red sepal lobe, the enlarged sepal lobes with slender pet- iole-like stipes 12-38 mm long and blades 3-10 cm long and 1-4 cm broad, ellipsoid to rhombic, flowers mostly sessile. Flowers with hypanthium 1-2 mm long, obconic and densely sericeous with ascending yellowish hairs, calyx tube ca. 1 mm long, calyx lobes 0.3-1 mm long and 1-1.3 mm broad, rounded distally, glabrous (1 lobe per lateral inflorescence branch greatly expanded; see above); corolla yellow-orange or orange and red, tube 3- 5 mm long, 1.5-2 mm diam., glabrous externally except for 4 longitudinal lines of appressed hairs, lobes 2-4 mm long and 1.5 mm broad, acute, anthers 1.5-2 mm long. Fruits 3-5 mm long, 3-4 mm diam., obconic to subglo- bose, appressed-puberulent; seeds 0.2-0.5 mm long.
Plants of evergreen lowland rain forest forma- tions, from 10 to 300 m elevation. Growing in forest light-gaps, forest edges, and secondary growth. Flowering throughout the year, but the principal flowering period appears to be in April- September. The species ranges from southern Nic- aragua to Peru and Bolivia.
Warszewiczia coccinea is readily identified by its brilliant and elongate racemiform inflores- cences with each opposing flower cluster bearing a single brilliant crimson leaf-like calyx lobe. Dis- tal branches often bear three inflorescences: a ter- minal one and two laterals. This is one of Costa Rica's most showy trees, but it is not common. Breeding biology was studied by Bawa and Beach (1983). Also referred to as bandera, curcero, guna, and sangrenaria.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
323
Literature Cited
ADAMS, C. D. 1 972. Flowering Plants of Jamaica. Uni- versity of the West Indies, Mona, Jamaica, 848 pp.
ALLEN, P. H. 1956. The Rain Forests of Golfo Dulce. University of Florida Press, Gainesville, 4 1 7 pp.
BAWA, K. S., AND J. H. BEACH. 1983. Self-incompat- ibility systems in the Rubiaceae of a tropical lowland wet forest. American Journal of Botany, 70: 1281- 1288.
CORRELL, D. S., AND H. B. CORRELL. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz, 1 ,692 pp.
CROAT, T. 1 978. Flora of Barro Colorado Island. Stan- ford University Press, Stanford, California, 943 pp.
DWYER, J. D. 1980. Rubiaceae. In Woodson, R. E., and R. Schery, eds., Flora of Panama. Annals of the Missouri Botanical Garden, 67: 1-522.
HABER, W. A., AND G. W. FRANKIE. 1989. A tropical hawkmoth community: Costa Rican dry forest Sphin- gidae. Biotropica, 21: 155-172.
HORTUS THIRD. 1976. A Concise Dictionary of Plants Cultivated in the United States and Canada. Compiled by staff of the Bailey Hortium, Cornell University. Macmillan, New York, 1,290 pp.
LERSTEN, N. R. 1974. Morphology and distribution of colleters and crystals in relation to the taxonomy and bacterial leaf nodule symbiosis of Psychotria (Rubi- aceae). American Journal of Botany, 61: 973-981.
MABBERLEY, D. J. 1987. The Plant Book. A Portable Dictionary of the Higher Plants. Cambridge Univer- sity Press, Cambridge, 706 pp.
PEMBERTON, R. W., AND C. E. TURNER. 1989. Occur- rence of predatory and fungivorous mites in leaf domatia. American Journal of Botany, 76: 105-1 12.
ROBBRECHT, E. 1 988. Tropical woody Rubiaceae. Ope- ra Botanica Belgica, 1: 1-271.
SANCHEZ V., P. E. 1983. Florula del Parque Nacional Cahuita. Edit. Universidad Estatal a Distancia, San Jose, 377 pp.
STANDLEY, P. C. 1938. Rubiaceae. In Flora of Costa Rica. Field Museum of Natural History Publications, Botanical Series, 18: 1264-1380.
STANDLEY, P. C., AND L. O. WILLIAMS. 1975. Rubi- aceae. In Flora of Guatemala. Fieldiana, Botany, 24(pt. XI): 1-274.
STEYERMARK, J. A. 1972. Rubiaceae. In Maguire, B., ed., The Botany of the Guyana Highlands, part IX. Memoirs of the New York Botanical Garden, 23: 227- 832.
. 1974. Rubiaceae. In Lasser, T., ed., Flora de
Venezuela, 9: 1-2070. Institute Botanico, Direccion de Recursos Naturales Renovables, Ministerio de Agricultura y Cria, Caracas, Venezuela.
TAYLOR, C. M. 1 99 1 . Rubiaceae. In Wilbur, R. L., ed., The Vascular Flora of the La Selva Biological Station, Costa Rica. Selbyana, 12: 141-190.
TAYLOR, C. M., B. E. HAMMEL, AND W. C. BURGER. 1991. New species, combinations and records in Ru- biaceae from the La Selva Biological Station, Costa Rica. Selbyana, 12: 134-140.
VERDCOURT, B. 1976. Rubiaceae (part 1). In Polhill, R. M., ed., Flora of Tropical East Africa. Crown Agents for Overseas Governments, London, 414 pp.
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List of Accepted Species
Key: ! = new species (15); # = new combinations or new names (12); CR = endemic to continental Costa Rica (68); (*) = endemic to Cocos Island; ** = endemic to Costa Rica and western Panama (21); CULT = introduced ornamental and useful species (11); (N) = in adjacent Nicaragua, not recorded for Costa Rica; (P) = in adjacent Panama, not recorded for Costa Rica. Total = 406 species in 84 genera.
Alibertia edulis Alibertia garapatica Allenanthus erythrocarpa Alseis sp. aff. A. hondurensis CR Amaioua corymbosa Amaioua pedicel lat a Amphidasya ambigua Appunia guatemalensis Arcytophyllum lavarum ** Arcytophyllum muticum
Bathysa veraguensis
Bertiera angustifolia
Bertiera guianensis
Borojoa atlantica
Borojoa panamensis (sensu laid)
Bouvardia glabra CULT
Calycophyllum candidissimum Chimarrhis latifolia ** Chimarrhis parviflora Chiococca alba Chiococca pachyphylla Chiococca phaenostemon Chiococca semipilosa Chione sylvicola Chomelia microloba Chomelia recordii Chomelia spinosa Chomelia venulosa CR! Chomelia sp. A, aff. C. spinosa Cinchona pubescens Coccocypselum cordifolium Coccocypselum guianense Coccocypselum herbaceum Coccocypselum hirsutum Coccocypselum lanceolatum Coffea arabica CULT Coffea liberica CULT Condaminea corymbosa Cosmibuena grandiflora Cosmibuena macrocarpa Cosmibuena valerii Coussarea austin-smithii CR Coussarea caroliana CR Coussarea chiriquiensis **# Coussarea enneantha Coussarea hondensis Coussarea impetiolaris CR Coussarea latifolia Coussarea nebulosa Coussarea nigrescens (P) Coussarea psychotrioides CR Coussarea talamancana ** Coussarea sp. A, aff. C. curvigemma Coussarea sp. B, aff. C. curvigemma Coutarea hexandra Crusea coccinea
Crusea hispida Crusea longiflora Crusea parviflora
Dedieuxia fruticosa Deppea grandiflora Didymaea alsinoides Diodia apiculata Diodia brasiliensis Diodia sarmentosa Diodia serrulata Diodia teres Duroia costaricensis Duroia utleyorum CR!
Elaeagia auriculata Elaeagia myriantha Elaeagia nitidifolia Elaeagia uxpanapensis Exostema caribaeum Exostema mexicanum
Faramea capulifolia **! Faramea eurycarpa Faramea hondurae Faramea multiflora Faramea myrticifolia CR! Faramea occidentalis Faramea ovalis Faramea parvibractea Faramea pauciflora Faramea scalaris (P) Faramea sessifolia Faramea stenura CR Faramea suerrensis Faramea trinervia ** Ferdinandusa panamensis **
Galium aschenbornii Galium hypocarpium Galium mexicanum Galium orizabense Galium uncinulatum Gardenia augusta CULT Genipa americana Genipa williamsii Geophila cordifolia Geophila gracilis (P) Geophila macropoda Geophila repens Gonzalagunia bracteosa CR Gonzalagunia brenesii Gonzalagunia ovatifolia Gonzalagunia panamensis Gonzalagunia rosea Gonzalagunia rudis Gonzalagunia stenostachya CR# Guettarda brenesii CR
Guettarda combsii (N) Guettarda conferta Guettarda crispiflora Guettarda foliacea Guettarda macrosperma Guettarda poasana CR Guettarda sanblasensis Guettarda tournefortiopsis Guettarda turrialbana CR
Hamelia axillaris Hamelia calycosa (P) Hamelia macrantha Hamelia magnifolia ** Hamelia patens Hamelia rovirosae Hamelia xerocarpa Hillia allenii Hillia grayumii CR Hillia longifilamentosa ** Hillia loranthoides Hillia macrophylla Hillia maxonii Hillia panamensis Hillia triflora Hippotis albiflora Hoffmannia aeruginosa Hoffmannia amplexifolia Hoffmannia arborescens CR Hoffmannia areolata ** Hoffmannia asclepiadea Hoffmannia bullata Hoffmannia congesta # Hoffmannia davidsoniae ** Hoffmannia decurrens CR Hoffmannia dotae ** Hoffmannia hamelioides CR Hoffmannia hammelii CR! Hoffmannia inamoena CR Hoffmannia laxa Hoffmannia leucocarpa CR Hoffmannia liesneriana CR Hoffmannia longipetiolata Hoffmannia nesiota (*) Hoffmannia pallidiflora Hoffmannia piratarum (*) Hoffmannia pittieri ** Hoffmannia psychotriifolia Hoffmannia subauriculata Hoffmannia valerii Hoffmannia vesiculifera
Isertia haenkeana
Isertia laevis
Ixora coccinea CULT
Ixorafinlaysoniana CULT
Ixora floribunda
Ixora nicaraguensis
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
325
Ladenbergia brenesii CR Ladenbergia sericophylla CR Ladenbergia valerii CR Lasianthus panamensis Lindenia rivalis
Machaonia acuminata (P) Machaonia martinicensis Macrocnemum glabrescens Malanea colombiana Manettia barbata Manet tia flexilis Manettia reclinata Mitracarpus hirtus Morinda citrifolia Morinda panamensis Morinda royoc (N) Mussaenda erythrophylla CULT
Neolamarckia cadamba CULT Nertera granadensis
Oldenlandia callitrichoides Oldenlandia corymbosa Oldenlandia herbacea Oldenlandia lancifolia Osa pulchra CR
Palicourea adusta CR Palicourea albocaerulea CR Palicourea angustifolia Palicourea bella Palicourea bellula CR Palicourea brenesii CR Palicourea copensis Palicourea crocea Palicourea discolor ** Palicourea garciae Palicourea guianensis Palicourea hammelii Palicourea lancifera CR Palicourea lasiorrhachis Palicourea macrocalyx CR Palicourea montivaga CR Palicourea orosiana CR Palicourea padifolia Palicourea purpurea ** Palicourea salicifolia CR Palicourea skotakii CR Palicourea spathacea CR Palicourea standleyana Palicourea tilaranensis CR Palicourea triphylla Palicourea vestita Pentagonia costaricensis **# Pentagonia donnell-smithii Pentagonia gymnopoda Pentagonia hirsuta CR Pentagonia macrophylla Pentagonia tinajita Pentagonia wendlandii Pent as lanceolata CULT Pentodon pentandrus Pogonopus speciosus Posoqueria coriaceae Posoqueria grandiflora CR
Posoqueria latifolia Psychotria acicularis CR! Psychotria acuminata Psychotria aggregata Psychotria alfaroana ** Psychotria allenii ** Psychotria angustiflora Psychotria aubletiana Psychotria aurantibractea CR# Psychotria berteriana Psychotria boquetensis Psychotria borucana # Psychotria brachiata Psychotria brachybotrya Psychotria calochlamys Psychotria camponutans Psychotria capacifolia Psychotria capitata (N) Psychotria cartagoensis CR Psychotria carthagenensis Psychotria chagrensis Psychotria chiapensis Psychotria chiriquiensis **# Psychotria chiriquina ** Psychotria chitariana CR Psychotria cincta Psychotria clivorum Psychotria cocosensis (*) Psychotria cooperi Psychotria correae # Psychotria deflexa Psychotria dichroa # Psychotria domingensis Psychotria dukei Psychotria elata Psychotria emetica Psychotria erecta Psychotria eurycarpa CR Psychotria fruticetorum Psychotria glomerulata Psychotria goldmanii Psychotria graciliflora Psychotria grandis Psychotria guadalupensis Psychotria guapilensis Psychotria haematocarpa Psychotria hazenii Psychotria hebeclada Psychotria hoffmannseggiana Psychotria horizontalis Psychotria insignis (?) Psychotria ipecacuanha Psychotria jimenezii CR Psychotria lamarinensis CR Psychotria laselvensis CR Psychotria limonensis Psychotria longipedunculoides # Psychotria luxurians Psychotria macrophylla Psychotria marginata Psychotria maxonii Psychotria mexiae Psychotria micrantha Psychotria microbotrys Psychotria microdon Psychotria monteverdensis CR
Psychotria mortoniana ** Psychotria neillii Psychotria nervosa Psychotria officinalis Psychotria orosiana Psychotria panamensis Psychotria parvifolia Psychotria phanaerandra Psychotria pilosa Psychotria pithecobia Psychotria pittieri Psychotria platypoda Psychotria poeppigiana Psychotria polyphlebia Psychotria psychotriifolia Psychotria pubescens Psychotria quinqueradiata Psychotria racemosa Psychotria remota Psychotria rosulatifolia Psychotria sarapiquensis Psychotria siggersiana Psychotria sixaolensis ** Psychotria solitudinum Psychotria steyermarkii Psychotria stockwellii ** Psychotria suerrensis Psychotria sylvivaga ** Psychotria tapantiensis CR! Psychotria tenuifolia Psychotria turrubarensis CR Psychotria uliginosa Psychotria umbelliformis Psychotria valeriana CR Psychotria viridis Psychotria sp. A (Cocos Island, in-
det.) Psychotria sp. B, aff. P. papantlense
Randia aculeata Randia altiscandens # Randia armata Randia brenesii Randia calycosa Randia genipoides CR! Randia gentryi Randia grandifolia Randia grayumii ! Randia loniceroides Randia matudae Randia mira Randia monantha Randia pittieri Randia pterocarpa Randia retroflexa Randia thurberi Randia vazquezii Raritebe palicoureoides Richardia brasiliensis Richardia scabra Rondeletia amoena Rondeletia aspera CR Rondeletia bertierioides Rondeletia brenesii CR Rondeletia buddleioides Rondeletia calycosa CR
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FIELDIANA: BOTANY
Rondeletia chaconii CR! Rondeletia cooperi Rondeletia costaricensis CR Rondeletia hamelifolia Rondeletia monteverdensis CR Rondeletia povedae CR! Rondeletia salicifolia Rondeletia stenostachya Rondeletia tayloriae CR! Rondeletia torresii CR Rondeletia urophylla CR Rudgea amplexicaulis Rudgea chiriquiensis (P) Rudgea cornifolia Rudgea monofructus CR Rudgea raveniana CR! Rudgea reducticalyx CR! Rudgea skutchii
Rudgea trifurcata CR Rustia costaricensis # Rustia Occident alls
Sabicea panamensis Sabicea villosa Schradera blumii (P) Schradera costaricensis CR! Serissa japonica CULT Sherardia arvensis Simira maxonii Sipanea biflora Sommera donnell-smithii Spermacoce assurgens Spermacoce confusa Spermacoce densiflora Spermacoce exilis Spermacoce latifolia
Spermacoce ocymifolia Spermacoce ovalifolia Spermacoce prostrata Spermacoce scabiosoides Spermacoce suaveolens Spermacoce tenuior Spermacoce vegeta Spermacoce verticillata
Tocoyena pittieri
Uncaria tomentosa
Vangueria madagascariensis CULT
Warszewiczia coccinea
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
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Index
The index includes all accepted names (in Roman type), synonyms (italics), common English names (Roman), and vernacular Spanish names (italics). New species, new names, and newly transferred names are in boldface, and the page numbers of illustrations are in boldface. Hyphenated words and multiple words are alphabetized by letter.
Alibertia 82 Alibertia acuminate 82 Alibertia edulis 39, 82 Alibertia garapatica 83 Alibertia latifolia 82 Allenanthus 83 Allenanthus erythrocarpa 83 Allenanthus hondurensis 83 Alseis 84
Alseis blackiana 85 Alseis sp. aff. hondurensis 54, 84 Amaioua 85 Amaioua corymbosa 85 Amaioua pedicellata 44, 86 Amphydasya 86 Amphydasya ambigua 21, 86 anileto 160 Anisomeris 99 Anisomeris englesingii 100 Anisomeris microloba 100 Anisomeris recordii 100 Anisomeris sylvicola 99 Anthocephalus 195 Anthocephalus cadamba 195 Anthocephalus chinensis 195 Anthocephalus indicus 195 Anthocephalus morindaefolius 195 Antirhea panamensis 100 Appunia 87
Appunia guatemalensis 33, 87 Appunia seibertii 88 aquijilla 184 Arachnothrix 292 Arachnothrix stenostachya 1 50 Arcytophyllum 88 Arcytophyllum chirropoense 88 Arcytophyllum lavarum 15, 88 Arcytophyllum muticum 15, 89 Arcytophyllum recurvatum 89 azulillo 160
bandera 323
Basanacantha altiscandens 282
Basanacantha grandifolia 285
Basanacantha monantha 287
Basanacantha pittieri 288
Bassanacanth subcordata 287
Bathysa 89
Bathysa veraguensis 89
Bertiera 90
Bertiera angustifolia 58, 90
Bertiera guianensis 58, 91
Bertirea panamensis 290
boca de vieja 220
Borojoa 91
Borojoa atlantica 40, 92 Borojoa panamensis 40, 92 Borreria 92
Borreria densiflora 3 1 6 Borreria exilis 316 Borreria gracilis 316 Borreria laevis 315 Borreria latifolia 316 Borreria ocymoides 318 Borreria ovalifolia 3 1 7 Borreria parviflora 3 1 8 Borreria repens 3 1 8 Borreria scabiosoides 318 Borreria spinosa 316 Borreria suaveolens 319 Borreria vegeta 320 Borreria verticillata 320 Bouvardia 92 Bouvardia glabra 93 Bouvardia longiflora 93 Breonia 195 Buena macrocarpa 109 Buena panamensis 149 Buena skinneri 109
cachitos 284
cafe 105, 107
cafe macho 139
cafecillo 139
Callicocca ipecacuanha 255
Calycophyllum 93
Calycophyllum candidissimum
30,93
Calycophyllum coccineum 323 caribee bark tree 1 30 carica 220 caruto 144
Casasia chiapensis 286 Cassupa laevis 1 8 1 Cassupa panamensis 1 8 1 Cephaelis 94 Cephaelis afflnis 238 Cephaelis axillaris 234 Cephaelis borucana 236 Cephaelis camponutans 238 Cephaelis chiriquiensis 24 1 Cephaelis chlorochlamys 252 Cephaelis correae 244 Cephaelis costaricensis 246 Cephaelis dichroa 245 Cephaelis discolor 25 1 Cephaelis elata 246 Cephaelis elata forma lutea 246
Cephaelis glomerulata 249 Cephaelis guapilensis 25 1 Cephaelis hirsuta 268 Cephaelis ipecacuanha 255 Cephaelis latistipula 239 Cephaelis nana 25 1 Cephaelis nicaraguensis 25 1 Cephaelis pittieri 235 Cephaelis psychotriaefolia 269 Cephaelis tetragona 241 Cephaelis tomentosa 268 Cephaelis tonduzii 251 Cephalanthus chinensis 195 Cephalanthus occidentalis 322 Chimarrhis 94 Chimarrhis cymosa 95 Chimarrhis decurrens 83 Chimarrhis latifolia 51, 95 Chimarrhis parviflora 51, 95 Chiococca 96 Chiococca alba 50, 96 Chiococca jefensis 1 29 Chiococca pachyphylla 50, 97 Chiococca phaenostemon 50, 97 Chiococca semipilosa 50, 98 Chione 98 Chione allenii 99 Chione chambersii 100 Chione costaricensis 99 Chione panamensis 99 Chione sylvicola 50, 98 chiritillo3\l Chomelia 99 Chomelia coclensis 189 Chomelia englesingii 100 Chomelia filipes 101 Chomelia microloba 47,48, 100 Chomelia panamensis 100 Chomelia peninsularis 100 Chomelia recordii 48, 100 Chomelia spinosa 48, 107 Chomelia sylvicola 98, 99 Chomelia venulosa 48, 101 Cinchona 102 Cinchona calisaya 102 Cinchona caribaeum 1 30 Cinchona grandiflora 1 09 Cinchona pubescens 51, 102 Coccocypselum 103 Coccocypselum canescens 105 Coccocypselum cordifolium 16, 104 Coccocypselum glabrum 105 Coccocypselum herbaceum 104 Coccocypselum hirsutum 16, 105 Coccocypselum hirsutum var. gla- brum 105
328
FIELDIANA: BOTANY
Coccocypselum hispidulum 104 Coccocypselum guianense 104 Coccocypselum lanceolatum 16, 105 Coccocypselum pleuropodum 1 04 Coccocypselum repens 104, 105 Coccocypselum rothschuii 104 Coffea 106 CofTea arabica 106 Coffea canephora 106 Coffea excelsa 107 Coffea liberica 107 Coffea robusta 106 colleters 1
Condalia lanceolata 105 Condaminea 107 Condaminea corymbosa 43, 107 coralillo 160, 283 corona de la reina 1 82 Cosmibuena 182 Cosmibuena grandiflora 42, 109 Cosmibuena macrocarpa 42, 109 Cosmibuena ovalis 109 Cosmibuena paludicola 109 Cosmibuena skinneri 1 09 Cosmibuena valerii 41, 1 10 Coussarea 1 10
Coussarea austin-smithii 62, 112 Coussarea caroliana 62, 112 Coussarea chiriquiensis 290 Coussarea colonis 290 Coussarea curvigemma 1 1 7 Coussarea enneantha 61, 113 Coussarea euryphylla 290 Coussarea hondensis 63, 113 Coussarea impetiolaris 61, 114 Coussarea jimenezii 114 Coussarea latifolia 63, 114 Coussarea nebulosa 1 1 5 Coussarea nigrescens 1 1 5 Coussarea psychotrioides 62, 115 Coussarea sp. A, aff. curvigemma
62, 116 Coussarea sp. B, aff. curvigemma 63,
117
Coussarea talamancana 63, 116 Coussarea veraguensis 112 Coutarea 1 1 7
Coutarea hexandra 45, 117 Coutarea speciosa 1 1 7 Crucianella hispida 1 1 9 crucilla 288 crucillo 283 crucito 283 Crusea 1 1 8
Crusea brachyphylla 1 1 9 Crusea coccinea 45, 119 Crusea coccinea var. chiriquiensis
119
Crusea hispida 119 Crusea longiflora 119 Crusea parviflora 18, 120 cruz de Malta 182 curcero 323
Declieuxia 120
Declieuxia fruticosa 15, 121
Declieuxia fruticosa var. mexicana
121
Declieuxia mexicana 121 Deppea 121
Deppea costaricensis 1 2 1 Deppea floribunda 121 Deppea grandiflora 52, 121 Deppea longipes 1 2 1 Deppea panamensis 128 Didymaea 122 Didymaea alsinoides 17, 122 Didymaea alsinoides var. australis
122
Didymaea alsinoides var. mollis 122 Didymaea australis 1 22 Didymochlamys 3 Diodia 123
Diodia apiculata 20, 124 Diodia brasiliensis 15, 124 Diodia brasiliensis var. angulata 1 24 Diodia maritima 125 Diodia ocymifolia 3 1 7 Diodia polymorpha 1 24 Diodia prostrata 125 Diodia rigida 1 24 Diodia sarmentosa 1 24 Diodia serrulata 20, 125 Diodia teres 20, 125 distylous 1 domatia 1
Dressleriopsis panamensis 185 Duggenea rudis 150 Dukea 290
Dukea euryphylla 290 Duroia 126
Duroia costaricensis 39, 126 Duroia genipifolia 1 26 Duroia hirsuta 126 Duroia panamensis 165 Duroia petiolaris 1 26 Duroia saccifera 126 Duroia ut ley or 11 m 127
Elaeagia 127
Elaeagia auriculata 53, 128
Elaeagia karstenii 128
Elaeagia myrianthes 1 28
Elaeagia nitidifolia 53, 129
Elaeagia utilis 127
Elaeagia uxpanapensis 53, 129
espino bianco 282
Evea dichroa 245
Evea elata 246
Evea guapilensis 25 1
Evea tomentosa 268
Exostema 130
Exostema caribaeum 45, 130
Exostema longicuspe 1 36
Exostema mexicanum 54, 1 30
Exostema occidentale 307
Faramea 131 Faramea belizensis 135 Faramea bocaturensis 123
Faramea bullata 138 Faramea capulifolia 1 22 Faramea cobana 1 34 Faramea eurycarpa 60, 123 Faramea hondurae 60, 123 Faramea luteovirens 1 36 Faramea multiflora 60, 134 Faramea myrticifolia 1 34 Faramea occidentalis 59, 134 Faramea ovalis 60, 135 Faramea parvibractea 60, 135 Faramea pa uciflora 136 Faramea quercetorum 135 Faramea scalaris 1 36 Faramea sessifolia 59, 137 Faramea sessilifolia 137 Faramea standleyana 135 Faramea stenura 60, 137 Faramea suerrensis 59, 137 Faramea talamancarum 1 34 Faramea trinervia 59, 138 Faramea trinervis var. suerrensis 137 Faramea zeteki 1 34 Ferdinandusa 138 Ferdinandusa panamensis 54, 138 flor de cruz 283 flordefuego 182 fruta de mono 220
Galium 139
Galium aschenbornii 17, 140 Galium hypocarpium 17, 140 Galium mexicanum 17, 141 Galium obovatum 142 Galium orizabense 17, 141 Galium uncinulatum 17, 141 gardenia 142 Gardenia armata 283 Gardenia augusta 142 Gardenia edulis 82 Gardenia florida 142 Gardenia jasminoides 1 42 Genipa 142
Genipa americana 39, 40, 143 Genipa americana var. caruto 143 Genipa caruto 143 Genipa codonocalyx 143 Genipa edulis 82 Genipa maxonii 3 1 1 Genipa oblongifolia 143 Genipa venosa 143 Genipa vulcanicola 143, 287 Genipa williamsii 40, 144 Geocardia pleuropoda 104 Geophila 144
Geophila cordifolia 16, 145 Geophila croatii 145 Geophila gracilis 145 Geophila herbacea 1 46 Geophila macropoda 16, 146 Geophila pleuropoda 104 Geophila repens 16, 146 Geophila trichogyne 145 Glossostipula concinna 143 Gomozia granadensis 195 Gonzalagunia 147
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
329
Gonzalagunia bracteosa 35, 148 Gonzalagunia brenesii 34, 148 Gonzalagunia exaltata 295 Gonzalagunia kallunkii 1 5 1 Gonzalagunia longithyrsa 149 Gonzalagunia ovatifolia 34, 148 Gonzalagunia panamensis 34, 149 Gonzalagunia rosea 34, 149 Gonzalagunia rudis 1 50 Gonzalagunia stenostachya 1 50 Gonzalagunia veraguensis 1 5 1 Gonzalea bracteosa 148 Gonzalea ovatifolium 149 Gonzalea panamensis 149 guatil 144
guayabo de mono 220 guayaba mica 220 Guettarda 151 Guettarda brenesii 47, 152 Guettarda chiriquiensis 153 Guettarda combsii 152 Guettarda conferta 153 Guettarda costaricensis 101 Guettarda crispiflora 46, 153 Guettarda divaricata 1 54 Guettarda foliosa 47, 153 Guettarda macrosperma 47, 154 Guettarda poasana 46, 154 Guettarda sanblasensis 47, 155 Guettarda tournefortiopsis 46, 155 Guettarda turrialbana 44, 156 guna 323
Hamelia 156
Hamelia subgenus Amphituba 157
Hamelia axillaris 57, 157
Hamelia calycosa 158
Hamelia costaricensis 1 60
Hamelia lutea 157
Hamelia macrantha 158
Hamelia magnifolid 57, 159
Hamelia panamensis 1 60
Hamelia patens 57, 159
Hamelia patens var. glabra 1 59
Hamelia rovirosae 48, 160
Hamelia rowlei 160
Hamelia storkii 160
Hamelia viridifolia 159
Hamelia xerocarpa 57, 160
Hamelia xerocarpa var. costaricen- sis 161
Hamelia xerocarpa var. xerocarpa 161
Hedyotis halei 217
Hedyotis herbacea 197
Hedyotis lancifolia 198
Hedyotis mutica 89
Hedyotis pentandra 2 1 7
Hemidiodia ocymifolia 3 1 7
heterotopy 203, 292
Higginsia psychotriaefolia 178
Hillia 161
Hillia allenii 162
Hillia chiapensis 164
Hillia chiriquiensis 1 10
Hillia grayumii 42, 162
Hillia hathewayi 163 Hillia ligulifolia 110 Hillia longifilamentosa 1 62 Hillia loranthoides 41, 163 Hillia macrophylla 42, 163 Hillia maxonii 41, 163 Hillia palmana 163 Hillia panamensis 41, 164 Hillia tetrandra 1 64 Hillia triflora 41, 164 Hillia triflora var. pittieri 164, 165 Hillia triflora var. triflora 165 Hillia valerii 110 Hintonia pulchra 198 Hippo tis 165 Hippo tis albiflora 39, 165 Hoffmannia 166 Hoffmannia aeruginosa 23, 169 Hoffmannia affinis 1 69 Hoffmannia amplexifolia 22, 169 Hoffmannia arborescens 24, 170 Hoffmannia areolata 23, 170 Hoffmannia asclepiadea 24, 1 70 Hoffmannia boraginoides 179 Hoffmannia bullata 23, 171 Hoffmannia capillacea 175 Hoffmannia carpinterae 175 Hoffmannia congesta 23, 171 Hoffmannia davidsoniae 21, 172 Hoffmannia decurrens 1 72 Hoffmannia discolor 1 7 1 Hoffmannia dotae 24, 173 Hoffmannia eliasii 1 76 Hoffmannia fimbrianther a 174 Hoffmannia gesnerioides 173 Hoffmannia hamelioides 25, 173 Hoffmannia hammelii 1 74 Hoffmannia haydenii 178 Hoffmannia inamoena 25, 174 Hoffmannia josefina 170 Hoffmannia kirkbridei 179 Hoffmannia lancistigma 1 72 Hoffmannia laxa 25, 174 Hoffmannia leucocarpa 24, 175 Hoffmannia liesneriana 22, 175 Hoffmannia longipetiolata 25, 176 Hoffmannia macrophylla 175 Hoffmannia morii 179 Hoffmannia nesiota 1 76 Hoffmannia nicotianaefolia 176 Hoffmannia pallidiflora 25, 177 Hoffmannia panamensis 111 Hoffmannia piratarum 177 Hoffmannia pittieri 177 Hoffmannia psychotriifolia 25, 178 Hoffmannia ramonensis 173 Hoffmannia refulgens 178 Hoffmannia rexmontis 111 Hoffmannia steinworthii 1 70 Hoffmannia subauriculata 22, 178 Hoffmannia tonduzii 176 Hoffmannia valerii 23, 178 Hoffmannia vesiculifera 22, 179 Hoffmannia woodsonii 176 Holtonia 179 Holtonia myriantha 128 horquetilla de montana 284
Houstonia 88 Houstonia fruticosa 1 2 1
ipecac, ipecacuana 255
Isertia 179
Isertia sect. Cassupa 1 80
Isertia deamii 1 80
Isertia deamii var. stenophylla 1 80
Isertia haenkeana 65, 180
Isertia hypoleuca 181
Isertia laevis 44, 181
Isertia panamensis 1 8 1
Ixora 181
Ixora coccinea 45, 182
Ixora fmlaysoniana 45, 182
Ixora floribunda 58, 182
Ixora nicaraguensis 58, 183
Ixora occidentalis 1 34
Ixora rauwolfioides 183
Ixora spinosa 101
jagua 144 jagua amarillo 95 jagua macho 283 jagua negro 144 Javorkia 292 jazmin 93, 142 jazmin de coral 1 82 jazmin de la virgen 93 jazmin del cabo 142 jazmin del volcdn 164 jazmin rojo 182 jazmincillo 187 jicarillo 220 Joosia 5
kadam, kedam 195
Ladenbergia 183
Ladenbergia brenesii 43, 184
Ladenbergia sericophylla 43, 184
Ladenbergia valerii 43, 185
lagartillo 82
Lagenanthus parviflorus 1 64
lagrimas de Maria 97
lagrimas de San Pedro 97
laran 195
Lasianthus 185
Lasianthus panamensis 21, 185
Lasionema glabrescens 188
lechuga 147
limoncillo 101, 283
Lindenia 186
Lindenia rivalis 29, 1 86
lirio de agua 187
Lonicera alba 96
Loranthus guadalupensis 250
Machaonia 187 Machaonia acuminata 187 Machaonia martinicensis 51, 188
330
FIELDIANA: BOTANY
Machaonia rotundata 188 Machaonia rotundata var. dodgei
188
Macrocnemum 188 Macrocnemum candidissimum 93 Macrocnemum coccineum 323 Macrocnemum corymbosum 107 Macrocnemum exsertum 218 Macrocnemum glabrescens 54, 188 Macrocnemum speciosus 2 1 8 madrono 94 malacaguite 101 malacahuite 101, 154 Malanea 189 Malanea colombiana 1 89 Malanea erecta 1 89 Mallostoma lavarum 88 Manettia 190 Manettia barbata 15, 190 Manettia bocaturensis 198 Manettia coccinea 191 Manettia costaricensis 191 Manettia cuspidata 1 9 1 Manettia estrellae 191 Manettia flexilis 15, 191 Manettia hydrophila 3 1 2 Manettia inflata 190 Manettia luteo-rubra var. paraguar-
iensis 190
Manettia orbifera 1 9 1 Manettia panamensis 1 9 1 Manettia reclinata 15, 191 Manettia seleriana 1 9 1 Manettia stenophylla 190 manzano de mico 220 Mapouria obovata 269 Mapouria remota 27 1 Mapouria trichogyne 145 medicinal uses 102, 118, 130, 247,
255
miel quema 283 mielcillo 3 1 7 Mitracarpus 191 Mitracarpus breviflorus 1 92 Mitracarpus hirtus 18, 192 Mitracarpus villosus 192 mono de reina 1 82 Montamans panamensis 233 Morinda 192 Morinda citrifolia 33, 193 Morinda guatemalensis 87 Morinda panamensis 33, 193 Morinda royoc 33, 194 mostrenco 283 Mussaenda 194 Mussaenda coccinea 323 Mussaenda erythrophylla 30, 194 Mussaenda spinosa 283
Nacibea coccinea 1 9 1 Nauclea aculeata 322 Nauclea cadamba 195 Nauclea tomentosa 322 Neolamarckia 195 Neolamarckia cadamba 195
Nertera 195 Nertera alsinoides 122 Nertera depressa 195 Nertera granadensis 17, 195 Nonatelia officinalis 263 Nonatelia racemosa 270 Nothophlebia costaricensis 2 1 3
Oldenlandia 196
Oldenlandia callitrichoides 197
Oldenlandia corymbosa 17, 197
Oldenlandia herbacea 197
Oldenlandia lancifolia 198
Oldenlandia pentandra 217
Oldenlandiodes callitrichoides 173
Ophrycoccus gesnerioides 173
Ophiorhiza lanceolata 217
Oregandra 98
Oregandra panamensis 98, 99
Osa 198
Osa pulchra 29, 198
Palicourea 199 Palicourea adusta 68, 2 1 1 Palicourea albocaerulea 68, 202 Palicourea angustifolia 202 Palicourea austin-smithii 209 Palicourea bella 202 Palicourea bellula 64, 203 Palicourea brenesii 67, 203 Palicourea caerulescens 209 Palicourea copensis 67, 204 Palicourea costaricensis 208 Palicourea crocea 65, 204 Palicourea discolor 67, 205 Palicourea domingensis 245 Palicourea galeottiana 209 Palicourea garciae 68, 205 Palicourea guianensis 65, 205 Palicourea hammelii 206 Palicourea lanceolata 202 Palicourea lancifera 68, 206 Palicourea lasiorhachis 66, 207 Palicourea leucantha 207 Palicourea longibracteata 2 1 2 Palicourea macrocalyx 64, 207 Palicourea macrosepala 205 Palicourea molliramis 253 Palicourea montivaga 68, 208 Palicourea nigrescens 259 Palicourea ochnoides 203 Palicourea orosiana 66, 203 Palicourea padifolia 66, 208 Palicourea panamensis 205 Palicourea parviflora 212 Palicourea phanderandra 265 Palicourea purpurea 67, 209 Palicourea salicifolia 68, 209 Palicourea skotakii 64, 210 Palicourea spathacea 64, 210 Palicourea standleyana 69, 2 1 1 Palicourea subruba 209 Palicourea talamancana 203 Palicourea thyrsiflora 209
Palicourea tilarensis 64, 211 Palicourea triphylla 65, 211 Palicourea veraguensis 207 Palicourea vestita 66, 212 pah comaron 1 60 palo cuadrado 1 89 palo de cruz 283 palo de Maria 183 Pentagonia 212 Pentagonia alfaroana 2 1 6 Pentagonia costaricensis 28, 213 Pentagonia donnell-smithii 28, 214 Pentagonia gymnopoda 214 Pentagonia hirsuta 2 1 5 Pentagonia macrophylla 2 1 5 Pentagonia tinajita 28, 216 Pentagonia wendlandii 2 1 6 Pentas216
Pentas lanceolata 45, 217 Pentodon 2 1 7 Pentodon halei 2 1 7 Pentas lanceolata 2 1 7 Pentodon pentandrus 2 1 7 picarito 220 pili-tso 160 pin flowers 1 pissi 160 Pogonopus 2 1 8 Pogonopus exsertus 2 1 8 Pogonopus speciosus 30, 218 Portlandia hexandra 1 1 7 Posoqueria 218 Posoqueria coriacea 2 1 9 Posoqueria grandiflora 29, 219 Posoqueria latifolia 29, 220 Posoqueria maxima 2 1 9 Posoqueria obliquinervia 321 Posoqueria panamensis 220 Posoqueria pittieri 321 princewood 130 Psychotria221 Psychotria acicularis 70, 23 1 Psychotria acuminata 71, 232 Psychotria aggregata 26, 232 Psychotria alboviridula 27 1 Psychotria alfaroana 81, 233 Psychotria allenii 233 Psychotria angustiflora 72, 234 Psychotria anomothyrsa 258 Psychotria apoda 249 Psychotria aubletiana 234 Psychotria aurantibractea 32, 234 Psychotria bella 202 Psychotria berteriana 73, 235 Psychotria beteriana ssp. luxurians
257
Psychotria boquetensis 236 Psychotria borucana 31, 236 Psychotria brachiata 73, 236 Psychotria brachybotrya 237 Psychotria brenesii 2 1 1 Psychotria caerulea 237 Psychotria calochlamys 237 Psychotria calophylla 238 Psychotria camponutans 238 Psychotria capacifolia 69, 238
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
331
Psychotria capitata 239 Psychotria cartagoensis 26,27, 239 Psychotria carthagenensis 80, 240 Psychotria chagrensis 75, 240 Psychotria chiapensis 32, 241 Psychotria chiriquiensis 31, 241 Psychotria chiriquina 76, 242 Psychotria chitariana 24 1 Psychotria cincta 73, 242 Psychotria clivorum 76, 243 Psychotria cocosensis 77, 243 Psychotria compressicaulis 264 Psychotria cooperi 27, 244 Psychotria copensis 204 Psychotria copeyana 205 Psychotria cornifolia 303 Psychotria cornigera 274 Psychotria correae 31, 244 Psychotria costaricensis 266 Psychotria costavenia 250 Psychotria crocea 204 Psychotria cuspidata 232 Psychotria deflexa 71 , 244 Psychotria dichroa 245 Psychotria dispersa 267 Psychotria domingensis 245 Psychotria dukei 246 Psychotria elata 31, 246 Psychotria elongata 262 Psychotria emetica 27, 247 Psychotria erecta 27, 247 Psychotria eugeniifolia 249 Psychotria eurycarpa 62, 248 Psychotria fimbriata 303 Psychotria fruticetorum 75, 248 Psychotria furcata 253 Psychotria glauca 269 Psychotria glomerata 278 Psychotria glomerulata 32, 249 Psychotria goldmanii 70, 249 Psychotria graciliflora 75, 249 Psychotria gracilis 145 Psychotria grandis 79, 250 Psychotria grandistipula 264 Psychotria guadalupensis 75, 250 Psychotria guapilensis 21, 251 Psychotria haematocarpa 71, 252 Psychotria hazenii 70, 252 Psychotria hebeclada 70, 252 Psychotria herbacea 146 Psychotria hoffmannseggiana 71,
253
Psychotria hondensis 1 1 3 Psychotria horizontalis 80, 254 Psychotria insignis 81, 254 Psychotria involucrata 253, 263 Psychotria ipecacuanha 27, 255 Psychotria iquitosensis 237 Psychotria jimenezii 78, 255 Psychotria lamarinensis 81, 255 Psychotria laselvensis 76, 256 Psychotria limonensis 79, 256 Psychotria limonensis var. angusti-
folia 243
Psychotria longicaulis 254 Psychotria longlpedunculata 257
Psychotria longipedunculoides 257 Psychotria lupulina 275 Psychotria luteotuba 265 Psychotria luxurians 72, 257 Psychotria macrophylla 26, 258 Psychotria macrophylla var. angus-
tissima 258 Psychotria macrophylla var. an-
omothyrsa 258 Psychotria macropoda 146 Psychtoria magna 264 Psychotria marginata 76, 258 Psychotria maxonii 75, 259 Psychotria mexiae 77, 259 Psychotria mexicana 208 Psychotria micrantha 78, 260 Psychotria microbotrya 73, 260 Psychotria microdesmia 278 Psychotria microdon 74, 261 Psychotria mima 234 Psychotria molliramis 252 Psychotria mombachensis 245 Psychotria monteverdensis 76, 261 Psychotria morae 270 Psychotria morii 272 Psychotria mortoniana 69, 26 1 Psychotria neilii 78, 262 Psychotria nervosa 78, 262 Psychotria nicaraguensis 258 Psychotria obovata 270 Psychotria officinalis 74, 263 Psychotria orchidearum 250 Psychotria orosiana 76, 263 Psychotria ostaurea 1 1 3 Psychotria padifolia 218 Psychotria palicourea 205 Psychotria panamensis 77, 79, 264 Psychotria parasitica 250 Psychotria parvifolia 75, 265 Psychotria patens 244 Psychotria pavetta 245 Psychotria pendula 250 Psychotria peperomiae 250 Psychotria phanerandra 265 Psychotria pilosa 69, 266 Psychotria pithecobia 75, 266 Psychotria pittieri 70, 267 Psychotria platypoda 267 Psychotria poeppigiana 31, 268 Psychotria polyphlebia 27, 268 Psychotria psychotriifolia 78, 269 Psychotria pubescens 74, 269 Psychotria quiinifolia 262 Psychotria quinqueradiata 80, 269 Psychotria racemosa 74, 270 Psychotria ramonensis 252 Psychotria remota 81, 271 Psychotria rosulatifolia 80, 271 Psychotria rufescens 260 Psychotria sarapiquensis 77, 271 Psychotria schippii 259 Psychotria siggersiana 272 Psychotria sixaolensis 79, 272 Psychotria solitudinum 72, 273 Psychotria sp. A 279 Psychotria sp. B 279
Psychotria steyermarkii 70, 273 Psychotria stipulosa 238 Psychotria stockwellii 77, 274 Psychotria subsessilis 274 Psychotria suerrensis 32, 274 Psychotria sylvivaga 77, 275 Psychotria tapantiensis 72, 275 Psychotria tenuifolia 80, 276 Psychotria tonduzii 232 Psychotria torresiana 249 Psychotria trichotoma 276 Psychotria triphylla 2 1 1 Psychotria turrubarensis 277 Psychotria tutensis 1 1 2 Psychotria uliginosa 26, 277 Psychotria umbelliformis 62, 277 Psychotria undata 262 Psychotria valeriana 71, 278 Psychotria vallensis 249 Psychotria viridis 80, 278 Psychotria wendlandiana 255
querica 220 quina 184, 294 quina falsa 294
raicilla 255
raicilla macho 247
Randia 279
Randia aculeata 36, 282
Randia aculeata var. dasyclada 282
Randia altiscandens 37, 282
Randia armata 36, 283
Randia brenesii 36, 283
Randia calycosa 36, 284
Randia crescentioides 289
Randia formosa 284
Randia genipifolia 126
Randia genipoides 38, 284
Randia gentryi 285
Randia grandifolia 37, 285
Randia grayumii 38, 285
Randia karstenii 282
Randia loniceroides 36, 286
Randia matudae 38, 286
Randia mira 38, 287
Randia monantha 37, 287
Randia pepoformis 282
Randia pittieri 37, 288
Randia pterocarpa 288
Randia retroflexa 288
Randia spinosa 283
Randia subcordata 287
Randia thurberi 36, 289
Randia vazquezii 289
rangayo 322
Raritebe 289
Raritebe palicoureoides 58, 290
Raritebe palicoureoides ssp. dwyer-
anum 290
Ravnia longifilamentosa 162 Ravnia panamensis 162 Ravnia pittieri 1 64 Ravnia triflora 164
332
FIELDIANA: BOTANY
Relbunium aschenbornii 140 Relbunium hypocarpium 140 Richardia 290 Richardia brasiliensis 29 1 Richardia scabra 18, 29 1 Richardsonia brasiliensis 29 1 Rogiera 292 Rogiera amoena 294 Ronabea erecta 247 Rondeletia 292 Rondeletia acuminata 297 Rondeletia affinis 295 Rondeletia amoena 55, 294 Rondeletia aspera 55, 294 Rondeletia bertierioides 295 Rondeletia biflora 3 1 2 Rondeletia brenesii 35, 295 Rondeletia buddleioides 35, 295 Rondeletia calycosa 55, 296 Rondeletia chaconii 56, 296 Rondeletia cooperi 297 Rondeletia costaricensis 55, 297 Rondeletia hamelifolia 56, 298 Rondeletia latifolia 294 Rondeletia leucophylla 300 Rondeletia macrocalyx 297 Rondeletia mexiae 297 Rondeletia microdon 261 Rondeletia monteverdensis 55, 298 Rondeletia pittieri 294 Rondeletia povedae 56, 299 Rondeletia repens 146 Rondeletia rugosa 294 Rondeletia salicifolia 299 Rondeletia scabra 297 Rondeletia stachyoidea 295 Rondeletia stenostachya 1 50 Rondeletia tayloriae 56, 300 Rondeletia torresii 55, 300 Rondeletia urophylla 35, 301 Rondeletia versicolor 294 Rosenbergiodendron formosum 284 Rudgea 301
Rudgea amplexicaulis 302 Rudgea ceratopetala 303 Rudgea chiriquiensis 173 Rudgea cornifolia 61 , 303 Rudgea fimbriata 303 Rudgea monofructus 303 Rudgea raveniana 304 Rudgea reducticalyx 61, 304 Rudgea skutchii 61, 305 Rudgea thyrsiflora 270 Rudgea trifurcata 61, 305 Rustia 306
Rustia costaricensis 52, 306 Rustia occidentalis 52, 307 Rustia panamensis 306
Sabicea 307 Sabicea ambigua 86 Sabicea costaricensis 308
Sabicea hirsuta 308 Sabicea panamensis 49, 308 Sabicea villosa 49, 308 Sabicea villosa var. adpressa 308 salamo 94 sangrenaria 323 Schradera 308 Schradera blumii 309 Schradera costaricensis 33, 309 Sherardia 310 Sherardia arvensis 3 1 0 Sickingia 3 1 1 Sickingia maxonii 3 1 1 Sickingia myriantha 128 Siderodendron floribundum 1 8 2 Simira 311
Simira maxonii 52, 31 1 Simira myriantha 128 Sipanea 3 1 1 Sipanea biflora 3 1 2 Solena latifolia 220 Sommera 312
Sommera donnell-smithii 49, 312 Sommera dunlapii 312 Sommera grandis 313 Sommera mesochroa 312 Sommera rivularis 312 Spermacoce 313 Spermacoce apiculata 1 24 Spermacoce assurgens 19, 315 Spermacoce confusa 19, 315 spermacoce densiflora 18, 316 Spermacoce ernestii 318 Spermacoce exilis 19, 316 Spermacoce hondurensis 3 1 3 Spermacoce glabra 3 1 9 Spermacoce hirta 192 Spermacoce laevis 315 Spermacoce latifolia 19, 316 Spermacoce longiflora 1 1 9 Spermacoce mauritiana 3 1 6 Spermacoce ocymifolia 20, 317 Spermacoce ovalifolia 19, 317 Spermacoce pringlei 317 Spermacoce prostrata 19, 318 Spermacoce rigida 1 24 Spermacoce riparia 319 Spermacoce scabiosoides 3 1 8 Spermacoce serrulata 125 Spermacoce spinosa 316 Spermacoce suaveolens 18, 319 Spermacoce tenuior 20, 319 Spermacoce vegeta 320 Spermacoce verticillata 18, 320 Spermacoce villosa 192 Stannia panamensis 220 Stomandra costaricensis 306 surra 94
tamarind of the Indies 323 Tapogomea tomentosa 268 Tarenna 99
teresa 294
Tedrtrea martinicensis 188 thrum flowers 1 Tocoyena 320 Tocoyena cuatrecasasii 321 Tocoyena obliquinervia 321 Tocoyena pittieri 44, 321 Tontanea canescens 105 Tontanea glabra 105 Tontanea guianensis 104 Tontanea herbacea 104 Tontanea hirsuta 104 Tontanea hispidula 104 Tontanea pleuropoda 104 Tournefortiopsis reticulata 155 Triodon angulatum 124 trompillo 82 trompito 82 tsus krd 1 60
Ugni myricoides 89
Uncaria 321
Uncaria tomentosa 51, 322
Vaillantia hypocarpia 140 Vangueria edulis 322 Vangueria madagascariensis 322 Varneria augusta 142 Virecta biflora 3 1 2 Viscoides pendulum 250
Warszewiczia 323 Warszewiczia coccinea 30, 323 Warszewiczia schwackei 129 Watsonamra donnell-smithii 214 Watsonamra gymnopoda 214 Watsonamra macrophylla 215 Watsonamra pubescens 215 Watsonamra wendlandii 216 wild coffee 220 wild guava 82 winged fruits 83
Xerococcus congestus 171 yema de huevo 95, 193
zorillo 160
zorillo Colorado 1 59
zorillo real 1 60
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
333
rding to thi
Typha
1 3 Hy
14 T"
17 Pa;
37 M 41 i
55 P:
5 9 An - 64 An
|
84 |
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|
Morin),, |
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|
'.ceae |
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Oxalida |
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Zygopl |
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|
Rutai |
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|
105 |
Simarubi |
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|
1 n/i |
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JUO |
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Malp. |
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HECKMAN
BINDERY INC.
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