Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora

Organization for Flora Neotropica 

Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera, Condaminea, Picardaea, Pogonopus, 

Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and Parachimarrhis) 

Author(s): Piero G. Delprete 

Reviewed work(s): 

Source: Flora Neotropica, Vol. 77, Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera, 

Condaminea, Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, 

and Parachimarrhis) (Jun. 25, 1999), pp. 1-225 

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FLORA NEOTROPICA MONOGRAPH 77 

RONDELETIEAE (RUBIACEAE)-PART I 

(Rustia, Tresanthera, Condaminea, Picardaea, Pogonopus, 

Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and 

Parachimarrhis) 

PIERO G. DELPRETE 

Publishedfor 

The Organization for Flora Neotropica 

by 

The New York Botanical Garden Press 

Bronx, New York 

Issued 25 June 1999

Flora Neotropica 

Published for 

Organization for Flora Neotropica 

by 

The New York Botanical Garden Press 

Executive Board 

Ariane Luna Peixoto, President, UFRRJ, Caixa Postal 74.582, 23851-970 Seropedica, Itaguai, 

RJ, Brazil 

Ren&e Fortunato, Vice President, Instituto de Recursos Biologicos, I.N.T.A., 1712 Castelar, Buenos 

Aires, Argentina 

Wm. Wayt Thomas, Executive Director, Ex Officio, Institute of Systematic Botany, The New York 

Botanical Garden, Bronx, NY 10458-5126, U.S.A. 

Dieter Wasshausen, Secretary, Department of Botany - NHB- 166, Smithsonian Institution, Washington, 

DC 20560-0001, U.S.A. 

Paul J. M. Maas, Treasurer, Institute of Systematic Botany, State University of Utrecth, Postbus 

80.102, 3508 TC Utrecht, The Netherlands 

Paul Berry, Botany Department, 132 Birge Hall, 430 Lincoln Drive, University of Wisconsin, 

Madison, WI 53706-1381, U.S.A. 

Mireya D. Correa A., Departamento de Botanica, Facultad de Ciencias Naturales, Exactas y 

Tecnologia, Universidad de Panama, Estafeta Universitaria, Panama, Panama 

Jose Rubens Pirani, Departamento de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, 

Caixa Postal 11461, 05422-970 Sao Paulo SP, Brazil 

Ghillean T. Prance, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K. 

Rodolphe Spichiger, Conservatoire et Jardin Botaniques, de la Ville de Geneve, Case Postale 60, 

CH-1292 Chambesy/GE, Switzerland 

Claudio Delgadillo M. (1996-2001) 

Laurence J. Dorr (1996-2001) 

Jose R. Pirani (1996-2001) 

Editorial Committee 

James L. Luteyn, Editor 

S. Rob Gradstein, Editor 

Julio A. Lombardi (1999-2004) 

Michael O. Dillon (1999-2004) 

Michael Nee (1999-2004) 

ORGANIZATION FOR FLORA NEOTROPICA 

Founded and Conducted under the Auspices of UNESCO

Rondeletieae (Rubiaceae)-Part I

Rondeletieae (Rubiaceae)-Part I 

(Rustia, Tresanthera, Condaminea, Picardaea, Pogonopus, 

Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and 

Parachimarrhis) 

PIERO G. DELPRETE 

CONTENTS 

A bstract/R esum en ........................................................................................................ 2 

Introduction to the Rondeletieae ............................. ............................... 3 

H istorical Sum m ary .................................................................................................. 4 

Phylogenetic Relationships ........................................ ................... 10 

Current Circumscription of the Rondeletieae, Including Notes on 

Genera Newly Included, Excluded, or Synonymized ........................................... 11 

Key to the Genera of Neotropical Rondeletieae ...................................................... 19 

Notes on Descriptions, Abbreviations, and Acronyms ............................................ 23 

Introduction to the Neotropical Genera Studied .......................................................... 23 

General Morphology .............................................................................................. 23 

Seed Morphology .......... ............................................................................. 33 

Pollen M orphology ................................................................................................... 42 

Wood Anatomy .......... .................................................... ................... 52 

Presence of Calcium Oxalate Crystals ..................................................................... 54 

K aryology ................................................................................................................. 54 

Pollination and Other Plant/Animal Interactions ..................................................... 54 

H abitat and Ecology ................................................................................................. 55 

U ses .......................................................................................................................... 55 

Species Concepts ............................................................. ................... 56 

Systematic Treatment of the Genera Studied ............................................................... 56 

1. Rustia . ................................................................................... 56 

2. Tresanthera ....... ............................................................................................ 96 

3. C on dam inea .......................................................................................................... 103 

4 . P icarda ea ............................................................................................................. 117 

5. Pogonopus ............................................................... .................. 121 

6 . C h im arrh is............................................................................................................ 13 7 

7. D ioicodendron ...................................................................................................... 187 

8. Molopanthera .............................................................. .................. 192 

9. D olichodelp hys ..................................................................................................... 196 

10. Parachimarrhis ...... ....................................................................................... 199 

A cknow ledgm ents ........................................................................................................ 203 

Literature Cited ......... ......................................................................... 203 

N um erical List of Taxa ................................................................................................ 209 

List of Exsiccatae .......... .................................................... ................... 210 

Nomenclatural List .............................................................. ................... 217 

Index of Local Names .................................. ........................... 220 

Index of Scientific Names ............................................................ 221

2 Flora Neotropica 

ABSTRACT 

Delprete, P. G. (Institute of Systematic Botany, The New York Botanical Garden, Bronx, 

NY 10458-5126, U.S.A.). Rondeletieae (Rubiaceae)-Part I (Rustia, Tresanthera, Condaminea, 

Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, 

and Parachimarrhis). Flora Neotropica Monograph 77: 1-226. 1999.-The tribe 

Rondeletieae, as here circumscribed, is a predominantly neotropical group of trees, shrubs, 

and rarely herbs. In the Neotropics it is represented by approximately 42 genera and 340 

species. The present volume is the first part of a series in which the monographic treatment 

of the neotropical Rondeletieae is planned. For this reason, a general definition and 

a taxonomic survey of this group, as well as a list of all the genera, their approximate 

number of species and their distribution, and a key to all the genera are included. 

This study originated as a revision of the tribe Condamineeae (sensu Hooker), a group 

of predominantly neotropical shrubs and trees (except Pinckneya from North America, 

and Bikkia and Badusa from the South Pacific), usually delimited by having valvate corollas 

and many-seeded capsules. The Condamineeae were subdivided by Hooker into three 

subtribes: Condamineinae, Portlandiinae, and Pinckneyinae. As a result of phylogenetic 

analyses using morphological characters, the Condamineeae (sensu Hooker) has been 

shown to be paraphyletic. The genera of the Portlandiinae have been transferred to the 

newly amended tribe Catesbaeeae sensu Delprete, and genera of the Condamineinae and 

Pinckneyinae transferred to the Rondeletieae, resulting in a complete dismantling of the 

Condamineeae. The Sipaneeae (Sipanea, Sipanaeopsis, Limnosipanea, and Neobertiera) 

are included in a broad delimitation of the Rondeletieae sensu Delprete. Additionally, 

Phyllomelia (with bi-seeded indehiscent fruits, i.e., pseudosamaras), recently treated as 

genus incertae sedis by Robbrecht, is also included in the Rondeletieae s.l., and believed 

to be related to Mazaea. 

The neotropical genera treated in the present monograph are those of the subtribes 

Condamineinae and Pinckneyinae, now included in the Rondeletieae, plus a few related 

genera: Rustia, Tresanthera, Condaminea, Picardaea, Pogonopus, Chimarrhis, 

Dioicodendron, Parachimarrhis, Molopanthera, and Dolichodelphys. Forty-six taxa, 40 

species, and 6 varieties, are recognized in the 10 genera listed above. 

RESUMEN 

Delprete, P. G. (Institute of Systematic Botany, The New York Botanical Garden, Bronx, 

NY 10458-5126, U.S.A.). Rondeletieae (Rubiaceae)-Part I (Rustia, Tresanthera, Condaminea, 

Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera, 

Dolichodelphys, and Parachimarrhis). Flora Neotropica Monograph 77: 1-226. 1999.- 

La tribu Rondeletieae, como esta aqui circunscrita, es un grupo predominantemente neotropical 

de arboles, arbustos y raramente hierbas. En el Neotr6pico la tribu esta representada 

por aproximadamente 42 generos y 340 especies. El presente volumen es la primera 

parte de una serie en la cual se esta programando el tratamiento monografico de los generos 

neotropicales de las Rondeletieae. Por lo tanto, se incluyen aqui una descripci6n de la 

tribu, un listado de todos los generos, con numero aproximado de especies y su distribuci6n 

geografica, y una clave para todos los generos neotropicales de este grupo. 

El presente estudio se origino a partir de una revision de la tribu Condamineeae (sensu 

Hooker), un grupo predominantemente neotropical (a excepci6n de Pinckneya, endemico 

de Norte America, y Bikkia y Badusa endemicos del Pacifico Sur) de arboles y arbustos, 

usualmente caracterizado por corolas valvadas y capsulas con muchas semillas. Las Condamineeae 

fueron divididas por Hooker en tres subtribus: Condamineinae, Portlandiinae 

y Pinckneyinae. Como resultado de un analisis filogenetico usando datos morfologicos, 

se demostro que la tribu Condamineeae (sensu Hooker) es un grupo parafiletico. Los 

generos de la subtribu Portlandiinae fueron transferidos a la tribu Catesbaeeae, recientemente 

enmendada por Delprete, y los generos de las Condamineinae y Pinckneyinae fueron

Introduction to the Rondeletieae 3 

transferidos a las Rondeletieae, causando asi el completo desmembramiento de la tribu 

Condamineeae. La tribu Sipaneeae (Sipanea, Sipanaeopsis, Limnosipanea and Neobertiera) 

esta tambien incluida en una mas amplia delimitacion de las Rondeletieae sensu Delprete. 

Adicionalmente, Phyllomelia (con frutos indehiscentes y con dos semillas, i.e., pseudosamaras), 

recentemente tratado como genus incertae sedis por Robbrecht, es tambien 

incluido en las Rondeletieae s.l. y certamente relacionado con Mazaea. 

Los generos neotropicales tratados en la presente monografia, tradicionalmente incluidos 

en las Condamineineae y Pinckneyinae y ahora transferidos a las Rondeletieae (a los cuales 

se afiaden unos generos relacionados), son: Rustia, Tresanthera, Condaminea, Picardaea, 

Pogonopus, Chimarrhis, Dioicodendron, Parachimarrhis, Molopanthera y Dolichodelphys. 

Cuarenta y seis taxa (40 especies y 6 variedades) son descritos en los 10 generos 

anteriormente listados. 

INTRODUCTION TO 

THE RONDELETIEAE 

The Rubiaceae is the fourth largest family of flow- 

ering plants, with approximately 650 genera and 

12,000 species, with mostly pantropical distribution 

and predominantly woody. In the Neotropics it is the 

largest dicotyledonous family, with approximately 

200 genera and 5000 species. Andersson (1992) cited 

about 4500 species, but considering the large amount 

of new taxa recently described from material collected 

in newly explored neotropical areas, his estimate of 

the neotropical species is, in my opinion, too low. 

This family is easily recognized by its opposite 

(rarely whorled, or exceptionally alternate) leaves 

with entire margins, interpetiolar (rarely intrapetiolar) 

stipules with colleters on the adaxial side, gamopetalous 

corollas, and inferior ovaries (exceptionally halfsuperior 

or superior). The latest comprehensive family 

classification was proposed by Robbrecht (1988, 

1993), where he recognized four subfamilies and 

44 tribes. Recent phylogenetic studies based on molecular 

data (Bremer et al., 1995; Rova et al., 1997) 

seem to indicate that the family should be divided 

into three subfamilies: Cinchonoideae, Ixoroideae, 

and Rubioideae. 

The Rondeletieae (subfam. Cinchonoideae), as 

here circumscribed, is a group of predominantly neotropical 

(except Pinckneya Rich. from North America; 

and Aleisanthia Ridl., Glionnetia Tirveng., Spathichlamys 

R. Parker, and Wendlandia Bartl. ex DC. 

being paleotropical) trees, shrubs, and rarely herbs, 

characterized by corollas valvate, contorted or imbricate, 

stamens variably inserted at basal, medial, or 

distal positions on the corolla tube, anthers dehiscing 

longitudinally orporicidally (Rustia and Tresanthera), 

and few- to many-seeded capsules or two-seeded 

pseudosamaras. 

This study originated as a revision of the Conda- 

mineeae (sensu Hooker, 1873) (Delprete, 1996f), a 

group of predominantly neotropical shrubs and trees, 

usually delimited by having valvate corollas and 

many-seeded capsules (Hooker, 1873). The tribe Con- 

damineeae was founded by Hooker (1873) and sub- 

divided by him into three subtribes: Condamineinae 

(as "Eucondamineae"), Portlandiinae (as "Port- 

landieae"), and Pinckneyinae (as "Pinckneyeae"). 

This group has been shown to be paraphyletic 

(Delprete, 1995a, 1996d, 1996f; also see Phylogenetic 

Relationships in this volume). 

Because of their corolla aestivation (imbricate 

lobes with reduplicate tubes) and their base chromo- 

some number (x = 12, large chromosomes; Kiehn, 

1995, and pers. comm.), the genera of the Portlandiinae 

(Bikkia Reinw., Catesbaea L., Ceuthocarpus 

Aiello, Coutaportla Urb., Coutarea Aubl., Cubanola 

Aiello, Hintonia Bullock, Isidorea A. Rich., Nernstia 

Urb., Osa Aiello, Phyllacanthus J. D. Hook., Portlandia 

P. Browne, Schmidtottia Urb., Siemensia Urb., 

and Thogsennia Aiello) have been included in the 

newly emended Catesbaeeae (Delprete, 1995a, 

1996d). Ongoing phylogenetic studies based on morphological 

and molecular data will definitely clarify 

the relationships within the Catesbaeeae-Chiococceae 

complex (Delprete et al., in progress). 

Based on previous phylogenetic analyses 

(Delprete, 1995a, 1996d, 1996f), the results of which 

are summarized below (see Phylogenetic Relationships), 

the subtribes Condamineinae and Pinckneyinae 

(sensu Hooker, 1873) were transferred to the tribe 

Rondeletieae (sensu Delprete). The inclusion of the 

two subtribes in the Rondeletieae is supported by their


base chromosome number (x = 11, small chromosomes; 

Kiehn, 1995, and pers. comm.), many-seeded 

capsules, and variable corolla aestivation. The monophyly 

of the Rondeletieae s.l. is yet to be tested (Rova 

et al., in progress). The inclusion of the Condamineeae 

(pro parte) in the Rondeletieae implies that the valvate 

aestivation is not a sufficient character for tribal 

delimitation. In addition, the minute flowers of 

Chimarrhis (usually reported as valvate), Molopanthera, 

Dolichodelphys, and Parachimarrhis have imbricate 

aestivation. 

The tribe Sipaneeae (sensu Robbrecht, 1988, 1993) 

is also here included in the wide circumscription of 

the Rondeletieae, where they are exceptional only in 

their herbaceous habit. Of this group, only Sipanea 

Aubl. has been the subject of chromosome studies, for 

which Kiehn (1995) reports: "In Sipanea, one diploid 

and one tetraploid taxon on a basic number of x = 11 

[small chromosomes, pers. comm.] are known. While 

chromosomes ofmitotic metaphase are not elongated 

as in Rondeletieae but rather compact as in Naucleeae, 

interphase nuclear structures of Sipanea resemble 

those of both Rondeletieae and Naucleeae." 

Additionally, Phyllomelia (monotypic, endemic to 

The present volume is the first part of a planned 

series of monographic treatments of neotropical Rondeletieae, 

as presently circumscribed. Being an introductory 

volume, the section Historical Summary introduces 

the complex nomenclatural history and 

delimitation of the Rondeletieae as currently recognized. 

At the end of this section, a description of this 

tribe is presented, followed by additional notes on the 

genera here included in (or previously referred to) the 

Rondeletieae. A table summarizes all the neotropical 

genera of this group, with their approximate number 

of species, habits, distributions, habitats, and elevation 

ranges. Finally, a Key to Neotropical Rondeletieae 

is presented. 

The genera treated in the Systematic Treatment of 

this volume are those of the subtribes Condamineinae 

and Pinckneyinae, previously treated as part of the 

Condamineeae and now included in the Rondeletieae, 

plus a few genera related to them. An overview of 

morphological, anatomical, palynological, and ecological 

studies of the genera is also presented. The 

Systematic Treatment includes monographs of Rustia, 

Tresanthera, Condaminea, Picardaea, Pogonopus, 

Chimarrhis, Dioicodendron, Parachimarrhis, Molop- 

anthera, and Dolichodelphys. Finally, the Nomencla- 

tural List gives all the accepted taxa, orthographic 

variants, and synonyms, as recognized by the author. 

HISTORICAL SUMMARY 

Rubiaceae classification has undergone a series of 

rearrangements as scientific knowledge and technology 

became more sophisticated. The taxonomic history 

of this family, with emphasis on the tribe Rondeletieae 

s.l., is here divided into six time periods. 

During the Linnean period (1753-1789) only a few 

genera and species were known. The post-Linnean 

period (1789-1828) is characterized by the exploration 

of tropical areas and the discovery of many new 

taxa; Jussieu during this period proposed the first 

Rubiaceae classification, using locule number and 

number of ovules per locule as main characters. The 

Candollean period (1829-1872) started with Achille 

Richard, who in 1829 presented the first monographic 

work dedicated to the whole family. Richard's system 

highly influenced the classification proposed by 

A. P. de Candolle, which remained the standard reference 

until the end of the 1800s. The late 19th cen- 

Cuba) with two-seeded pseudosamaras (Spjut, 1994), tury (1873-1900) is marked by the innovative work 

is also included in the Rondeletieae s.l., and is believed of J. D. Hooker, the first rubiologist who followed 

to be related to Mazaea (see notes after Rondeletieae, evolutionary theories. Hooker's classification influbelow). 

enced the work of the other two great scholars of this 

period: Baillon and Schumann. The modem period 

(1900-1990) is punctuated by the milestone works of 

Bremekamp (1934, 1940, 1954, 1966), Verdcourt 

(1958), and Robbrecht (1988). Their classifications 

attempted to break away from phenetic systems; in 

constructing them they evaluated character evolution, 

using total evidence (i.e., morphology, anatomy, cytology, 

biogeography, etc.) and defined groups using 

sets of characters (vs. single character classifications). 

During the last decade of the 20th century (after 1990) 

Rubiaceae phylogenetic studies evolved quite rapidly, 

being influenced by the advent of cladistic methods 

and by the science of molecular biology. The various 

delimitations of the Rondeletieae, Condamineeae, 

Sipaneeae, Catesbaeeae, and Chiococceae are summarized 

in Figure 1. 

Linnean Period (1753-1789) 

Linnaeus (1753) simply listed and described the 

several genera known to him (a few of them neotro- 

pical), which he included in his Tetrandia Monogynia 

and Pentrandria Monogynia. In this work he de- 

scribed Rondeletia L., the type genus of this tribe. 

Adanson (1763) simply divided the Rubiaceae [as


SIPANEAE 

CaMl.do He mr 3.i3. Selhum.. Verort Brmahmp Rebobrert Brner DdMpre 

130 1873 1 191 158 193 1s2 i m tbreataert 

RONDELTI|EA| | | | | 

-SIA | | 

? 

'CON.. ........ ' .............. ............... ...................................... 

CONDAMINE_A_ 

-o- H U H 11 U H 11 | 

C,HIOCOCCEA 

.11 

I 

I I I I 

FIG. 1. Diagrammatic representation of the delimitations of the Rondeletieae, Sipaneeae, Simireae, Condamineeae 

(and its three subtribes), Catesbaeeae, and Chiococceae according to the various authors. Vertical bars indicate tribal delimitation: 

Rondeletieae (black), Sipaneeae (vertical lines), Simireae (horizontal lines), Condamineeae (white), Catesbaeeae 

(dotted), and Chiococceae (oblique lines). 

"Aparines"] into two sections: section I with two ovules 

per ovary, and section II with more than two ovules per 

ovary, without giving any further subdivision. 

Post-Linnean Period (1789-1828) 

Antoine Laurent de Jussieu (1789, 1807, 1820) 

produced the first important Rubiaceae classification, 

dividing them into four arbitrary sections: Sect. I, with 

dicoccous fruits and uniovulate locules; Sect. II, with 

bilocular fruits, uniovulate locules, and capsular or 

baccate; Sect. III, with bilocular fruits, multiovulate 

locules, capsular or baccate; and Sect. IV, with multilocular 

fruits, uniovulate locules, baccate or 

drupaceous. In the second section he included 

Chimarrhis, due to Jacquin's (1763) erroneous observation 

that genus had two-seeded capsules. 

Rafinesque (1820) proposed a new elaboration of 

the Linnean classification for his "Classe Endogynie," 

which he arbitrarily divided into three orders based 

on the number of locules and number of seeds per 

locule. In this system the members of the Rubiaceae 

were represented as a mixture of previously described 

and Rafinesque's odd new genera, spread throughout 

his several ill-defined "families" ("Famille[s] Sambucidia, 

Dilarnia, Polarnia, Lamospermia, Aparinia, 

Cephalanthia, Viburidia"), which were in turn disseminated 

within the last two orders (Polyspia and 

Sphanidia) of his classification. Needless to say, 

Rafinesque's system was never adopted. 

Candollean Period (1829-1872) 

The two leading Rubiaceae classifications (A. Ri- 

chard, 1830; A. P. de Candolle, 1830) of this period 

were still pervaded by fixed species concepts. 

Achille Richard (son of Louis Claude Marie Ri- 

chard) presented a new Rubiaceae classification of 11 

tribes and 154 genera at the Royal Academy of Sci- 

ences in Paris on 7 July 1829. As Richard explained 

in the preface of this work, during that year Augustin 

Pyramus de Candolle was writing the treatment of the 

Rubiaceae for his Prodromus, and Richard was invited 

to submit his unpublished manuscript to be included 

in the Prodromus. Richard's Memoire sur lafamille 

Rubiacees (the first Rubiaceae monograph) went to 

press separately, in Paris in 1829, and was eventually 

published in December 1830 (and reprinted in 1834). 

De Candolle's Prodromus was published in Septem- 

ber 1830, anticipating the publication of Richard's 

Memoire by three months. Richard's taxa (tribes and 

genera) newly published in the Prodromus, were not 

or only inconsistently cited by de Candolle, some fol- 

lowed by "Rich." and others without any citation. I 

agree with Steam's (1957) conclusion that all of the 

new taxa and new combinations established by Rich-


ard and published in the Prodromus should be cited 2. Portlandiinae [as "Portlandieae. Calycis limbus 

as: "A. Richard in de Candolle." Richard (1830), in brevis v. elongatis, lobi nunquam in laminan 

his Memoire, founded and described the tribe foliaceam dilatati. Corollae tubus elongatus, 

Cinchoneae so as to include those rubiaceous genera saepissime angulatus, lobi reduplicatim valwith 

bilocular many-seeded capsules and winged vati"], with Portlandia, Isidorea A. Rich. in 

seeds, a group that included nine genera of the Ron- DC., Bikkia, and Morierina Vieill. 

deletieae s.l. 

3. Pinckneyinae [as "Pinkneyeae. Calycis lobus 

Augustin Pyramus de Candolle 

unus hic illic in laminam foliaceam 

(September 1830) 

amplam 

largely followed (and sometimes 

coloratum 

literally reproduced) 

petiolatam dilatatus"], with Pinck- 

Richard's classification, but with several modifica- neya [as "Pinkneya"] and Pogonopus. 

tions and innovations, presenting a system of 13 tribes 

and 223 genera. Following Richard, he maintained the In the Rondeletieae, Hooker included Warszetribe 

Cinchoneae (as "Cinchonaceae"). Adding to wiczia Klotzsch, Pallasia Klotzsch (= Wittmackanthus 

Richard's classification, de Candolle (1830) validated Kuntze), Bathysa C. Presl, Elaeagia Wedd., Chalepothe 

tribe Hedyotideae proposed by Chamisso and phyllum J. D. Hook., Lindenia Benth. (= Augusta 

Schlechtendal (1829), and established the subtribe Pohl), Gloneria Andre (= Psychotria L.), Augusta, 

Rondeletiinae [as "Rondeletieae"], where he placed Greenea Wight & Am. [as "Greenia"], Deppea Cham. 

nine genera of the Rondeletieae s.l. 

& Schltdl. (Hamelieae), and Carphalea Juss. [as 

Endlicher (1838) largely followed de Candolle's "Carphalia"]. 

classification and divided the tribe Hedyotideae into Baillon (1880) followed both de Candolle's and 

the subtribes Hedyotidinae (as "Hedyoteae") and Hooker's classifications but thought that corolla aes- 

Rondeletiinae (as "Rondeletieae"). 

tivation alone was insufficient to separate rubiaceous 

tribes. Baillon's tendency was to lump similar tribes 

Late 19th Century (1873-1900) 

and genera superficially, producing a classification 

that subdivided the Rubiaceae into 13 tribes and 194 

Joseph Dalton Hooker (1873) produced the first 

Rubiaceae classification that followed 

genera. He included the Condamineeae and the Ronevolutionary 

deletieae in the "Portlandia Series" or "Portlandieae" 

concepts, which were proposed by his friend and colleague 

Charles Darwin (1859). Hooker's classifica- 

(both of them nom. illeg. because the group contains 

tion was a major breakthrough in the classification of 

Rondeletia). Baillon went into lengthy speculative 

this increasingly large family, where he proposed 25 

explanations in which he hypothesized relationships 

tribes and 337 genera. Hooker was the first to intro- among the genera that he included in the same "series" 

or 

duce corolla aestivation as a 

tribes, 

cardinal tribal character 

increasing the understanding of this 

in constructing a phylogenetic classification. In this 

large family and supporting evolutionary theories, 

which were still debated 

work he established the tribe Condamineeae 

during his times. 

[as 

Herman Karsten 

"Condaminieae"], separating it from the Rondeletieae. 

(1887) commented on the newly 

established Condamineeae of J. D. Hooker 

He defined the Condamineeae as a group with valvate 

(1873: 

corollas (vs. imbricate or contorted in the Rondele- 

356) as follows (free translation from German): 

tieae), and distinguished it from the Hedyotideae in Bentham & [J. D.] Hooker, like their predeceshaving 

simple stipules, sometimes bifid, and with sors, believe that the Condamineeae are natuwoody 

habit (vs. dentate to setose stipules and herba- ral genera, when they use the deciduous or perceous 

habit). Hooker also stated that the Condami- sistent calyx lobes and the basifixed or 

neeae and the Hedyotideae were similar in having dorsifixed anthers together with the mode of 

capsular fruits with numerous horizontal seeds [al- dehiscence [of the anthers] as principal charthough 

Chimarrhis (Condamineeae sensu Hooker) has acters. The related Cascarilla [Wedd. (= Ladenvertical 

peltate seeds] that are non-winged to minutely bergia Klotzsch)] shows, however, that each 

winged. Hooker (1873) subdivided the Condamineeae character of the calyx lobes appears variable in 

into three subtribes: 

some of its species and has no generic value. 

Condaminea and Chimarrhis cannot be distin- 

1. Condamineinae [as "Eucondamineae. lobi 

guished from each other in the mode of openaequalis 

v. subequales. Corollae tubus teres v. 

ing of their capsules alone, therefore [the capsub-0; 

lobi simpliciter valvati"], with Conda- sule dehiscence] cannot be treated as an 

minea, Chimarrhis, and Rustia (incl. Henlea 

important character. 

and Tresanthera)


Karl Schumann (1888a, 1889, 1891) presented a A few years later Bremekamp (1940) restated his 

classification of 21 tribes and 346 genera that influ- skepticism about the Rubiaceae classifications availenced 

Rubiaceae systematics throughout the 20th cen- able at that time: "All botanists acquainted with the 

tury. His work was influenced by all his predecessors, family Rubiaceae will agree that the present subdivimost 

strongly by Hooker (1873). Following Hooker's sion [cf. Schumann, 1891] is far from satisfactory and 

classification, Schumann (1888a, 1891) maintained that more of its tribes are either artificial or ill-defined 

the Condamineeae [as "Cinchonoideae-Cinchoninae- or both." In this little-known paper he predicted the 

Condamineeae"] separate from the Rondeletieae, but course of future Rubiaceae studies during the 20th 

he did not recognize the subtribal ranks proposed by century, declaring that fruit succulence should be re- 

Hooker. In addition, Schumann (1888a, 1891) pro- garded as being of secondary importance, and that the 

duced the first comprehensive keys to the 10 genera primary taxonomic characters were to be found in 

that he included in the Condamineeae and to the 15 

genera of his Rondeletieae. Of the genera here included presence or absence of hairs in the corolla 

in the Rondeletieae, Schumann 

throat; the 

(1888b) 

place where the stamens are inserted 

presented 

in the 

detailed discussions of Sickingia Willd. [= Simira tube; the presence or absence of floral 

Aubl.], Chimarrhis, Pinckneya sensu Allemao & dimorphism; the uni- or bisexuality of the flow- 

Saldanha 

ers; the insertion and 

(Saldanha, 1872) [= Simira], Tresanthera, 

shape of the placenta; the 

number of 

Henlea [= Rustia], Rustia, and Phitopis J. D. Hook. 

ovary cells and the presence or absence 

of false septs [septa]; the nature of the 

Modern Period 

stipules; the position of the inflorescences, etc. 

(1901-1990) 

These characters, however, are at present but 

During the first half of the 20th century, system- imperfectly known, and therefore a satisfactory 

atists strived to construct phylogenetic classifications subdivision can be given, the genera themselves 

by attempting to detect "primitive" or "derived" char- will have to be studied in more detail. 

acters. The concept was taken to extremes by [Bremekamp, 1940: 172] 

Hutchinson (1926), who divided the Angiosperms into 

Verdcourt 

two main, supposedly phylogenetic, branches: herba- 

(1958), strongly influenced by 

ceous and woody. Nevertheless, in his key to 

Bremekamp's early work, was the second rubiologist 

tropical to 

West African Rubiaceae (Hutchinson & 

classify the Rubiaceae on the basis of total evidence, 

Dalziel, 

1931) he maintained corolla aestivation, number of 

adding new sets of characters, such as chromosomes, 

ovules per locule, and fruit fleshiness as cardinal tribal 

pollen grains, hairs, stipules, raphides, seed albumen, 

and 

characters for this 

embryos, and constructing the Rubiaceae classifamily. 

fication 

In 1934 Bremekamp published a highly influenusing 

sets of characters. He divided the Rubiaceae 

into three subfamilies 

tial, and at that time quite controversial, article. In it 

(Rubioideae, 

he criticized Baillon's and Schumann's systems for 

Cinchonoideae, and Guettardoideae) and 28 tribes, 

being constructed on groups formed by a dichotomous 

using presence or absence of raphides (calcium oxclassification 

based on single characters, and noted 

alate), more or less developed embryos, and presence 

or absence of albumen in the seeds as main characthat 

several of their groups were paraphyletic. He 

ters. He combined the Condamineeae with the Rondeclared 

that the wings of certain seeds were not significant 

taxonomic characters, contrary to Baillon's 

deletieae, stating: 

opinion, and doubted that Rubiaceae classifications The Rondeletieae are well-characterized by 

based on ovule number and ovules per locules were their capsular fruits containing large numbers 

natural. Bremekamp was also the first to group cer- of minute horizontal seeds. The testa cells are 

tain genera according to their pollination mechanism pitted and the aestivation not valvate. The Con- 

(the "ixoroid pollination syndrome"). In the same damineeae are very similar but the aestivation 

paper, Bremekamp placed Chimarrhis in the tribe is valvate and the seeds often winged. In both 

Chimarrheae, but without giving any description or the habit is mainly arborescent and the stipules 

delimitation of the group, so this tribal name is as a are entire or at the most bifid. I would combine 

nomen nudum; he apparently abandoned this tribal these two groups into one tribe and consider 

name because it never appeared again in his later them to be subtribes at the most [subtr. 

work. In addition, he separated the tribe Sipaneeae Condamineinae Verdcourt]. In the Cinchoneae 

from the Rondeletieae, principally because of its her- placed next to them the aestivation character 

baceous habit (Sipaneeae is here returned to the Ron- is considered valueless by all previous workdeletieae). 

ers because if used it would divide an other-


wise compact tribe-all three types ofaestiva- inae are worthy of recognition at tribal level." This 

tion occurring in it. I do not think this one char- was probably the first suggestion that the Condamiacter 

is sufficient to separate the Rondeletieae neeae sensu Hooker (1873) were a paraphyletic asand 

Condamineeae particularly as two types of semblage. 

aestivation occur in the Rondeletieae. Garcia Kirkbride (1982) presented a linear system 

[Verdcourt, 1958: 245] 

for the Rubiaceae to be used for floristics. In this work 

she constructed two hypothetical phylogenetic trees, 

In his last paper on Rubiaceae classification, based on the presence or absence of albumen and the 

Bremekamp (1966) analyzed Verdcourt's classifica- presence or absence of raphides, treating the most 

tion, with which he partly agreed. In contrast to his common state of these two characters as the primiprevious 

classification (Bremekamp, 1934), he di- tive one. After her rather speculative phylogeny, she 

vided the Rubiaceae into 8 subfamilies and 43 tribes. decided to adopt the same subfamilial and tribal clas- 

In agreement with Hooker (1873) and Schumann sification proposed by Verdcourt (1958), including the 

(1891), but contrary to Verdcourt (1958), he main- Condamineeae in the Rondeletieae, but she listed the 

tained the Condamineeae separated from the Ronde- taxa with the same linear arrangement presented by 

letieae, stating: "The Rondeletieae differ from the Bremekamp (1954b, 1966) "because it better illus- 

Condamineae [sic] in the aestivation of the corolla trates the subfamilial phylogeny of the Rubiaceae" 

lobes, which in this tribe is either imbricate or con- (Garcia Kirkbride, 1982). 

torted. However, as these two types of aestivation are, Robbrecht (1988), in his comprehensive treatment 

in contrast to imbricate and valvate, but rarely met Tropical Woody Rubiaceae, divided the family into 4 

with in nearly related genera, it is not impossible that subfamilies (Rubioideae, Antirheoideae, Ixoroideae, 

further study will lead to splitting up of this tribe [Ron- and Cinchonoideae) and 44 tribes (and estimated 637 

deletieae]." In his general discussion Bremekamp genera and 10,700 species). In this classification he 

(1966) also added: "With regard to his [Verdcourt's maintained the views of Hooker (1873) and 

(1958)] reduction of the Condamineeae to the Ron- Bremekamp (1966), and maintained the Condamineeae 

deletieae I feel rather skeptical, but I admit that both separated from the Rondeletieae, without recognizing 

tribes are very imperfectly known and deserve a any subtribal divisions. He described the Condamithrough 

revision." 

neeae as follows: "Trees or shrubs. Stipules mostly 

Aiello (1979) followed Bremekamp's (1966) clas- interpetiolar, entire, rarely bifid. Raphides absent. 

sification, and treated seed insertion and position as Inflorescences mostly terminal and many-flowered. 

key tribal and generic characters. She placed Corolla aestivation valvate, rarely imbricate. Ovary 

Portlandia and some of the genera that she separated 2-locular, with many horizontal ovules in each locfrom 

Portlandia in the Condamineeae, because of ule. Capsules with many horizontal seeds. Exotestal 

their horizontal seeds; transferred Siemensia from the cells with pitted thickening along the inner tangential 

Condamineeae to the Hedyotideae (subfam. Rubioideae) walls. Pollen mostly 3-colporate, rarely porate 

because of their raphides, multicellular hairs, and (Bikkia)." He listed the following genera under the 

numerous tiny seeds; and transferred Coutarea Aubl. tribe Condamineeae: Badusa A. Gray, Bikkia, 

to the Cinchoneae because of its winged vertical seeds. Ceuthocarpus, Chimarrhis, Condaminea, Cubanola 

Finally, she treated Hintonia Bullock, Ceuthocarpus, Aiello, Dioicodendron, Flexanthera [= Simira], 

Coutaportla, Cigarilla Aiello [= Nernstia Urb.], and Isidorea, Kerianthera J. H. Kirkbr. [Isertieae], 

Schmidtottia as genera of uncertain position, and de- Morierina, Nernstia, Osa, Parachimarrhis, Picarclared 

that "none of the above placements can be made daea, Pinckneya, Pogonopus, Portlandia, Rustia, 

with great certainty since a large amount of work re- Schmidtottia, Siemensia, Stomandra [= Rustia], 

mains to be accomplished with regard to tribal rela- Thogsennia, and Tresanthera, and tentatively included 

tionship within the Rubiaceae." 

Kajewskiella Merr. & Perry and Pseudomussaenda 

Ridsdale (1982) reported: "In aestivation the sub- Wernham. 

tribe Portlandiinae differs from the remainder of the The Rondeletieae were described by Robbrecht 

Condamine[e]ae; Coutarea, Exostema (Pers.) Humb. (1988) as follows: "Trees or shrubs, rarely (Sipane- 

& Bonpl., Hintonia, and Schmidtottia differ from this opsis Steyerm.) herbaceous. Stipules entire, rarely 

subtribe only in the orientation of the seeds and their bifid. Raphides absent. Further little differing from the 

mode of attachment. However, the remainder of the Condamineeae: corolla lobes imbricate or more rarely 

Cinchoneae is also heterogeneous in this respect contorted. Flowers frequently heterostylous. Pollen 

(Ridsdale & Friskus, unpublished observations). Fur- mostly 3-colporate." In this tribe he included 

ther study might as well indicate that the Portlandi- Acrobotrys K. Schum & K. Krause, Acunaeanthus


Borhidi, Koml. & M. Moncada, Aleisanthia, Arach- 

nothryx Planch. [= Rondeletia s.l.], Augusta, Bathysa, 

Blandibractea Werham [= Simira], Chalepophyllum, 

Cuatrecasasiodendron Standl. & Steyerm., Dendro- 

sipanea Ducke, Eizia Standl., Elaeagia, Glionnetia, 

Gloneria Andre [= Psychotria L.], Greenea, Holsti- 

anthus Steyerm., Javorkaea Borhidi & Koml., 

Lindenia [=Augusta], Macbrideina Standl., Maguire- 

othamnus Steyerm., Neblinathamnus Steyerm., Pteri- 

docalyx Wernham, Rogiera Planch. [= Rondeletia 

s.l.], Roigella [= Rondeletia s.l.], Rondeletia, Simira, 

Sipaneopsis, Spathichlamys, Steenisia Bakh.f., 

Stevensia Poit., Warszewiczia, and Wendlandia. 

Following Bremekamp (1966), Robbrecht provi- 

sionally maintained the Sipaneeae (with Limno- 

sipanea, Sipanea, and Steyermarkia, and treating 

Neobertiera as a dubious genus) separate from the 

Rondeletieae because of their herbaceous to 

subfrutescent habit, but noted that "a revision of the 

Rondeletieae/Condamineeae 

complex may well show 

that this is not justified." 

The Last Decade of the 20th Century 

The last decade of the 20th century has seen the 

advent, and the first applications in the Rubiaceae, of 

a new methodology and a new branch of science that 

revolutionized the apparently stable system proposed 

by Robbrecht (1988): cladistic analyses and molecular 

biology. In addition, during this decade the Rubiaceae 

has finally received the increased attention of 

the botanical community, as shown by the organization 

of two international Rubiaceae congresses and by 

the considerable proliferation of botanists dedicated 

to Rubiaceae systematics. 

Andersson and Persson (1991) published the first 

Rubiaceae cladistic analysis in which, using morphological 

characters, they attempted to circumscribe the 

tribe Cinchoneae. Partly as a result of this analysis, 

the genera Cigarilla (= Nernstia), Coutarea, Exostema, 

Molopanthera, Syringantha Standl., and Wernhamia 

S. Moore (= Simira) were transferred from the 

Cinchoneae to the Condamineeae. Andersson and 

Persson (1991) commented on these transfers as follows: 

"Coutarea and Exostema seem to have their true 

relationship with Portlandia and related genera. 

Portlandia is presently referred to the tribe Condamineeae 

[sensu Robbrecht, 1988], to which Coutarea 

and Exostema may thus be referred. Unfortunately, it 

seems that this might only create new problems, since 

a relationship between Condaminea and the Portlandia 

group is not supported by our analysis." Their 

comments support the inclusion of the "Portlandiagroup" 

in the Catesbaeeae sensu Delprete (1996d). 

Bremer and Jansen (1991) presented the first cladistic 

analysis of the Rubiaceae using molecular data 

(chloroplast DNA restriction site mutations) obtained 

from 33 representative genera from 18 tribes (sensu 

Robbrecht, 1988). In their cladograms, Pogonopus 

and Pinckneya shared the same clade with Calycophyllum 

DC. (Calycophylleae; cf. Andersson & 

Persson, 1991) and Mussaenda L. (Isertieae sensu 

Robbrecht, 1988), and the same basal clade with 

Erithalis P. Browne and Chiococca P. Browne, Exostema, 

Coutarea, and Catesbaea. The results of their 

study suggested for the first time a relationship between 

the subtribe Portlandiinae (Condamineeae) with 

the tribe Chiococceae. This is also supported by several 

morphological similarities (i.e., filaments attached 

at the base of corolla tube, anthers basifixed, 

stigmatic area consisting of two narrow lines, etc.). 

Bremer (1992) presented a cladistic analysis using 

20 morphological characters of genera in subtribe 

Portlandiinae (Condamineeae) and tribe Chiococceae. 

She concluded that the former group should be included 

in the latter. Many morphological characters 

were not considered in her analysis: fruit morphology, 

number and position of ovules (many horizontal or 

imbricate in Portlandiinae vs. two pendulous in 

Chiococceae s.str.), placentation (central in Portlandiinae 

vs. apical in Chiococceae s.str.), anther shape and 

attachment (linear and basifixed in Portlandiinae vs. 

elongate and dorsifixed in Chiococceae). 

At the First International Conference on the Systematics 

of the Rubiaceae (4-6 October 1993, St. 

Louis, Missouri; cf. Taylor, 1995), Bremer (1996) 

presented a phylogenetic analysis using rbcL molecular 

sequences that indicated that, in contrast to 

Robbrecht's (1988, 1993) classifications, the Rubiaceae 

are better divided into three major groups, 

namely, the Cinchonoideae s.str., Ixoroideae s.l., and 

Rubioideae. During this conference, Delprete (1993) 

presented the results of a preliminary phylogenetic 

analysis of 43 genera using 44 morphological characters 

of all Condamineeae (sensu Robbrecht, 1988), 

all Chiococceae (sensu Robbrecht, 1988), several 

Rondeletieae, two Cinchoneae (Cinchona L. and 

Joosia H. Karst.), and Coffea L. (Coffeae) as 

outgroups. He concluded that at the tribal level the 

Portlandiinae (Hooker, 1873) should be separated 

from both the Condamineeae s.str. and the Chiococceae 

s.str., and instead be included in the Catesbaeeae 

(a tribe treated as incertae sedis by Robbrecht, 

1988). For further information about this analysis see 

Phylogenetic Relationships, below. 

Robbrecht (1993), following the conclusions presented 

by Delprete (1993), treated the Portlandiinae 

as an informal "Portlandia-group" within the Conda-


mineeae, but had reservations about including the PHYLOGENETIC RELATIONSHIPS 

group in the Catesbaeeae (sensu Hooker). The genera 

listed under the Condamineeae by Robbrecht 

(1993) were Chimarrhis, Condaminea, Dioicodendron, 

Flexanthera [= Simira], Kerianthera [Isertieae], 

Parachimarrhis, Picardaea, Pinckneya, Pogonopus, 

Rustia (incl. Stomandra), Tresanthera, and tentatively 

Kajewskiella Merr. & Perry, with Pseudomussaenda 

Wernham transferred to the Isertieae. Robbrecht 

(1993) made several changes in the genera that he 

previously included in the Rondeletieae (Robbrecht, 

1988). Following Lorence (1991), Robbrecht (1993) 

treated Arachnothryx, Rogiera, and Javorkaea as synonyms 

of Rondeletia, and also added Mazaea Krug 

& Urb. He maintained the Sipaneeae (to which he 

added Neobertiera Wernham) and separated the 

Simireae (here included, and proposed to be closely 

related to Parachimarrhis) from the Rondeletieae. 

At the Second International Rubiaceae Conference 

(13-15 September 1995, Meise, Belgium), Bremer 

(1996) presented a second phylogenetic analysis using 

rbcL molecular sequences that supported her previous 

results (Bremer, 1995) concerning the monophyletic 

origin of the Rubiaceae and their subdivision 

into three main lineages. Delprete (1995a, 1996d) 

presented a second set of tribal-level phylogenetic 

analyses using morphological data, from 170 species 

of 44 genera, including all Condamineeae and Catesbaeeae 

sensu Hooker and selected Chiococceae, Rondeletieae, 

Cinchoneae, and Coffeae. According to the 

results of these analyses (Delprete, 1995a, 1996d), the 

two subtribes Condamineinae and Pinckneyinae were 

transferred into a broader, tentative delimitation of the 

Rondeletieae, which is adopted, with few modifications, 

in the present treatment. He also included the 

subtribe Portlandiinae (Condamineeae sensu Hooker, 

1873) in the Catesbaeeae. The tribe Catesbaeeae 

(sensu Hooker, 1873) has nomenclatural priority over 

the subtribe Portlandiinae (also, Portlandieae Baillon 

is a nomen illegitimum because it includes Rondeletia); 

therefore, the newly emended tribe should be called 

"Catesbaeeae J. D. Hooker emend. Delprete" (Delprete, 

1996d). An abbreviated version of these analyses is 

presented in Phylogenetic Relationships, below. 

Classification at subfamilial and tribal levels in the 

Rubiaceae has been debated since its establishment, 

and up to the present day no overall consensus among 

workers has been reached. As certain taxonomic characters 

are carefully studied, or new ones are added to 

phylogenetic analyses, the macrosystematics of the 

Rubiaceae have been reexamined and modified accordingly. 

The number of recognized subfamilies within the 

Rubiaceae varied according to systematists' interpretation 

of the usefulness and taxonomic importance of 

certain characters. Using similar data sets, Verdcourt 

(1958) proposed three subfamilies (Rubioideae, 

Cinchonoideae, and Guettardoideae) whereas 

Bremekamp (1966) divided the Rubiaceae into the 

largest number of subfamilies ever proposed, eight: 

Cinchonoideae, Urophylloideae, Pomazotoideae, 

Gleasonioideae, Guettardoideae, Ixoroideae, Rubioideae, 

and Hillioideae. The most recent comprehensive 

classification using morphological, anatomical, 

and cytological data was proposed by Robbrecht 

(1988, 1993), who recognized four subfamilies 

(Cinchonoideae, Rubioideae, Antirheoideae, and 

Ixoroideae) and 44 tribes (a compromise between 

Verdcourt's and Bremekamp's classifications). 

Bremer (Bremer et al., 1995; Bremer, 1996) presented 

phylogenetic analyses using rbcL molecular sequences 

showing evidence for recognition of only 

three subfamilies (merging the Antirheoideae within 

the Ixoroideae s.l.), which was later supported by 

trnL-F (Rova et al., 1997) and rps 16 (Andersson, 

unpubl. data) molecular sequences. 

The phylogenetic analyses using molecular and 

morphological data presented by Bremer (1992), 

Bremer and Eriksson (1992), and Bremer and Struwe 

(1992) supported Bremekamp's and Verdcourt's opinions 

about the ambiguous usefulness of certain flower 

and fruit characters. In my opinion, fruit and flower 

characters have often been superficially or erroneously 

interpreted in trying to detect phylogenetic relationships. 

In order to avoid the repetition of erroneous 

observations, morphological characters and characters 

states should be coded after direct observation of fresh 

Rova et al. (1997) presented a phylogenetic analysis 

using trnL molecular sequences of more than 100 

species representative of the most tribes in 

Robbrecht's (1988, 1993) classification. These analyses 

supported the rbcL phylogenies of Bremer et al. 

(1995) and Bremer (1996), and rpsl6 phylogenies of 

Andersson (in progress, unpubl. data) in dividing the 

Rubiaceae in three main groups: Cinchonoideae s.str., 

Ixoroideae s.l., and Rubioideae. 

or preserved material, instead of being reported from 

previous literature. The taxonomic significance of 

characters such as number of ovules per locule, placentation, 

mesocarp fleshiness, seed insertion and 

position within the fruit, dioecy, heterostyly, and corolla 

aestivation should be evaluated with accurate 

anatomical and morphological studies, and compared 

with phylogenies obtained with alternative sets of data 

(i.e., molecular, biochemical, cytological data).


Recently, tribal rearrangements have occurred ses. In all analyses Rustia and Tresanthera were 

within subfam. Cinchonoideae, and mostly among the placed as sister genera on one clad,, supported by 

Cinchoneae, Rondeletieae, Condamineeae, Cates- three synapomorphies (8:1 = leaves with pellucid 

baeeae, and Chiococceae. The Condamineeae and the glands; 11:2 = presence of secundiflorous panicles; 

Sipaneeae have been historically treated either as sis- 42:1 = testa with ridges and small pits). Augusta and 

ter tribes (Hooker, 1873; Schumann, 1891; Lindenia were consistently placed as sister taxa on one 

Bremekamp, 1966) or as included (Baillon, 1880; clade, supported by testa sculpturing with small pits 

Verdcourt, 1958) in the Rondeletieae. The Chiococ- to ? smooth (42:0), which justifies Kirkbride's 

ceae were treated as a tribe (Hooker, 1873; Schumann, (1997b) recent inclusion of Lindenia in Augusta. 

1891; Bremekamp, 1934, 1966; Verdcourt, 1958; These analyses demonstrated that the Condami- 

Robbrecht, 1988) defined by two-seeded fleshy fruits neeae (sensu Hooker) were paraphyletic, and their 

until Bremer (1992) included the Portlandiinae (with three subtribes were rearranged and delimited accordmany-seeded 

capsules). Catesbaea and Phyllacanthus ingly. The Condamineinae and the Pinckneyinae, 

were separated from the Gardenieae by Hooker (1873) which included the genera treated in the present monoto 

establish the Catesbaeeae; these two genera were graph (Rustia, Tresanthera, Condaminea, Picardaea, 

referred to the Gardenieae (Ixoroideae) (Grisebach, Pogonopus, Pinckneya, Chimarrhis, Dioicodendron, 

1861; Baillon, 1880; Schumann, 1891), treated as a Molopanthera, and Dolichodelphys), were merged in 

separate tribe near the Gardenieae (Hooker, 1873; a broad delimitation of the tribe Rondeletieae, result- 

Verdcourt, 1958), or with uncertain relationship ing in the complete dismantling of tribe Condami- 

(Robbrecht, 1988). 

neeae. 

Monophyly and phylogenetic relationships of the According to these analyses, the subtribe Port- 

Condamineeae, Rondeletieae, Chiococceae, and landiinae (of the former Condamineeae sensu Hooker) 

Catesbaeeae (sensu Robbrecht, 1988) were tested in was included within the newly emended Catesbaeeae 

various cladistic analyses by Delprete (1993, 1995a, (treated by Robbrecht [1988] as incertae sedis), for 

1996d) using morphological data. The final analyses which a revised tribal delimitation was presented 

involved 170 species of 44 genera of all Condami- (Delprete, 1996d). The inclusion of the Portlandiinae 

neeae, all Catesbaeeae s.str. and selected Chiococceae, into the emended Catesbaeeae is supported by the 

Rondeletieae, Cinchoneae, and Coffea (Coffeae). karyological studies of Kiehn (1995, and pers. 

Forty-four morphological characters were scored from comm.), in which they are reported to be x = 12 and 

direct observation of herbarium specimens and/or liv- very large. 

ing material, and entered into a data matrix. Three In these analyses the Chiococceae and the Catesphylogenetic 

analyses were performed (using PA UP baeeae (both sensu Delprete, 1996d) appeared to be 

3.1 [Swofford, 1993]), each with a different outgroup: closely related, and in one analysis they were mapped 

the first using Cinchona and Joosia, the second using as sister groups. Nevertheless, recent studies using 

Coffea, and the third using Cinchona, Joosia, and trnL-F sequence data (Rova, Andersson, Delprete & 

Coffea simultaneously. A strict consensus tree and a Albert, in progress) indicated that Catesbaeeae and 

majority rule tree were produced for each analysis. Chiococceae (both sensu Delprete, 1996d) form a 

The consensus trees were produced from PA UP and strongly supported monophyletic group. 

were then transferred to MacClade (Maddison & 

Maddison, 1992) for character mapping and interpre- CURRENT CIRCUMSCRIPTION OF THE 

tation of state changes on the cladogram. Finally, RONDELETIEAE, INCLUDING NOTES ON GENERA 

synapomorphies and parallel homoplasies ("parallel- NEWLY INCLUDED, EXCLUDED, OR SYNONYMIZED 

isms") were mapped on the majority rule consensus 

trees obtained in each analysis. The majority rule con- The current definition and delimitation of the Ronsensus 

tree using Cinchona, Joosia, and Coffea as deletieae are tentative, and its monophyly has yet to 

outgroup is shown in Figure 2 (for detailed informa- be tested. The description of this group is intended for 

tion see Delprete, 1996d). In this analysis, the this floristic treatment and is not meant to be strictly 

Chiococceae, Portlandiinae, and the Catesbaeeae phylogenetic. Table I summarizes the neotropical 

formed a monophyletic group supported by one genera of Rondeletieae, and gives for each the apsynapomorphy 

(44:0 = pollen exine echinate), with proximate number of species, occurrence in neotrothe 

Catesbaeeae nested within the Portlandiinae. The pical countries, habitat, and elevation ranges. 

rest of the Condamineeae (Condamineinae and The following notes are necessary for clarification 

Pinckneyinae sensu Hooker) were interspersed with of the taxonomic position, delimitation, and recent 

the selected members of the Rondeletieae in all analy- synonymy of certain taxa that have at some point been


-'------ I'--------ID 

213:01 M:0 

8:1 1: 2 42:1 

,________ 

: 

|, 

2o:2 

3:2 i 

| 

CUB 

DAD 

CAT | 

It---------r 

"- 26:3 3 :1 - - -. ..T 

.PAR 

lobes absent; 35:3 =stigmatic surface linear I2:i3 along ,,.2 11 style; II 36:0 * =ovules 1 * 4:3 * pendulous; 37:1 =ovules - 

I= one =fruitM per locule; 38:1 

ss 

FOG 

CO . 

CHI 

BAT 

rr ~ ~ * BAUG 

JOO 

BAT 

COT =Coutaportla, NER=Nernstia, CUB Cubanola, THO =Thogsennia, CATC=Catesbaea, PHY Phyllacanthus, CEU 

FIG. 2. Majority rule consensus tree from the phylogenetic analysis of all genera of'the Condamineeae and Catesbaeeae 

(sensu Delprete): Catesbaeeae (black), Chiococceae (white) and Rondeletieae s.l. (oblique lines). Key to generic acronyms: 

RUS=Rustia, TRE =Tresanthera, CON =Condaminea, P00 =Pogonopus, PIN =Pinckneya, DOL=Dolichodelphys, DIO 

leathery FIG. 2.a berry; 38:2 =fruleit stronsengly esus tree =fruom the ple; phylogenetic 38:3 analysis of all genera of the Condamratineeae andg into two mericarps;baeeae 

=Ceuthocarpus, SCH =Schmidtottia, BIK--Bikkia, MOR-Morierina, EXO =Exostema, BAD =Badusa, CHI Chiococca, 

Joo41:2 =seed laterally flattened; 42:1 =testa ridged and withng strmall pits; 43:1 =testalr a indicated with dashed lines. Char- 

7:177,ERI Erithalis, fi Au-4. 1996). 

SCO Scolosanthus, PLA Placocarpa, CHO Chione, ALL Allenanthus, indenia, AUG 

entire marge indicated by double bars. Numbers indicate character and character states. Vertical bars indicate tribal delimitationg.


referred to the Sipaneeae, Condamineeae, and Ron- 1991), to which its only species was transferred as R. 

deletieae sensu Robbrecht (1988, 1993), but are now brandegeeana Lorence. 

excluded or not treated in the first part of the Ronde- Acunaeanthus Borhidi, Koml. & Moncada was 

letieae. The following discussion is organized into treated as distinct from Rondeletia by Delprete 

informal groups according to observations stemming (1996d). Acunaeanthus is similar to Mazaea in havfrom 

this research. These informal groups do not im- ing septicidal, oblong capsules, 4-merous flowers, 

ply any taxonomical status. 

tubular corollas, and imbricate corolla lobes; these 

characters are also shared with Rondeletia. Acunae- 

The Bathysa Group 

anthus differs from Mazaea in having terminal inflo- 

The two species of rescences 

Schizocalyx (Weddell, 1854; 

(vs. axillary), corollas pinkish white (vs. 

Standley, 1929) were recently shown to be conspe- orange), ovary with 15-30 ovules per locule (vs. 2-6 

cific and were transferred to ovules 

Bathysa by Delprete per locule), and winged seeds (vs. unwinged 

(1997b), as B. bracteosa (Wedd.) Delprete. The seeds). The two taxa are closely related and probably 

calycophylls of Schizocalyx have been proven to be congeneric, but further studies are necessary to ascerof 

little taxonomic significance, and this genus is tain their 

syn- relationships with Rondeletia. 

onymous with Bathysa. 

Phitopis was described by Hooker (187 la), who 

The Mazaea Group 

stated that it had the same habit as Hippotis Ruiz & Mazaea was published by Krug and Urban 

Pav. (tribe Hippotieae; see Garcia Kirkbride, 1981), (1897a). As explained by Robbrecht and Bridson 

the first name being an anagram of the other. 

(1993), Neomazaea Krug & Urb. (1897b) is a nomen 

Schumann (1888b) analyzed the position of Phitopis 

superfluum, and is treated as a synonym of Mazaea. 

and tentatively included it in the Rondeletieae. The type species of Ariadne Urb. was originally de- 

Standley (1931 d) added one species to this genus and scribed by Standley (1918) as Neomazaea shaferi 

maintained it in the Rondeletieae (Standley, 1936). Standl., and was later transferred by Urban (1922, 

Phitopis is here kept in the Rondeletieae as closely 1923) to his newly established Ariadne. Delprete 

related to Bathysa, with which it might be congeneric. (1999a) recently compared the monotypic Ariadne 

Bathysa was described by Presl (1845), based on with Mazaea and treated them as congeneric, and cre- 

Coffea stipulata Vell. With recent transfers (Delprete, ated the new combination Mazaea shaferi (Standl.) 

1996b, 1997b) and the discovery of new taxa, this 

Delprete. Ariadne is treated as a synonym of Mazaea 

genus now comprises approximately 16 species dis- 

(incl. Neomazaea Krug & Urb.; see Robbrecht & 

tributed in Central and South America. An account 

Bridson, 1993) because of its axillary inflorescences, 

of the Brazilian species in this genus was recently 4-merous flowers, imbricate corolla lobes, 1-3 ovules 

completed by Germano Filho (1993). 

per locule, and narrowly oblong capsules. 

The monotypic Phyllomelia was founded by 

The Rondeletia Complex 

Grisebach (1866), who placed it between Machaonia 

The generic limits of Rondeletia and its closely and Phialanthus (Chiococceae), probably because of 

related taxa are in a state of flux. In the present treat- the two-seeded indehiscent fruits. Phyllomelia was 

ment, a Rondeletia-complex is adopted which includes placed in the Ixoreae by Standley (1934) and treated 

Arachnothryx (Planchon, 1849), Javorkaea (Borhidi as genus incertae sedis by Robbrecht (1988, 1993). 

& Jarai-Komlodi, 1983), Rogiera (Planchon, 1849), Because of the axillary, few-flowered, cymose inflo- 

Roigella (Borhidi & Fernmndez, 1981a) and Suber- rescences with capilliform rachis, 4-5(-6)-merous 

anthus. The morphological characters used to segre- corollas, filaments inserted in the middle of the cogate 

these genera from Rondeletia s.str. (Planchon, rolla tube, anthers narrowly oblong, and absence of 

1849; Steyermark, 1967,1974; Borhidi & M. Fernndez, raphides in the blades, I concluded that Phyllomelia 

1981a, 1981b; Borhidi et al., 1980) have been shown is closely related to Mazaea (Delprete, 1999a). Both 

to be variable among species of Rondeletia s.str. genera are endemic to the serpentine soils of Sierra 

(Lorence, 1991). The inclusion of the above listed taxa del Rosario and Sierra de los Organos, Prov. Pinar del 

in the Rondeletia-complex does not necessarily as- Rio, Cuba. Phyllomelia is exceptional to the Rondesume 

their reduction to synonymy under Rondeletia. letieae s.l. by having indehiscent 2-seeded fruits with 

The Rondeletia-complex is here adopted for the prac- persistent rotate corolla, which are released as distical 

reasons of the presentation in this treatment. persal units, called "pseudosamaras" by Spjut (1994). 

The monotypic Otocalyx Brandegee has been For additional information about Phyllomelia, see 

shown to be congeneric with Rondeletia (Lorence, Delprete (1999a).

Table I 

Summary of the neotropical Rondeletieae (35 genera, ca. 340 species). Genera in boldface are 

No. of 

Genera species Countries" Habitat 

Acrobotrys I Co Cloud forests 

Acrosynanthus 6 Cu, J Pine forests, serpentine barrens 

Acunaeanthus 1 Cu Serpentine formations 

Augusta 4 M, G, Be, Ho, ES, N, CR, Pa, Among rocks in turbulent creeks 

Br (and South Pacific) 

Bathysa 16 CR, Pa, Co, V, Gu, E, Pe, Br, Lowlands of Central America, cloud forests, Amazoni 

Bo rain forests, Brazilian Atlantic forest 

Chalepophyllum 3 V, Gu Wet savannas and plateaux of the Guayana shield 

Chimarrhis 14 Cu, J, Ha, LA, Co, V, Gu, E, Caribbean seasonal forests, Andean slopes and lowlan 

Pe, Br, Bo Amazonian rain forests 

Condaminea 1(2?) CR, Pa, Co, V, E, Pe, Br, Bo Deciduous lowland forests, cloud forests, and rain 

forests 

Cuatrecasasiodendron 2 Co Colombian cordilleras 

Dendrosipanea 3 V, Br Flooded and non-flooded savannas 

Dioicodendron 1 Co, V, E, Pe Andean seasonal forests, and cloud forests 

Dolichodelphys 1 Co, V, E, Pe Andeail cloud forests, and rain forests 

Elaeagia 15 M, G, Be, Ho, Cu, CR, Pa, Co, Dry and wet seasonal forests, Cuban Sierra Maestra 

V, Gu, E, Pe, Br, Bo thickets, South American cloud forests 

Holstianthus 1 V Venezuelan mesetas 

Limnosipanea 5 Pa, Co, V, Br Llanos and wet savannas, pond margins and standing 

water 

Macbrideina 1 Co, E, Pe Amazonian slopes of the Andes and lowland Amazoni 

rain forest 

Maguireothamnus 4 V, Gu, Br Tepuis of the Guayana shield 

Mazaea 2 Cu, Ha Pine woodlands and serpentine barrens

Molopanthera 1 Br Seasonal and Atlantic forests 

Neblinathamnus 4 V, Br Cerro de la Neblina 

Neobertiera 1 Gu Lowland rain forest 

Parachimarrhis 1 Co, E, Br Amazonian rain forest 

Phitopis 2 Co, (E?), Pe Amazonian slopes of the Andes 

Phyllomelia 1 Cu Serpentine barrens 

Picardaea 1 Cu, Ha, DR Seasonal forests 

Pogonopus 3 M, G, ES, CR, Pa, Co, V, Pe, Seasonal and evergreen deciduous forests, and Andean 

Br, Bo, Ar cloud forests 

Pteridocalyx 2 Gu Lowland rain forests 

Rondeletia complex ca. 150 US, M, G, Be, Ho, ES, N, CR, Deciduous and evergreen seasonal forests; mostly 

Pa, Cu, J, Ha, DR, PR, VI, Caribbean,few species in Andean cloud forests (rarely 

LA, Co, V, Gu, E, Pe, Bo lowland Amazonian forests) 

Rustia 14 G, N, CR, Pa, Co, V, E, Pe, Br Andean cloud forests, seasonal forests, Amazonian 

slopes of the Andes, swampy coastal areas of Pacific 

coast, Brazilian Atlantic forests 

Simira 37 M, G, Be, Ho, ES, N, CR, Pa, Seasonal lowland forests, Andean cloud forests and 

Co, V, Gu, E, Pe, Br, Bo Amazonian lowland rain forests Brazilian Atlantic for 

Sipanea 18 CR, Pa, Co, V, TT, Gu, Br, Bo Guayana shield, and Caribbean coast of Central and 

South America, in poorly drained moist soil 

Sipaneopsis 8 Co, V, Br South American savannas 

Stevensia 9 Ha, DR Forested thickets on lateritic or serpentine soils 

Steyermarkia 1 M, G Coastal Caribbean lowlands 

Tresanthera 1 V, TT Rain forest of coastal cordillera 

Warszewiczia 7 N, CR, Pa, Co, V, Gu, E, Pe, Lowland rain forests, cloud forests, and Amazonian ra 

Br, Bo forests 

'Ar, Argentina; Ba, Bahama Archipelago; Be, Belize; Bo, Bolivia; Br, Brazil; Ca, Cayman Islands; Co, Colombia; CR, Costa R 

ES, El Salvador; G, Guatemala; Gu, The Guianas; Ha, Haiti; Ho, Honduras; J, Jamaica; LA, Lesser Antilles; M, Mexico; N, N 

United States; TT, Trinidad and Tobago; V, Venezuela; VI, Virgin Islands.


Stevensia Poit. was described by Pierre Antoine is exceptional in having 2-seeded pseudosamaras. 

Poiteau (1804). Since then, several species have been Future flowering collections will shed light on the 

added and this genus is currently represented by nine identity and systematic relationships of this taxon. 

species endemic to Haiti and the Dominican Republic. 

Liogier (1995) recently published an updated de- 

The Simira Group 

scription of Stevensia and a key to all the species. 

Acrosynanthus Urb. was described by Urban 

Simira was established by Aublet (1775), and 

(1913) as a monotypic genus with 6-merous flowers. Sickingia by Willdenow (1801), with Sickingia 

Additional species described by other authors and erythroxylon Willd. and Sickingia longifolia Willd. 

himself (Standley, 1918; Urban, 1923; Howard & 

Baillon (1879) wrongly reduced Sickingia to a sub- 

Proctor, 1958) proved that this taxon has 4-5(-6)genus 

of Chimarrhis (see Excluded Species in the 

merous flowers. 

treatment of Chimarrhis, below). Bremekamp (1954a) 

convincingly demonstrated that Sickingia and Simira 

are congeneric, and transferred the two Willdenowian 

The Augusta Group 

species to Simira. Steyermark (1972), following 

Lindenia was described by Bentham (1841), and Bremekamp's conclusion, transferred all the South 

a detailed account of the taxonomic history and mono- American species of Sickingia to Simira. Bremekamp 

graphic treatment of this taxon was published by Dar- (1966) also separated Simira from the Rondeletieae 

win (1976). Augusta was established by Pohl (1828- because of the morphology and insertion of its seeds, 

1829), under which he described five species endemic establishing the tribe Simireae. Robbrecht (1988) into 

Brazil. The complex taxonomic history of the Bra- cluded the tribe Simireae in the Rondeletieae but a few 

zilian taxa of Augusta has been summarized by years later (Robbrecht, 1993) reestablished the tribe. 

Delprete (1997a), who treated Pohl's five species as In the current treatment the Simireae are included in 

synonyms ofAugusta longifolia (Spreng.) Rehder. On the Rondeletieae according to a cladistic analysis usthe 

basis of its rheophytic habit, fruit dehiscence, and ing morphological characters (Delprete, 1996d). Reinflorescence 

position and architecture, Kirkbride cent accounts of Simira from selected Brazilian states 

(1997b) included this taxon in Augusta, as subgen. were published by Peixoto (1981, 1982) and Barbosa 

Lindenia. Kirkbride attributed the differences in the and Peixoto (1989). 

floral morphology of the two taxa to a pollination Flexanthera was established by Rusby (1927). In 

syndrome. With Kirkbride's inclusion, Augusta is the protologue he stated that Flexanthera is "allied to 

represented by four species of rheophytic shrubs dis- Rustia [but without discussing their similarities],... 

tributed in Central America, Brazil, and islands of the from which it differs in its smaller flowers, reflexed 

South Pacific (Fiji and New Caledonia). 

and recurved dehiscent anthers [poricidal in Rustia] 

and deciduous calyx-limb and in the character of the 

Eosanthe 

placentae." Standley (1930a, 1931b) transferred the 

two known species of Flexanthera to Sickingia (Ron- 

Eosanthe was established by Urban (1923), who deletieae) without further comments, and Steyermark 

related it to Phyllomelia. Eosanthe is a monotypic (1972) transferred the species of Sickingia to Simira. 

genus endemic to the serpentine soils of Sierra de Flexantherafragrans Rusby and F. subcordata Rusby 

Cristal, Cuba (Borhidi, 1991; Delprete, 1999a). Its 4- are synonyms with Simirafragrans (Rusby) Steyerm. 

lobed persistent calyx functions in a similar manner and Simira cordifolia (J. D. Hook.) Steyerm., respecas 

the pterophyllous rotate calyx of Phyllomelia but, tively, and Flexanthera is synonymous with Simira. 

because of its general habit (small multi-stemmed The monotypic Exandra Standl. was described by 

shrubs; cf. Borhidi, 1991), ridged corolla tube and Standley (1923). This taxon was later treated by 

thick-coriaceous leaves, Eosanthe resembles Schmidt- Steyermark (1972) as congeneric with Simira, to which 

ottia (Catesbaeeae sensu Delprete, 1996d). On the it was transferred as S. rhodoclada (Standl.) Steyerm. 

other hand, the sheathing stipules, axillary inflores- The monotypic Holtonia Standl. was established 

cences, persistent 4-lobed calyx, filaments not by Standley (1932) with a species that he previously 

connated to the corolla tube, and 2-seeded indehiscent described in Sickingia (S. myriantha; Standley, 

fruits of Eosanthe resemble those of Phialanthus 1930a). He suggested that Holtonia is closely related 

(Chiococceae sensu Delprete, 1996d). In conclusion, to Chimarrhis and therefore belonged in the Condabecause 

of its resemblance to both Schmidtottia and mineeae (sensu Hooker). Holtonia has recently been 

Phialanthus, Eosanthe is here tentatively placed in the transferred to Elaeagia (Burger & Taylor, 1993) and 

Catesbaeeae-Chiococceae-complex, within which it a new combination [E. myriantha (Standl.) C. M.


Taylor & Hammel] created, a decision with which I subglobose), both characters of little taxonomic sigagree. 

The suggested relationship with Chimarrhis nificance. General evidence strongly suggests that the 

(Standley, 1932) was appropriate, and corresponds to two taxa are congeneric, but the two species of 

my idea (Delprete, 1996b) that Elaeagia and Bathysa Pteridocalyx are known only from single flowering 

are closely related to Chimarrhis. 

specimens. Their mature fruits are unknown, and 

Wernhamia (Moore, 1922), traditionally placed in therefore fruiting collections are necessary to confirm 

the Cinchoneae and transferred to the Condamineeae their identity. Due to its probable synonymy with 

by Andersson and Persson (1991), has recently been Pteridocalyx, Neobertiera is not included in the key 

synonymized with Simira by Delprete and Nee (1997). to neotropical Rondeletieae given below. 

The taxa Wernhamia boliviensis S. Moore and Simira 

catappifolia (Standl.) Steyerm. are synonyms of 

Standleya and Bradea 

Simira macrocrater (K. Schum.) Steyerm. 

Blandibractea (Wernham, 1917), after careful The Brazilian endemics Standleya Brade and 

morphological comparison, was also treated as con- Bradea Standl. are closely related, and both belong 

generic with Simira by Delprete (1998c), B. to the Hedyotideae. They are similar in having tetbrasiliensis 

being synonymous with S. glaziovii (K. 

ramerous flowers, campanulate corollas, stamens in- 

Schum.) Steyerm. 

serted about the middle of the corolla tube, linear to 

ovate calyx lobes, few-seeded capsules, and ovoid 

seeds attached to a stalked 

The 

placenta. Standleya was 

Sipanea Group 

listed in the Rondeletieae by Robbrecht (1988, 1993) 

Limnosipanea, Sipanea, and Steyermarkia were and Delprete (1996d), and Bradea in the Hedyotideae 

transferred by Bremekamp (1934) from the Rondele- by Robbrecht (1988, 1993). Standleya differs from 

tieae to his newly founded tribe Sipaneeae. The only Bradea in having herbaceous habit (vs. subfruticose) 

morphological element that makes Sipanea and and unwinged seeds with tuberculate testa (vs. shal- 

Limnosipanea exceptional within the Rondeletieae is lowly winged with smooth testa). The tribal delimitheir 

herbaceous habit. Sipaneopsis and Neobertiera tation of the Hedyotideae, and their supposed relation- 

Wernham have habits that range from herbaceous (10 ships with the Rondeletieae are still uncertain, and 

cm tall) to subfrutescent (to 2-4 m tall), and could be need to be carefully analyzed. 

interpreted as the phylogenetic links between the 

Sipaneeae and the Rondeletieae s.str. Robbrecht 

Eizia 

(1988, 1993) maintained the Sipaneeae provisionally 

separate from the Rondeletieae because of their her- Eizia was listed by Delprete (1996d) in the Ronbaceous 

habit. In agreement with Verdcourt (1958) deletieae, but Lorence and Dwyer's (1988) observaand 

Steyermark (1974), the tribe Sipaneeae is here tions and recent personal studies have convinced me 

included in a broad circumscription of the Rondele- that this monotypic genus is better placed in the 

tieae. 

Hamelieae. 

Neobertiera and Pteridocalyx 

Pinckneya 

The monotypic Neobertiera was described by Pinckneya (Richard, 1803) is a monotypic genus 

Wernham (1917), who erroneously stated that its fruits endemic to the southwestern United States. It is the 

were berries and placed it in the Hamelieae. only North American genus included in the Rondele- 

Steyermark (1967: 289-290) corrected Wernham's tieae. Two Brazilian species, P. rubescens Allemao 

errors, declaring that the fruits of Neobertiera are & Saldanha and P. viridiflora Allemao & Saldanha, 

many-seeded capsules, and transferred the genus to were attributed to Pinckneya (Allemao & Saldanha da 

the Rondeletieae. The similarities between Gama, in Saldanha da Gama, 1872), and a third Bra- 

Neobertiera and Pteridocalyx Wernham (1911, 1913) zilian species, P. erubescens, was added to this geare 

striking, both having dichasial-cymose inflores- nus by Glaziou [in herb.]. Schumann (1888) transcences 

terminating in monochasial branchlets, con- ferred the Allemao & Saldanha species to Sickingia 

torted corolla aestivation, many-seeded capsules, [as S. rubra (Martius) K. Schum. and S. viridiflora 

subglobose seeds with reticulate exotesta, and both (Allemao & Saldanha) K. Schum.], and synonymized 

coming from low-elevation Guyanan rain forests; the Glaziou species under S. glaziovii K. Schum. 

Pteridocalyx differs from Neobertiera by the presence Steyermark (1972) transferred these three species to 

of calycophylls and by narrow, elongated fruits (vs. Simira, as S. glaziovii (K. Schum.) Steyerm., S. rubra


(Martius) Steyerm., and S. viridiflora (Allemao & 

Saldanha) Steyerm., respectively, leaving the one remaining 

species, P. bracteata (W. Bartr.) Raf., as a 

North American endemic. Additional information can 

be found in Delprete, 1996c. 

Steenisia (5 spp.) from Borneo, and Wendlandia (ca. 

56 spp.) widespread in Indomalesia. Augusta (including 

Lindenia) is the only genus present in both the 

Neotropics and Paleotropics, with one species in Central 

America, one endemic to Brazil, and two species 

in the Pacific Islands. The paleotropical Gloneria was 

Kerianthera 

listed by Delprete (1996d) in the Rondeletieae, but it 

is here excluded because recent personal observations 

Kerianthera J. H. Kirkbr. is a monotypic genus demonstrated that this genus is synonymous with 

endemic to the Brazilian Amazon. It was placed in the 

Psychotria L. (Psychotrieae). 

Condamineeae by Kirkbride (1985b) but, after phylogenetic 

and morphological studies, was transferred 

to the Isertieae by Delprete (1996e). 

Rondeletieae (A. P. de Candolle) Miquel, Flora 

Nederl. Indie 2: 130, 156. 1856. Rondeletiinae A. 

The Paleotropical Genera 

P. de Candolle (as "Rondeletieae," tribus 

Pseudomussaenda (5 spp., endemic to Africa) was 

Hedyotideae), Prodr. 342, 401. 1830. Rondeletieae 

A. P. de Candolle ex Reichenbach, stat. non indic., 

placed in the Condamineeae-Pinckneyinae by Der Deutsche Botaniker 1: 77. 1841. 

Wernham (1916) and later transferred by Verdcourt 

Type genus. 

Rondeletia Linnaeus. 

(1958) to the Isertieae. This transfer has recently been 

supported by Puff et al. (1993), who suggested a close Condamineeae J. D. Hooker (as "Condaminieae"), Gen. 

relationship to Schizomussaenda H. L. Li and PI. 8, 12. 1873. Condamineinae J. D. Hooker (as 

Mussaenda. Pseudomussaenda, Schizomussaenda, "Eucondaminieae," nom. inadmiss.), Gen. P1. 12. 

Mussaenda, and Kerianthera are exceptional within 1873. Type genus. Condaminea A. P. de Candolle. 

the Isertieae in having capsular fruits and Pinckneyinae J. D. Hooker (as "Pinckneyeae"), Gen. PI. 

calycophylls. 

12. 1873. Type genus. Pinckneya L. C. M. Richard 

Kajewskiella (endemic to the Solomon in 

Islands) 

A. Michaux. 

was established by Merrill and Perry (1947). It was Chimarrheae Bremekamp, nom. nud., Rec. Trav. Bot. 

Neerl. 31: 253. 1934. 

tentatively placed in 

Chimarrhidoideae 

the Condamineeae by Jansen 

Rafinesque 

(as "Sous-famille 

(1978) because of its valvate aestivation, absence of 

Chimarhidia"), Ann. G6n. Sci. 

Phys. 6: 84. 1820. Type genus. Chimarrhis Jacquin. 

raphides, and seed shape and position. Tange (1995) Simireae Bremekamp ex S. Darwin, Taxon 25: 606. 

found raphides in both species of Kajewskiella, which 1976. Simireae Bremekamp, sine descr. lat., Acta 

are in turn "indistinguishable from those found in Bot. Neerl. 15: 22. 1966. Type genus. Simira Aublet. 

Xanthophytum [Reinw. ex Blume]. These observa- Sipaneeae Bremekamp (as "Sipaneae"), Rec. Trav. Bot. 

tions lead to the conclusion that Kajewskiella belongs Neerl. 31: 253. 1934. Type genus. Sipanea Aublet. 

to the tribe Hedyotideae as defined by Robbrecht 

(1988), where together with Pomazota [Ridl.], Shrubs to tall canopy trees, rarely herbs or sub- 

Xanthophytum and Lerchea [L.] probably forms a shrubs; wood whitish, yellowish, or reddish; bark ofnatural 

group." (Tange, 1995). After studying flower ten containing quinine-related compounds. Raphides 

and seed morphology of Kajewskiella, I agree with absent. Stipules mostly entire, rarely bifid (four stipu- 

Tange that this genus should be included in the lar units in Condaminea), mostly interpetiolar or rarely 

Hedyotideae. 

intrapetiolar (Elaeagia), frequently with colleters on 

Badusa (Pacific Islands; cf. Soejarto et al., 1996) the adaxial side secreting resinous compounds, perwas 

transferred from the Cinchoneae to the Conda- sistent to readily caducous. Leaves opposite, decusmineeae 

(subtr. Portlandiinae) by Ridsdale (1982). sate (rarely in whorls of 3-4 leaves per node in a few 

Delprete (1996d) treated Badusa as related to the herbaceous taxa), petiolate to sessile; blades folia- 

Catesbaeeae sensu Delprete. Recent additional infor- ceous to thick-coriaceous; domatia variably present. 

mation about this genus can be found in Soejarto et Inflorescences terminal or axillary, often with 2-8 

al., 1996. 

orders of branching. Flowers hermaphroditic (dioe- 

The paleotropical genera belonging to the Ronde- cious in Dioicodendron), mostly actinomorphic (zyletieae, 

as presently circumscribed, are Aleisanthia (2 gomorphic in Molopanthera, and subzygomorphic in 

spp.) from the Malay Peninsula, Glionnetia (1 sp.) Cuatrecasasiodendron), 4-6-merous. Calyx persisfrom 

the Seychelles Islands, Greenea (12 spp.) from tent or caducous; lobes often minute, sometimes foli- 

Indomalesia, Spathichlamys (1 sp.) endemic to Burma, ose, frequently expanded into calycophylls. Corolla


white, red, green, or yellow, membranous to fleshy; 

aestivation valvate, contorted, or imbricate. Stamens 

mostly as many as corolla lobes, inserted near the base, 

on the medial zone, or near the orifice of corolla tube; 

anthers included or exserted, oblong to narrowly el- 

liptic to button-shaped, 2-locular, commonly opening 

by longitudinal slits or rarely by terminal pores (Rustia 

and Tresanthera), dorsifixed near the base or around 

the middle, introrse. Pollen exine reticulate or foveolate 

(not echinate). Style branches present, with stigmatic 

surface smooth to verrucate. Ovary inferior, 

bilocular, with few to many ovules per locule attached 

to a central placenta, or exceptionally 1 ovule per locule 

apically attached (Phyllomelia). Fruits woody, 

loculicidal or septicidal capsules, mostly opening 

basipetally (opening in central portion in Picardaea), 

or exceptionally pseudosamaras (Phyllomelia); pla- 

centa central, rarely apically incomplete, or shortly 

stalked. Seeds horizontal, imbricate, or peltate and 

vertical; winged, with lateral (Simira and 

Parachimarrhis) or concentric wing, or unwinged 

(apically inserted, ellipsoid-ovoid in Phyllomelia). 

The Rondeletieae and Condamineeae have traditionally 

been described as having many-seeded capsules. 

Nevertheless, the inclusion of Phyllomelia (related 

to Mazaea, see below), brings a new fruit type 

into the delimitation of the Rondeletieae: the 

pseudosamara (Spjut, 1994). In Phyllomelia the 2- 

seeded indehiscent fruits, with a persistent, rotate 

calyx, detach at maturity and function as a dispersal 

unit. I regard this fruit type as being the result of a 

derived set of characters that contributed to the adaptation 

to wind dispersal into this tribe. 

KEY TO THE GENERA OF NEOTROPICAL RONDELETIEAE S.L. 

1. Herbs to small subshrubs, usually 15 seeds per fruit; seeds angular, polyhedral; mouth of the 

corolla without triangular projections. 

4. Anthers exserted beyond the corolla; leaves in whorls of 3 or more per node (but leaves in 

pairs in L. schomburgkii J. D. Hook.) (Amazon basin) ............................................................. Limnosipanea 

4. Anthers included within the corolla; leaves in pairs (mostly South America, except S. biflora L.f. 

in Central America) ............................................................................. Sipanea 

1. Small shrubs to large canopy trees, >1 m tall. 

5. Stipules 4 per node, 2 intrapetiolar and 2 interpetiolar (Costa Rica to Bolivia) .................................... Condaminea 

5. Stipules 2 per node. 

6. Stipules intrapetiolar or calyptrate, mostly copiously exuding resin when still in bud, in several 

species solidifying into a globose mass at terminal bud (Central and South America, and E. cubensis 

Britt. endemic to Cuba) .............................................................................. Elaeagia 

6. Stipules interpetiolar, mostly scantly exuding resin when still in bud (copiously exuding resin in 

Chalepophyllum), not solidifying into a globose mass at terminal bud. 

7. One or more calyx lobes of some flowers expanded into 1 or 2 foliose extensions (calycophylls). 

8. Inflorescences axillary, subterminal, in pairs at the last two nodes below the apical bud, 

caducous at fruit maturity; seeds peltate on the central placenta, with central hilum (a few 

South American species with calycophylls) ............................................................. Chimarrhis 

8. Inflorescences terminal and solitary, persistent after fruit maturity; seeds not peltate on a 

central placenta, with lateral hilum. 

9. Corollas 2.5-8 cm long, long-tubular. 

10. Corolla tube with a ring of hairs at base inside; corolla lobes glabrous within; 

seeds not winged (mountains of Central and South America) .................................. Pogonopus 

10. Corolla tube without a ring of hairs inside; corolla lobes pubescent within; 

seeds winged (swamp areas of SE United States) ....................................................... Pinckneya 

9. Corollas 2 cm long; wood usually turning reddish when 

exposed to air (calycophylls present in several South American species) ...................... Simira


11. Seeds smaller,


mouth of the tube, partially exserted; corolla tubes 

green and lobes red throughout anthesis; capsules 

thin-woody, oblong, 2.5-3.5 cm long; seed tetrahedral, 

1-2 mm long, unwinged (Amazonian slopes of the 

Andes from Colombia to Peru) ........................................ Macbrideina 

23. Corollas 2 cm 

long; wood usually turning reddish when exposed to 

air (Central and South America) . ........ ......................... Simira 

26. Seeds small, 2-3.5 mm long; capsules small,


entire margin; stipules obovate to ellipticobovate, 

15-30 mm long, foliaceous; seeds 

prominently winged (Colombia) 

......................................... Cuatrecasasiodendron 

33. Corollas actinomorphic, 4-5-lobed; the 

innermost lobe equal in size to the other, 

with entire margin; the outer 1 to 2 lobes 

undulate-crenulate; stipules deltoid to 

lanceolate, 3 cm long. 

35. Corollas narrowly campanulate, red; corolla tube narrowly imbutiform 

(Cerrado and Atlantic forests of Brazil) ... Augusta subgen. Augusta 

35. Corollas hypocrateriform, white; corolla tube narrowly tubular. 

36. Calyx lobes linear; seeds with lateral hilum (truncate at apex), 

not winged, horizontally attached to the central placenta 

(Central America and South Pacific Islands) .. Augusta subgen. Lindenia 

36. Calyx lobes oblong-obovate; seeds with central hilum, 

with shallow concentric wing, vertical, peltately attached 

to the central placenta (Venezuelan tepuis) ........... Maguireothamnus 

34. Corollas 2 x 1 cm. 

38. Ovary 4-locular, with 5-6 ovules per locule; mature 

fruits dividing into 4 cocci (Colombia) .............................. Acrobotrys 

38. Ovary 2-locular, with numerous ovules per locule (>6); 

mature fruits not dividing into 4 cocci (but old capsules of 

Chalepophyllum 

splitting into 4 parts, though not into 4 cocci). 

39. Inflorescences axillary (not subterminal), at the axils 

of the last 5-7 nodes below the apical bud, in congested 

cymules of 1-4 flowers. 

40. Corollas hypocrateriform, white (the tubes 

sometimes suffused with pinkish-reddish tinge 

at base); resinous exudates from young stipules 

copious; calyx lobes narrowly oblong to narrowly 

lanceolate, subcoriaceous, unequal, 1-3 mm wide, 

fully expanded before anthesis; seeds with a narrow 

marginal expanded wing, 3-4 mm long (high 

elevations of Venezuela and Guianas)......... Chalepophyllum 

40. Corollas narrowly campanulate, salmon-red; 

resinous exudates from young stipules scant; 

calyx lobes lanceolate, membranous, subequal, 

4-6 mm wide,expanding after anthesis; seeds 

unwinged, to 1 mm long (high elevations of 

Amazonian Venezuela) ...................................... Holstianthus 

39. Inflorescences terminal, in corymbose panicles of 

9-40 flowers. 

41. Inflorescences 10 cm long; flowers subcampanu- 

late; calyx lobes widely ovate; stipules readily 

caducous (Colombia and Peru) ................................. Phitopis

Introduction to the Neotropical Genera Studied 23 

NOTES ON DESCRIPTIONS, (20-30 m) with much expanded buttresses in South 

ABBREVIATIONS, AND ACRONYMS American species. Some Amazonian species of 

Chimarrhis (C. glabriflora, C. hookeri, C. barbata, 

In the preparation of herbarium specimens, veg- and C. turbinata) can grow into forest giants, 40-50 

etative and reproductive parts are subjected to minor m tall and 2 m diam. 

shrinkage. Measurements given in keys and descrip- The tree architecture of this group has been studtions 

have been obtained from such specimens. Col- ied in only a few species of Chimarrhis. Halle et al. 

ors are from personal observations and label data, (1978) reported that C. cymosa (from the Lesser 

unless otherwise indicated. The abbreviation "BA" 

Antilles) has a Petit model of branching, while C. 

represents the basal angle from the leaf margin to the 

microcarpa (from eastern Amazon) has an Aubreville 

midrib of the leaf blade, and "L/W" symbolizes the model ("Terminalia branching"). I have observed C. 

length to width ratio of the leaf blades. Nomenclature 

glabriflora and C. hookeri (both from western Amaof 

leaf domatia follows Robbrecht (1988: fig. 12), and zon forest), and both have Petit's model. 

that of vestiture types follows Radford et al. (1974: Large buttresses are present only in the large trees 

140-141). In the descriptions of inflorescence archi- of the Amazonian species of Chimarrhis, extending 

tectures, "L/A" is the ratio of the total length (L) to to 4 m in height near the trunk and expanding outthe 

length of the branched portion (A). The general ward 8-10 m from the trunk (per. obs.). Additionally, 

shape and vestiture of the disk, and whether the disk at least two arborescent species of Rustia (R. alba and 

exceeds the calyx, refer to mature capsules, unless R. rubra) can develop small buttresses. 

otherwise indicated. The term "microscopically (40x)" Lateral roots are occasionally present in the genmeans 

that the structures or characteristics referred to era studied. Rustia occidentalis, R. thibaudioides, and 

can be seen using a dissecting microscope at 40x 

Pogonopus speciosus (all multistemmed shrubs growmagnification. 

All specimens cited, including types, ing in proximity to streams or standing freshwater) 

have been seen unless indicated by "n.v." immediately have been personally observed to produce lateral roots 

following the citation. Specimens cited as "B*" were at the basal nodes of the external branches. The three 

destroyed in the Berlin herbarium during the Second 

species of Pogonopus are very ornamental shrubs due 

World War (Hiepko, 1987). Herbarium acronyms to their colourful calycophylls, and their propagation 

follow Holmgren et al., 1990. Authors of plant names could be easily accomplished through cuttings of 

have been abbreviated throughout the text according young woody branches. 

to Brummitt & Powell, 1992. Genera included in the 

present systematic treatment are in boldface in the key. 

Leafy Branchlets 

INTRODUCTION TO THE 

Leafy branchlets are here referred to as the terminal 

NEOTROPICAL GENERA STUDIED portion of the vegetative shoots of the last nodes 

(commonly known as "twigs"). Leafy branchlets are 

GENERAL MORPHOLOGY commonly terete in all genera, with Dioicodendron 

being exceptional in having subtetragonal leafy 

Habit 

branchlets. Young leafy branchlets vary among spe- 

The Condamineeae sensu Hooker (Robbrecht, cies of the same genus in being either thin and woody 

1988, 1993) is a paraphyletic group of genera repre- or thick and semisucculent. I consider these two catsented 

by shrubs, small trees, and tall canopy trees egories of vegetative shoots as a reliable taxonomic 

(i.e., Chimarrhis). The genera Pogonopus, Condaminea, character, which is quite consistent at the specific 

Picardaea, and Rustia are most commonly found as level, mostly in Chimarrhis and secondarily in Rustia 

shrubs. Several species of Rustia (R. costaricensis, R. and Tresanthera. 

dressleri, R. alba, R. bilsana, R. rubra, R. viridiflora, 

R. venezuelensis, and rarely R. formosa), Dioico- 

External Indumentum 

dendron, Dolichodelphys, and Molopanthera are usually 

encountered as medium-sized trees 10-15 m tall The indumentum of the genera here treated, when 

(exceptionally to 20-30 m). Tresanthera is usually present, consists of unicellular, nonglandular hairs. It 

encountered as pyramidal single-stemmed trees. is extremely variable among and within populations 

Trees of Chimarrhis vary from medium-sized [10- of the same species (Rustia thibaudioides, R. alba, 

20(-30) m tall] in Central America and the Caribbean pers. obs.; Condaminea corymbosa, pers. obs.; Pogon- 

Islands (C. parviflora, C. latifolia, C. cubensis, C. opus speciosus var. sandwithianus, pers. obs.; Pinckjamaicensis, 

and C. cymosa) to large canopy-trees neya bracteata, pers. obs.) and even among the foli-


age of a single individual (Rustia thibaudioides, pers. 

Domatia 

obs.; Condaminea corymbosa, pers. obs.; Pogonopus Domatia are here defined as presence of hairs, 

speciosus, pers. obs.; Chimarrhis hookeri, pers. obs.). correlated or not with a pit or a crypt, in the axils of 

For the above reasons the occurrence, density, and 

the secondary veins, or rarely of the tertiary veins of 

quality of the indumentum have been largely considthe 

leaf blades. Domatia are commonly associated 

ered characters of little taxonomical significance. with the presence of mites. In many of the species of 

Almost all the genera studied have species with wide the genera studied, larval and adult stages of unidengeographic 

distributions that have been described as tified mites have been found. 

a complex of species based on their variable The general classification of leaf-domatia in the 

indumentum. Standley (1918, 1921, 1930a, 1930b, 

present treatment (and in the terminology used in the 

1931 a, 1931b, 1931 c, 1934a, 1934b) and Steyermark 

species descriptions) is in accordance with Robbrecht's 

(1963, 1964b, 1965, 1967, 1972, 1974, 1975) are 

typology (1988: 50). 

among the many botanists who have especially em- 

Leaf domatia have not been found (and are most 

phasized the density and quality of the indumentum 

probably absent) in Tresanthera, Condaminea, Dioof 

vegetative parts in describing new species and new 

icodendron, Molopanthera, and Dolichodelphys; and 

varieties of Rubiaceae. Nevertheless, in some very are found to be variably absent or present in Rustia, 

special cases, where a good number of collections 

Pogonopus, Chimarrhis, and Picardaea. The absence 

were available, and very distinct forms of indument or presence, morphology, and vestiture ofdomatia was 

were consistently present and were correlated with a 

studied in detail and found to be very variable within 

particular area, I have used vestiture as a secondary 

species, within populations of the same species, and 

taxonomic character at the varietal rank. 

even within the same individual. Domatia are therefore 

considered a character of secondary taxonomical 

Leaves 

importance, to be used, at the most, at the specific level. 

The leaves of woody Rubiaceae are mostly oppo- Nevertheless, several species of Chimarrhis dissite, 

rarely ternate or whorled, with stipules constantly play a certain degree of consistency in the type of 

present, and entire blades; these three vegetative fea- domatia, and some species can be recognized solely 

tures make the family easy to recognize even with by the presence of their characteristic domatia (see 

sterile material. Opposite leaves are usually decussate. Fig. 58). The most typical domatia are those of C.ja- 

In the genera here treated, the leaves are opposite, maicensis, unique in the genus in being tuft-pit domatia, 

decussate, and petiolate. The leaves of Condaminea with hairs departing from the veins and minutely pitcorymbosa 

are exceptional, varying from sessile (in ted at the axils of secondary veins, and rarely of ter- 

Central America) to petiolate (in South America) (see tiary veins. Chimarrhis cubensis has minutely winged 

Fig. 47). Petioles are mostly adaxially flattened to veins, forming "pocket-domatia" in axils of secondconcave, 

thickened at the base, and apulvinate. How- ary veins. In C. latifolia, C. brevipes, C. gentryana, and 

ever, in two species of Rustia (R. occidentalis and R. C. barbata the blades and the veins are entirely glathibaudioides) 

I have observed rudimentary pulvini brous, and no domatia have been observed. In the rest 

on petioles of young leaves, which become thicker of the species of Chimarrhis, domatia were variably 

(sometimes corky) and nonfunctional on older leaves. present or absent and of various kinds (see Fig. 58). 

Blades are always entire and mostly stiff-foliaceous, 

but coriaceous in Condaminea, and semicoriaceous 

Foliar Pellucid Glands 

in Tresanthera and in some species of Rustia (R. Solereder (1893), in his Anatomischen Charakdressleri, 

R. alba, R. bilsana, R. venezuelensis, R. teristik und zur Systematik, reported the presence of 

formosa, R. simpsonii, R. angustifolia, and variably "inneren Drilsen" [internal druses] in the Rubiaceae, 

so in R. thibaudioides). 

which he found previously described as "intercellu- 

Venation of all the genera studied is brochido- lare Secretbehilter" ["intercellular secretion-containdromous, 

with secondary veins ascending, arcuate, ers"], "secretfiihrende" [secretory cells] of differing 

and anastomosing near the blade margins. Bacterial forms and various contents, and also as "eingenthiimnodules 

are found in blades of Rustia (R. occidentalis), liche Drisen" ["characteristic druses"] present in 

Pogonopus (P. speciosus var. sandwithianus, being some taxa in the form of secretory cells. Such secrepunctate 

on the adaxial surface due to bacteria), and tory glands have recently been called "pellucid 

Chimarrhis (C. glabriflora and C. hookeri). To my glands" (Steyermark, 1974; Dwyer, 1980), "transluknowledge, 

these are the first reports of bacterial nod- cent dots" (Robbrecht, 1988), or "pellucid-dots" 

ules in neotropical Rubiaceae. 

(Burger & Taylor, 1993).


In the Rubiaceae, schizogenous secretory cavities, biaceae. Unfortunately, stipules are sometimes readily 

filled with a resinous secretion, have been found only caducous, and in that case they can only be found at 

in the leaves of the closely related genera Rustia and the youngest nodes (and sometimes only at the apical 

Tresanthera (which are also unique in the family in bud). I found that caducous or persistent stipules are 

having poricidal anthers). To my knowledge no de- consistent within each genus of the Condamineeae 

tailed work has been done on the anatomy of such sensu Hooker (except for Chimarrhis; see below). For 

secretory structures, nor has any attempt been made descriptive purposes the stipule persistency is here 

to identify the content of their cavities (Delprete et divided into three degrees: persistent (present after 

al., in progress). The only overview on pellucid-punc- leaves of the same node fall off), readily caducous 

tate leaves was presented by Solereder (1893: 279; (present only at the vegetative bud), and caducous 

free translation from German) as follows: (present during leaf development but caducous before 

leaves fall 

Intercellular secretory space, in the form of 

off). Of the genera here presented, the 

schizogenous secretory cavities filled with 

stipules are readily caducous in Rustia and Tressticky 

[resinous] secretion, lined with a thin 

anthera, caducous in Condaminea and Pinckneya, and 

walled epithelium, are found in Rustia persistent in Pogonopus, Picardaea, Dioicodendron, 

(incl. 

Henlea) and Tresanthera, which were united Molopanthera, and Dolichodelphys. Chimarrhis is 

by Bentham and Hooker [1873] in a exceptional to the group in having most species with 

single genus 

(Rustia), which I already noted briefly in stipules readily caducous (subgen. Chimarrhis), and 

my paper on Hymenocnemis in Bot. Centralbl. 

a few species with persistent stipules (subgen. 

1891. These secretory cavities occur in the spe- 

Pseudochimarrhis). 

cies of Rustia and Tresanthera in large leaves, Stipular position is also an important taxonomic 

usually as translucent points, which were alcharacter 

in the Rubiaceae. All the genera here treated 

ready described by Karsten [1858, 1861; cf. 

have two interpetiolar stipules, except for Conda- 

Karsten, 1858-1869] for Rustia in general, for minea, which is commonly described as having four 

Tresanthera condamineoides and Henlea interpetiolar and intrapetiolar stipules. The nature of 

splendens [= Rustia thibaudioides], and the 

by stipules in Condaminea is rather ambiguous, be- 

Bentham [1844] for Exostem[m]a occidentale ing all basally connate and not truly intrapetiolar (Fig. 

justly transferred to Rustia by Hemsley [1881 ]), 3F-H), because no stipules are positioned between the 

and by Schumann (1889, 1891) [incorrectly de- stem and the petioles. The stipules of Condaminea 

scribed as "characteristic fat bodies"] for could also be interpreted as two interpetiolar stipules 

Rustia. I have described the secretory cavities highly dissected and connate at the base. To underfor 

certain available species, namely T. conda- stand fully the position of stipules in Condaminea, 

mineoides, R. angustifolia, R.formosa, R. gra- developmental studies are needed. 

cilis, R. occidentalis, R. secundiflora, R. The most common shape of the stipules is trianwarczewicziana 

[two former = R. thi- gular with entire margins (Fig. 3), varying from delbaudioides], 

as well as R. pauciflora [= T. toid to narrowly triangular, with the exception of 

pauciflora] described by me, which Schumann Condaminea (narrowly lanceolate and foliose) and 

[in contrast to Bentham & Hooker (1873)] Dioicodendron (obovate stipules). Stipule size in the 

treated in a distinct genus, Tresanthera genera here studied varies from small and scale-like 

pauciflora. 

(Picardaea) to small and deltoid (Pogonopus, Pinckneya, 

Molopanthera, and Chimarrhis turbinata) to 

I examined all the species of Rustia (incl. Henlea) medium-size and narrowly triangular (Rustia, most 

and Tresanthera using light microscopy, and all of 

species of Chimarrhis, and Dolichodelphys) to large 

them have pellucid glands. Such glands represent a 

and foliose (Tresanthera, Condaminea, Chimarrhis 

good field character to easily identify these two rubicymosa). 

aceous genera. The density and size of the pellucid 

glands in the blades is variable (rarely consistent 

within the same species), and for the most part it is 

Colleters 

not a reliable taxonomic character. 

The term colleters is "derived from the Greek Colla 

[= glue], referring to the sticky excretion from this 

Stipules 

structure," and "they frequently produce a mixture of 

Stipules are the paramount feature of the family terpenes and mucilage" (Esau, 1965). Colleters are 

and represent one of the easiest and most apparent commonly defined as "pluricellular mucilage-secretfield 

characters for recognizing members of the Ru- ing structures occurring inside stipules and on other

26 

A 

' 


FIG. 3. Stipule and colleter variation in the genera referred to the Condamineeae (sensu Hooker, 1873). A-C. Rustia. 

A. R. occidentalis (Delprete 5267, TEX), adaxial view. B. R. alba (Delprete & Verduga 6420, TEX), adaxial view. C. R.formosa 

(Kirkbride 5431, F), adaxial view. D. Kerianthera preclara (Prance et al. 24293, NY), adaxial view. E. Tresanthera 

condamineoides var. condamineoides (Steyermark & Rabe 96163, US), adaxial view. F-H. Condaminea corymbosa (Delprete 

& Webster 6104, TEX). F. View from above showing stipular insertion. G. Detail of base of stipule showing colleters in 

adaxial view. H. Habit of stipules. I, J. Pinckneya bracteata (Delprete & Sharif 6472, TEX). I. Adaxial view. J. Lateral 

view of stipule insertion (white area is petiole attachment). K, L. Pogonopus speciosus var. sandwithianus (Delprete & 

Apreza 6359, TEX). K. Adaxial view. L. Lateral view of stipule insertion and tuft of hairs in stipules' axils (white area is petiole 

attachment). M. Picardaea cubensis (Shafer 7850, NY), adaxial view. N. Kajewskiella trichantha (Kajewski 1667, A), 

adaxial view. O-Q. Chimarrhis. 0. C. cymosa (Webster et al. 9205, DAV), adaxial view. P. C. glabriflora (Delprete & 

Verduga 6423, TEX), abaxial view. Q. C. turbinata (Irwin et al. 55443, GH), adaxial view. R. Dioicodendron dioicum 

(Hammered et al. 5233, NY), adaxial view. S. Dolichodelphys chlorocrater (Ramirez & Cardenas 1017, COL), adaxial view. T. 

Molopanthera paniculata var. paniculata (Mori et al. 10328, NY), adaxial view. All drawn to the same scale, except G.


organs" (Robbrecht, 1988). Additional information 

Inflorescences 

about colleter structure and function can be found in The terminology and classification of inflores- 

Esau, 1965 (311); Halle, 1967 (95-97); Lersten, cences in the present work has been heavily influenced 

1974a, 1974b, 1975; Robbrecht, 1988 (64-68); and by the recent work of Weberling (1992). More spe- 

Rogers, 1987. 

cifically, Weberling (1977) presented a comprehen- 

Colleters in the Rubiaceae are most commonly sive review of the generalized typology of Rubiaceae 

found inside of the stipules, usually at the basal por- inflorescences, and diagrammatic representations can 

tion, and are also frequently found inside the calyx, be found in Delprete (1996d: fig. 2B). When differat 

the base of the calyx lobes or alternate to them. ent interpretations on inflorescence morphology were 

For the group of genera under study, colleters have presented by various authors, Weberling's (1977, 

been consistently found inside the stipules at the basal 1992) conclusions were used. 

portion. 

Inflorescence position, in the genera studied, is 

When apical stipules are about to open, their in- most commonly terminal, with the exception of 

ternal colleters begin to secrete a resinous material Chimarrhis (axillary and subterminal). I consider the 

which is believed to protect the young vegetative buds axillary inflorescences of Chimarrhis to be the result 

against herbivory. The presence of"sticky" resinous of extreme reduction of the last terminal nodes 

exudates in vegetative buds has been observed in (Weberling, 1977, 1992; Robbrecht, 1988). 

Rustia, Condaminea, Pogonopus, and Chimarrhis. I Particular attention should be called to the infloconsider 

absence or presence, arrangement, and gen- rescences ofPogonopus, Molopanthera, and Dioicoeral 

morphology of stipular colleters significant taxo- dendron, which have been variously described as ternomic 

characters, especially when used at the generic minal or terminal and lateral. Standley (1918, 1930a, 

and specific levels (Fig. 3). 

1931 b, 1931 c) described the inflorescence of Pogon- 

In Rustia, stipular colleters are present in a trian- opus as "cymes arranged into terminal leafy panicles"; 

gular area on the base in R. occidentalis (Fig. 3A), in Steyermark (1974) as [literally translated from Spana 

single basal row in R. alba (Fig. 3B), or as a narrow ish] "densiflorous subcorymbose-cymose with 

basal area in R.formosa (Fig. 3C). In Picardaea (Fig. plurichothomous ramification"; Dwyer (1980) as "ter- 

3M) stipular colleters are present in a single basal row. minal paniculate and with small lateral cymes," 

In Tresanthera (Fig. 3E) colleters are long-slender and though he described P. speciosus as having "inflorespresent 

on the base. In Condaminea (Fig. 3G) colleters cences terminal and axillary and thyrsoid-cymose"; 

are present in an irregular basal area on each of the Burger and Taylor (1993) [as in Dwyer, 1980] defour 

stipular units. In Pogonopus (Fig. 3K,L) the scribed Pogonopus with "inflorescences terminal, 

colleters are in one to two irregular basal rows and paniculate and subcorymbose-cymose," but P. 

intermixed with sericeous hairs. In Chimarrhis (Fig. speciosus with "inflorescences terminal or axillary." 

30-Q) the position of the colleters is variable, but In my opinion (and according to Weberling, 1992), 

usually present on a basal area in C. cymosa (Fig. 30), Pogonopus, Molopanthera, and Dioicodendron have 

in a single basal row in C. glabriflora (Fig. 3P), or on terminal frondose paniculate inflorescences, with cya 

densely packed narrow basal area in C. turbinata mose branches subtended by foliose bracts, similar to 

(Fig. 3Q). In Dioicodendron (Fig. 3R) the stipular foliage leaves. The basal "leaves" (pherophylls) subcolleters 

are sparsely distributed in the basal area and tending the inflorescence branches are identical in size 

intermixed with golden densely pubescent hairs. In and shape to foliage leaves, and are gradually smaller 

Dolichodelphys (Fig. 1S) the colleters are sparsely toward the distal portion of the inflorescence. In gendistributed 

on the basal portion of the stipule. In eral, the presence of"leaves" (pherophylls) in termi- 

Molopanthera (Fig. 3T) only a few colleters are nal inflorescences of the Rubiaceae has led the above 

present at the base of the stipules and intermixed with cited workers to incorrectly describe the infloresgolden-sericeous 

hairs. [In Pinckneya (Fig. 3I,J) stipu- cences as lateral and cymose instead of terminal and 

lar colleters are sparsely distributed in a basal trian- paniculate. This argument was convincingly made by 

gular area. In Kerianthera (Fig. 3D) the colleters are Troll (1950) and again by Weberling (1977, 1992). 

sparsely distributed in an ovate area up to half the Inflorescence position of Chimarrhis is unique 

length of the stipules and (as in Pogonopus) are in- among the genera studied in having what I call "axiltermixed 

with gold-sericeous hairs. In Kajewskiella lary subterminal inflorescences," being borne in pairs 

(Fig. 3N) the colleters are long-slender, in a basal area in the leaf axils on the last terminal nodes. In my field 

one-third the length of the stipule and intermixed with observations of Amazonian species (i.e., C. glabrisericeous 

hairs]. 

flora, C. hookeri, C. barbata, and C. duckeana), the


apical shoot stops growing during anthesis and fruit 

maturation, and the axillary inflorescences are 

abscissed just after seed dispersal (sometimes even 

before capsules dehiscence), immediately resuming 

apical shoot growth (see phenology of C. glabriflora). 

In some species growing in Central America and the 

Caribbean region, where the climate is markedly seasonal, 

the inflorescences sometimes remain attached 

to the branchlets up to one year after seed dispersal 

(C. parviflora, C. cymosa, and C. microcarpa). The 

axillary-subterminal inflorescence position of 

Chimarrhis is a good taxonomic character that easily 

separates it from related genera. 

The inflorescence morphology of the taxa presented 

here is quite variable. The terminal panicle is 

the most common, with architecture variable among 

and within genera. The panicles of Rustia vary from 

many-branched pyramidal and once- to thrice-compound 

(R. costaricensis, R. alba, R. bilsana, R. rubra, 

R. viridiflora, R. venezuelensis, R. formosa, R. simpsonii) 

to few-branched and reduced (R. occidentalis, 

R. dressleri, R. gracilis) to few-branched and longpedunculate 

(R. angustifolia) to racemoid or thyrsoid 

secundiflorous (R. schunkeana, R. thibaudioides). 

ers are perfect in all genera examined, except in Dio- 

icodendron, which has individuals with either male 

or female flowers only. 

In most of the genera studied the flowers are acti- 

nomorphic, except for Condaminea and Molopan- 

thera. The anthers of Condaminea are reflexed toward 

the lower base of the corolla tube and all open upward, 

forming a landing area for visitors/pollinators. The 

flower buds of Molopanthera are zygomorphic, curving 

upward; its anthers and corolla lobes are longer in 

the lower portion of the minute flowers; the two lower 

stamens have filaments curved upward, and their anthers 

often remain connected by the cuneate extensions 

present at both ends (forming a cup-shaped unit). 

The flowers of this group are usually presented in 

an erect position, with the exception of some pendulous-flowered 

species of Rustia (R. thibaudioides, R. 

schunkeana, R. viridiflora, and R. rubra; see Delprete, 

1996a). The floral morphology in Rustia is very diversified, 

in response to the process of coevolution 

with a variety of pollinators. This speciation has probably 

been facilitated by Rustia's poricidal anthers, 

which can be variably exserted or included, adapting 

to the several pollination syndromes found in this 

Tresanthera inflorescences vary from densely thyrs- genus. In R. costaricensis the flowers are minute, 

oid panicles (T. condamineoides var. thyrsiflora) to erect, and exserted spreading anthers, and are pollipauciflorous 

thyrses (T. condamineoides var. conda- nated by small bees (lap-pollination). The fleshy flowmineoides). 

Condaminea have terminal, long-pedun- ers ofR. occidentalis, R. bilsana, and R. dressleri have 

culate, corymbose panicles, once- to twice-compound. anthers only half-exserted and convex toward the cen- 

Pogonopus, Dioicodendron, and Molopanthera have ter, which are instead buzz-pollinated by bees (perterminal 

frondose panicles with corymbose branches sonally observed in R. occidentalis). 

(tertiary thyrsoid in Molopanthera). Picardaea has 

short-pedunculate, corymbose, trichotomous panicles 

Calyces 

terminating with triflorous cymules. Dolichodelphys All the genera studied have gamosepalous calyces 

has laxly branched panicles with many secondary cy- positioned at the apices of the ovaries. The general 

mose branches. 

shape of the calyces varies from small-cupular to ex- 

Inflorescence branches of the genera studied are comtremely 

reduced, but this character is consistent at 

monly opposite to subopposite and decussate, and the most at the specific level. Calyx lobes vary from easrachis 

is usually basally terete and decussately comily 

distinguishable to extremely reduced (i.e., a wavy 

pressed distally, where the lateral branches are inserted. 

margin) or absent. The calyces are reduced to a mi- 

Inflorescence bracts subtending primary and secnutely 

undulate margin (sometimes with barely visondary 

branches are consistently present, and vary ible lobes) in Rustia, Tresanthera, Pogonopus, 

from leaf-like (pherophylls) to extremely reduced and Chimarrhis, and Dioicodendron. In Condaminea, 

deltoid, to narrowly linear. The bracteoles subtend- Picardaea, Molopanthera, Dolichodelphys, and 

ing flowers are not constantly present, are always Parachimarrhis the calyces are small-cupular and 

minute, and vary from deltoid to linear. 

vary from truncate to obviously lobed. 

The persistence of the calyces in mature fruits is 

Flowers 

an important taxonomic character for the taxonomic 

The floral morphology of the genera here treated delimitation of the genera here examined. The calyis 

rather variable. Flowers are protandrous in all the ces are consistently persistent in mature capsules of 

genera except for Chimarrhis, where the receptive Rustia, Tresanthera, Chimarrhis, Dioicodendron, 

stigmas protrude above the still-closed corollas (for Dolichodelphys, and Parachimarrhis, and caducous 

more details see discussion under Chimarrhis). Flow- in those of Condaminea, Pogonopus, and Picardaea.


Calycophylls 

Calycophylls are defined as the expansion of one 

or more calyx lobes into colorful foliose units that serve 

mainly to attract visitors/pollinators, and probably to 

effect seed dispersal in mature capsules (Delprete, 

1966b). The oft-used term "semaphyll" was coined by 

Leppik (1956) and refers to "all colored leaves ofplants, 

like petals, sepals, bracts, ligulate flowers, etc., which 

serve to attract pollinators," and therefore is not specific 

to calyx lobes. In the Rubiaceae the presence of 

calycophylls is usually correlated with capsular fruits, 

variably occurring in many genera (Robbrecht, 1988; 

Delprete, 1996b), and which I treated as a secondary 

taxonomic character at the generic and specific ranks. 

A survey of the presence and general morphology of 

calycophylls in Rubiaceae was recently published by 

Classen-Bockhoff (1996). As discussed elsewhere 

(Delprete, 1996b, 1996c, 1996e, 1997b, 1998b, 1998c), 

some genera that were previously separated from their 

sister taxa based on the presence of calycophylls need 

to be reconsidered and probably returned to their origividuals 

growing in undercanopy and shady places 

have white to cream-white calycophylls, while the 

individuals in sunny exposures have pink to pale-lilac 

calycophylls.) In Chimarrhis the calycophylls are 

cream-white to greenish white (as in Calycophyllum). 

In Pogonopus (and Pinckneya), as the capsules 

approach maturity the calycophylls gradually lose 

their color, drying out, becoming brown, and are 

sometimes eliminated by breaking off of their driedup 

fragile stalks, or persisting on the mature capsules. 

(I speculated that in some cases the dried-up 

calycophylls of Pogonopus remain attached to the 

mature capsules while the seeds are released, probably 

assisting in the dispersal of the minute seeds by 

shaking the capsules as a result of wind turbulence.) 

In Parachimarrhis several flowers per inflorescence 

have a small calycophyll. The calycophylls of 

this genus are white to cream-white and 7-13 mm 

long, with the abaxial side covered with a dense mat 

of glandular hairs. The function of this mat of hairs, 

not seen in any other rubiaceous genera, is unknown. 

nal taxonomic position. For example, Chimarrhis has 

been historically described as lacking calycophylls, but 

Corollas 

the newly described C. gentryana (Delprete, 1996b) dis- The corollas of this group vary considerably in 

plays an abundance of cream-colored calycophylls (for size, shape, and thickness among and within genera 

which it has been often confused with Calycophyllum), (Fig. 2), and are commonly actinomorphic (except 

which are also sporadically found in C. brevipes, C. Condaminea and Molopanthera). The Condamineeae 

barbata, and C. turbinata. Calycophylls are typically (sensu Hooker) have been classically distinguished 

pigmented (white, cream, pink, or red) and have pal- from the Rondeletieae in having valvate aestivation, 

mate venation. In some species of Chimarrhis (C. but careful observation showed that only a few genbarbata 

and C. turbinata) bracts are present as foliose era of this group have truly valvate corollas. All the 

expansions in the terminal portions of the inflores- genera placed in the subtribe Portlandiinae sensu 

cences, not to be confused with calycophylls, because Hooker (Bikkia, Ceuthocarpus, Coutaportla, Couthey 

are extrafloral, not colored, and pinnately veined. tarea, Cubanola, Hintonia, Isidorea, Nernstia, Osa, 

Pogonopus consistently has inflorescences with a Portlandia, Schmidtottia, Siemensia, and Thogsennia) 

multitude of calycophylls. Because this genus has have imbricate lobes with reduplicate corolla tubes. 

corymbose inflorescences, the calycophyllous flow- and have been removed to the newly amended tribe 

ers are positioned on the older external flowers of the Catesbaeeae (Delprete, 1996d). 

inflorescences (the central flowers of the ultimate Valvate aestivation is commonly defined as nonbranchlets). 

In Pogonopus the number of calycophyl- overlapping floral parts with continuous contact along 

lous flowers per inflorescence is highly variable, rang- their margins. In the genera studied, valvate aestivaing 

from 2-3 to 20-30. 

tion is correlated with fleshy corollas. Typical valvate 

The main function of calycophylls is obviously to aestivation (without induplications or reduplications) 

attract potential pollinators. In Pogonopus, whose 

most probable pollinators are hummingbirds, the 

calycophylls (and flowers) are dark pink to bright red 

(whereas in Pinckneya, whose most probable pollinators 

are butterflies, the calycophylls [and flowers] 

are cream-white to pale-pink). In Pogonopus (as in 

Pinckneya) the color intensity of calycophylls varies 

according to the solar exposure of the inflorescences, 

being pink in shaded individuals and deep-pink to 

dark-red in sunny exposures. (In Pinckneya the indiis 

present in some species of Rustia (i.e., R. occidentalis, 

R. dressleri, and R. bilsana) and in Dioicodendron. 

Valvate aestivation with reduplicate corolla 

lobes is present in the remaining species of Rustia, 

and consistently in Tresanthera, Condaminea, 

Picardaea, and Pogonopus (and Pinckneya). Chimarrhis 

and Dolichodelphys have usually been reported 

to have valvate aestivation (and for this reason were 

previously included in the Condamineeae sensu 

Hooker), but have instead corolla lobes narrowly im-


bricate. Molopanthera shares with Chimarrhis and 

Parachimarrhis narrowly imbricate corollas (Fig. 

2N), the latter differing by having flower buds curved 

upward (and unequal stamens). 

Corolla shapes and sizes are also quite variable 

(Fig. 4). In Rustia corollas vary from minute and cam- 

panulate (R. costaricensis, Fig. 4B) to medium-sized 

and campanulate (Fig. 4A) to long and narrowly tu- 

bular (Fig. 4C). In Tresanthera, Condaminea, and 

Picardaea the corollas are medium-sized and cam- 

panulate (Fig. 4D-F), while in Pogonopus (Fig. 4G) 

(and Pinckneya, Fig. 4H) they are medium-sized and 

long-tubular. Chimarrhis (Fig. 4J,K), Dioicodendron 

(Fig. 4L,M), and Molopanthera (Fig. 4N) have mi- 

nutely campanulate flowers (creamy-white and fra- 

grant). Dolichodelphys is unique in this group in hav- 

ing corollas that vary greatly in size (from 3 cm long 

down to a few millimeters long) and shape (from tu- 

bular to urceolate) depending on their position in the 

inflorescences (see Dolichodelphys below). 

Fleshiness of the corolla is variable among species 

of Rustia, from fleshy (R. occidentalis, R. simpsonii, 

and R. angustifolia) to semifleshy (R. dressleri, R. 

alba, R. bilsana, R. rubra, R. viridiflora, R. schunke- 

ana, R. thibaudioides, R. formosa) or thin-delicate (R. 

gracilis). Corolla thickness varies also among species 

of Pogonopus, being either semifleshy (P. speciosus) 

or membranaceous (P. exsertus and P. tubulosus). In 

Pinckneya the corollas are semifleshy, whereas in 

Tresanthera, Condaminea, and Picardaea the corol- 

las are thick-fleshy (and all with similar pigmenta- 

tion). The minute corollas of Chimarrhis, Dioico- 

dendron, and Molopanthera are relatively thick with 

respect to their dimensions (a few millimeters). 

Dolichodelphys has thinly membranous corollas. 

Corolla color is variable among the species of 

Rustia, being white (R. costaricensis, R. dressleri 

[with yellow lobes], R. alba, R. simpsonii), pinkish 

white (R. thibaudioides, R.formosa, R. gracilis), red 

(R. rubra [with green lobes]), green (R. viridiflora), 

greenish yellow (R. schunkeana), and purple to flesh- 

red outside and white-perlaceous inside (R. occi- 

dentalis, R. angustifolia, and R. bilsana). The fleshy 

corollas of Tresanthera, Condaminea, and Picardaea 

are red outside and white-perlaceous inside. The tu- 

bular flowers of Pogonopus are deep-pink to red [and 

those of Pinckneya a creamy-white with red streaks]. 

The minute flowers of Chimarrhis, Dioicodendron, 

and Molopanthera are cream-white to pale-green. The 

membranous corollas of Dolichodelphys are variable 

in size, shape, and color, ranging from whitish cream 

to yellowish green to pale yellow. 

The genera reported to have a sweet fragrance are 

the minutely flowered trees of Chimarrhis, Dioico- 

dendron, Molopanthera, and Parachimarrhis, which 

have flowers that are cream-white and bee-pollinated. 

At least two species of Rustia have sweet-fragrant 

flowers: R. alba and R.formosa. Pogonopus (and Pinckneya) 

have a sweetish scent that is barely perceptible 

to humans. 

Corollas of the genera studied have an internal ring 

of hairs, usually located toward the base of the corolla 

tube (near the orifice in minute flowers), close 

to filament attachment (the only exception being 

Pinckneya, where the internal ring of hair is absent). 

The function of the internal ring of hairs (usually basal 

in position) is believed to be protection against possible 

nectar-robbers that might enter through the orifice 

of the corollas. This strategy obviously does not 

protect the flowers against wasps, bumblebees, and 

other insects that rob the flowers by lateral incisions, 

below the ring of hairs. 

Stamens 

Stamens in the group studied are most commonly 

exserted, and the anthers open by longitudinal slits, 

except for the poricidal anthers of Rustia and Tresanthera. 

Anther presentation is exceptional in Rustia, 

which also includes a few pendulous-flowered species 

with narrowly tubular corollas: R. thibaudioides 

(completely included), R. viridiflora (included but 

with tips excluded), and R. schunkeana (partially exserted). 

Anther shapes are commonly broadly elliptic 

to narrowly elliptic in all genera, but are long and 

narrow in the pendulous-flowered species of Rustia. 

Rustia and Tresanthera are unique among the genera 

presented in having poricidal anthers, but they 

differ from each other in being apical and biporate in 

the former, and lateral and with a common pore in the 

latter. The modality of anther opening also differs in 

the two genera. The anthers of Rustia open by reflexing 

the apical deltoid flap placed in a dorsal (abaxial) 

position, and leaving the two cylindrical thecae open 

at their apical ends. In Tresanthera the two thecae are 

apically fused and curved toward the center; they open 

by reducing their curvature, causing the lateral common 

pore below the apex to expand in a ventral 

(adaxial) position. Rustia occidentalis (see details 

under this species) has been personally observed to 

be buzz-pollinated by bees, which is the first official 

report of such specialized pollination within the Rubiaceae 

(most common in the Solanaceae and Melastomataceae). 

Other species, R. dressleri and R. bilsana, 

with fleshy campanulate corollas and half-exserted anthers, 

are also probably buzz-pollinated by bees. The 

tubular pendulous flowers of Rustia thibaudioides have 

included anthers and are pollinated by hummingbirds.


A 

C 

tern 

FIG. 4. Flower variation 

in the genera referred 

to the Condamineeae 

(sensu Hooker, 1873). A-C. Rustia. A. R. occidentalis 

(from fresh material and Delprete 6329, TEX). B. R. costaricensis (Gomez-Laurito 11698, F). C. R. thibaudioides (from 

fresh material and Delprete & Fagua 6378, TEX). D. Tresanthera condamineoides var. condamineoides (Sandwith 1757, 

K). E. Picardaea cubensis (Wright 2661, GH). F. Condaminea corymbosa (from fresh material and Delprete & Verduga 

6412, TEX). G. Pogonopus speciosus var. sandwithianus (from fresh material and Delprete & Apreza 6359, TEX). H. 

Pinckneya bracteata (from fresh material and Delprete & Sharif 6474, TEX). I. Kerianthera preclara (Santos & Coelho 

759, K, NY). J, K. Chimarrhis glabriflora (from fresh material and Delprete & Verduga 6424, TEX). J. Flower bud with 

receptive stigma protruding above still closed corolla. K. Later stage (after J), with anthers exserted above open corolla and 

without style (abscissed). L, M. Dioicodendron dioicum. L. Female flower (Barklay et al. 3424, US). M. Male flower 

(Cuatrecasas 8665, US). N. Molopanthera paniculata var. paniculata (Williams & Assis 6861, GH), curved floral bud 

(zygomorphic).


Peculiar anther extensions were observed in Rustia 

and Molopanthera. In R. simpsonii the base of the 

anthers have papillose spheroid extensions. 

In M. paniculata the flower buds are bent upward, 

and when the flowers open the two lower filaments 

often remain bent upward, with two pairs of adjacent 

anthers connected to each other by their pointed ex- 

tensions at both ends (forming two cup-shaped units), 

with the fifth filament remaining free (see Fig. 85C). 

The peculiar floral morphology of Molopanthera 

seems to be a mechanism of pollen projection, which 

has been reported by Schumann (1891: 10) but not 

personally observed. Apparently the pollen is shed, 

when the flower is still closed, into a spherical mass, 

and the single filament projects this mass onto the 

pollinators (most probably bees). [A detailed descrip- 

tion of such pollen projection mechanism in Poso- 

queria has been explained by Halle (1967: fig. 40)]. 

Styles 

Styles in these genera are terete, glabrous, exserted 

beyond the corollas, and terminated by two style 

branches (stigmas). Most species of Chimarrhis are 

exceptional in having styles minutely costate (C. 

parviflora, C. latifolia, C. cubensis, C. jamaicensis, 

C. cymosa, C. microcarpa, C. speciosa, C. turbinata, 

and C. duckeana) or strigose to puberulent (C. 

turbinata, C. barbata, C. duckeana, and C. gentryana). 

When the flowers are presented in a horizontal 

or oblique position, the styles are curved upward in 

Rustia, Condaminea, Chimarrhis, and Pogonopus 

(and Pinckneya). The stigma branches are generally 

ovate-oblong, with stigmatic surfaces minutely (40x) 

papillose. In Chimarrhis, the stigmas are exserted (and 

receptive) beyond the still-closed corollas (protogynous). 

Dioicodendron is exceptional in being dioecious; 

in its female flowers the styles are exserted 

beyond the corollas, and the stigmas are arcuate with 

their tips still connected at maturity. None of the genera 

studied are heterostylous [Chimarrhis was erroneously 

reported as such by Urban (1899)]. 

Ovaries 

In the genera studied the ovaries are inferior and 

bilocular, with axile placenta on which numerous 

ovules are horizontally (vertically in Chimarrhis and 

Molopanthera) inserted. The ovaries are topped with 

a nectariferous disk, composed mainly of parenchyma 

cells. In some genera (Rustia, Tresanthera, Chimar- 

rhis) the shape, persistence, and vestiture of the caly- 

ces in mature capsules proved to be useful taxonomic 

characters at the specific and varietal levels. Shape 

and vestiture of the disks in the generic and specific 

descriptions below are referred to as they appear in 

mature capsules. 

Fruits 

In the genera here studied, fruits are multiseeded, 

bilocular, woody capsules with a central placenta. In 

early stages the exocarp and mesocarp of immature 

fruits are green and fleshy, becoming brown and woody 

at maturity. Mesocarp fleshiness has been considered 

by many botanists (Hooker, 1873; Baillon, 1880; 

Schumann, 1889, 1891; Verdcourt, 1958; Bremekamp, 

1966; Steyermark, 1974; Robbrecht, 1988, 1993) to 

be a key character in establishing tribal and generic 

delimitations. It has been recently demonstrated 

(Bremer, 1992; Bremer & Eriksson, 1992; Bremer & 

Struwe, 1992) that fleshiness of the mesocarp in mature 

fruits has evolved independently several times 

within the Rubiaceae. Nevertheless, its total dismissal 

as a taxonomic character would deprive systematists 

of a useful source of data (Delprete, 1996d). 

Size and shape of capsules are rather consistent 

within the same genus, offering a good practical aid 

for immediate generic identification. The shapes and 

dimensions of capsules are quite variable among genera: 

Rustia globular, obovoid to turbinate, 4-23 x 4- 

9 mm; Tresanthera ellipsoid to obovoid, 15-24 x 8- 

12 mm; Condaminea obovoid-oblong, 14-40 x 6-11 mm; 

Picardaea oblong, 15-19 x 6.5-8 mm; Dolichodel- 

phys narrowly cylindrical, 14-23 x 3.5-5 mm; 

Pogonopus obovoid to ellipsoid, 5-10 x 4-9 mm (and 

Pinckneya bilobed-subglobose, 10-25 x 15-25 mm); 

Chimarrhis obovoid to turbinate, 1.5-12 x 1.5-6 mm; 

Dioicodendron obovoid to turbinate, 3-4.5 x 2-3.5 mm; 

Molopanthera bilobed-subglobose, 2-3 x 4-5 mm; 

and Parachimarrhis, biglobular, 3-4.5 x 1.5-2.5 mm. 

The mode of capsular dehiscence is a valuable 

character in establishing generic boundaries among 

the genera studied. Mature capsules of Chimarrhis, 

Dioicodendron, and Dolichodelphys have septicidal 

dehiscence, whereas Rustia, Tresanthera, Picardaea, 

Condaminea, and Pogonopus (and Pinckneya) have 

loculicidal dehiscence. Picardaea has a unique dehiscence 

pattern, splitting loculicidally but with the two 

halves remaining attached at the apex and base (no 

apical dehiscence), as in Macrocnemum (under which 

it was originally described). 

I recently introduced the term "secondary dehiscence" 

(Delprete, 1995a, 1996d), referring to the splitting 

that occurs in advanced stages of capsule maturity, 

usually after and perpendicular to primary 

dehiscence. Secondary dehiscence may or may not 

occur, depending on the genus, usually occurring only 

at the apices of old capsules; it is a character of some


taxonomic importance at the generic level. Second- examined and mounted on SEM stubs using a light 

ary dehiscence has been observed in old capsules of microscope. The mounted samples were gold-coated 

some species of Rustia, Tresanthera, Pogonopus, with a Ladd sputter-coater, and then studied, mea- 

Chimarrhis, and Dioicodendron. Secondary dehis- sured, and photographed using a Phillips 515 scancence 

has not been found (and is probably absent) in ning electron microscope (SEM) housed at the Cell 

Condaminea, Picardaea, and Dolichodelphys. Research Center of the University of Texas at Austin. 

The dimensions of the seeds are given first by the 

SEED MORPHOLOGY 

ranges of length and second by the ranges in width. 

Voucher specimens are cited in the explanations of 

Seed morphology has been considered for a long the microphotographs. 

time to be of cardinal importance in Rubiaceae systematics 

(Hooker, 1873; Baillon, 1880; Schumann, Rustia (Figs. 5A-F, 6A-H). Seeds of irregular size 

1888a, 1888b, 1889, 1891; Verdcourt, 1958; and shape, flattened, 0.67-2.9 x 0.25-1.33 mm; acute 

Bremekamp, 1966; Steyermark, 1974; Robbrecht, at base; acute, truncate, or uncinate at distal end; testa 

1988, 1993). Recent investigations (Bremer, 1992; shallowly reticulate, cells elongated and longitudinally 

Bremer & Eriksson, 1992; Delprete, 1995a, 1996d) oriented, interspaces smooth to sparsely granular. 

have shown that the importance given to seed mor- Species of which seeds were available were observed 

phology (i.e., endosperm fleshiness, presence of to have the following dimensional ranges: R. costawings, 

etc.) in the classification of this family needs ricensis (Fig. 5A,B) 0.75-1 x 0.25-0.4 mm; R. occireevaluation. 

Exotesta sculpturing is an important dentalis (Fig. 5E) 1.06-1.26 x 0.67-0.83 mm; R. alba 

taxonomic trait at the generic and specific levels. The (Fig. 6F) 2.25-2.9 x 0.65-0.75 mm; R. rubra (Fig. 

presence of wings (testal outgrowths) in seeds was 5F) 1.16-1.5 x 0.33-0.53 mm; R. viridiflora (Fig. 6E) 

considered a very important taxonomic character by 0.87-1.27 x 0.33-0.83 mm; R. schunkeana (Fig. 6A) 

Baillon (1879), Schumann (1888a, 1888b, 1889, 1.17-1.67 x 1-1.33 mm; R. thibaudioides (Fig. 6C,D) 

1891), and Andersson and Persson (1991), but 1.4-1.5 x 0.75-1.25 mm; R. venezuelensis (Fig. 6G,H) 

Bremekamp (1966) and Verdcourt (1958) considered 0.93-1 x 1 mm; R.formosa (Fig. 5C) 1.4-1.6 x 0.48it 

to be of little importance. Bremer and Eriksson 0.5 mm; R. angustifolia (Fig. 6B) 1.33-1.5 x 0.4-0.73 

(1992) concluded that winged seeds are "not predomi- mm; R. gracilis (Fig. 5D) 0.87-1.16 x 0.43-0.67 mm. 

nant in any of the six groups" of their cladistic analysis, 

which "may indicate that winged seeds are more Tresanthera (Fig. 7A-C). Seeds generally ob- 

'evolutionary plastic' than the fleshy fruits." As a triangular, flattened, 1-1.5 x 0.33-0.53 mm; acute at 

result of a phylogenetic analyses using morphologi- base, truncate at distal end; testa reticulate, cells eloncal 

characters, Delprete (1995a, 1996d) concluded that gated and longitudinally oriented, interspaces smooth to 

the presence and general shape of seed wings is de- sparsely granular at margins. The seeds of this monopendent 

on the insertion of the seeds (whether later- typic genus have been measured as follows: T. 

ally or ventrally attached) on the placenta and their condamineoides var. condamineoides (Fig. 7C) 1.33-1.5 

final position at maturity (horizontal or imbricate). x 0.37-0.53 mm; T. condamineoides var. thyrsiflora (Fig. 

Seed wings of different genera should not be assumed 7A,B) 1-1.2 x 0.33-0.5 mm. 

to be homologous, because of their different origins 

and developments. When properly used, seed mor- Condaminea (Fig. 7D-H). Seeds generally obphology 

provides a good set of taxonomic characters long, irregularly 3-4-angular, 0.57-0.77 x 0.13-0.33 

for inferring the phylogeny of the Rubiaceae. mm; acute at base, truncate at distal end; testa shal- 

In most genera, the seeds are laterally attached and lowly reticulate, cells elongated and longitudinally 

horizontally positioned within the capsules (Fig. 5A), oriented, interspaces smooth. The seeds of C. corymexcept 

for Chimarrhis and Molopanthera, where the bosa (Fig. 7D-F) are 0.57-0.77 x 0.13-0.33 mm. The 

seeds are peltate on the placenta and umbilically at- dubiously placed C. elegans have seeds trapezoid (Fig. 

tached. In Chimarrhis the placenta is central and ver- 7H), 1.3-1.87 x 0.9-1.33 mm, testa densely reticutical, 

while in Molopanthera the placenta is globular late, cells variously oriented, and granular interspace. 

and supported by a short stalk in a central position. In All the previous characters place this species within 

most genera, seeds are minute and of irregular shape Rustia, but its floral morphology is similar to that of 

(but in Parachimarrhis are hemi-elliptic in outline, with Condaminea, where it is tentatively maintained. 

lateral linear hilum) and are adapted to wind dispersal. 

Seeds for SEM studies were removed from mature Picardaea (Fig. 8F). Seeds generally oblong to 

capsules of herbarium specimens. The seeds were first trapezoid, irregularly 3-4-angular, 0.5-0.7 x 0.2-0.4


i~ d.J Cri " ... . . ; lOlllslil~i . ......====================================. 

. . .. `~~~l~~''' .::ri :. 

, 

.El;$ifii~ c~~$~llsl3.... .-..l 

. 

... 

1s 74 I | _ 

o. 

"-'~ ,.^ ~... . ... . . :~~-w~ _~ 

:. ':" ' 

5 : __ BF 0.3r / 1F m : i 29 F.. _~~~~~~~~~~~~~~~~~~ 

?::-:?:: ..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

_e~~~~~~:i- ? *_ 

mm;~_ B-F 0, mm.I8_^. s 

FIG. 5. SEM photomicrographs of seeds of Rustia. A, B. R. costaricensis (Croat 22191, MO). A. Capsule in longitudi- 

:.q 

_f~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i# 

I: 

A _ 

_1_.?"iF s~; re??~~ 

_ 

~~8r* :e>. 

a t a - E~, i 

i,. 4l.~ 

* t .,.*? ... 's 

. 

FIG. 5.SM htomcrgrph ofses ofRsi.A .R otriess(ra 29,M).A asl nlniui 

na etin hoigsed aerlyatahd B ed.C .oroa(Iwne a. 8881, MO),U sed.R.gaiis(lzo 

1493 M),sed.E.R.ocidntli (elree 32, EX, ee. . . uba 8021,.;? Ster US) Scal~Y~~ se.,; e br: 3~ A, 

mm; B-F,?" ~ 0. mm.""" ~ I 

::~~l..? 

mm; acute at base, truncate at distal end; testa irregu- 

larly reticulate, cells mostly with irregular shape, 

without particular orientation, interspaces densely 

granular. Even though this genus is usually placed 

close to Condaminea, its seed morphology closely 

resembles that of Pogonopus. 

Pogonopus (Fig. 8A-E). Seeds generally oblong 

to trapezoid, irregularly 3-5-angular, 0.38-0.75 x 

0.23-0.49 mm; acute at base, truncate at distal end; 

testa irregularly reticulate, cells mostly with irregu- 

lar shape and undulate margins, generally longitudi- 

nally oriented, interspaces densely granular. The three

Introduction to the Neotropical Genera Studied 

-i 

35


_16; 

% 

___~~~~~r 

.6. .:A. ,::.'.,::' . 

~.,~. 

~~ ~'~..~~~~~~~~~W. 

.' ...?-.~'.. 

'>B~.. 

' 

~il 

t; .~-~ ~I'. . ' ~ ... '........... 

GiE9~~~~4 ~.~1 ~ ~ %1 A 

~5?FsI~i7:, ?t:,,,~...~. ~ 

-LAW 

~'"~'-'"~:::'~ ~:~:"'~ 

~~~~~S~~~~~~~C ~ ~ ~ ~ ~ 

~ii, 

' ~ 

~?. ~.~~,~.-~.. . .,,.?',fi~ . . 

aIl&j ? 

....~_,~~"?:I;- ???~:~ 

? !~ ...... 

!~ . .~.~ . . 

~~~~~uh ~~~~~~~~~~~4. 

/ 

. .. ~~ .... 

FIG. 7. SEM photomicrographs of seeds of Tresanthera and Condaminea. A, B. Tresanthera condamineoides var. 

thyrsiflora (Bernardi 5829, K). A. Seed. B. Close-up of seed coat. C. Tresanthera condamineoides var. condamineoides 

(Broadway 3076, GH), seed. D-G. Condaminea corymbosa. D. Seed (Delprete et al. 6049, TEX). E. Seed (Schunke 10512, 

MO). F. Close-up of seed coat (Schunke 10512, MO). G. Seed (Fosberg 28672, US). H. Condaminea elegans (Foster & 

Vivar 13333, MO), seed. Scale bars: A, C-E, G, H, 0.3 mm; B, F, 0.01 mm. 

~! 

"~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG. 7 SEM potomirograps of seds o Tresatheraand Codamine. A, . Tresntheracondaineoids var 

rsfor ed .Coeufse d C.i ot Tresanhrodaiedsvr.cnanods 

(Broaday 307, GH),seed. -G. Codamine corymosa. D Seed Delprte eta.64,TX.E.Se Shne152,~ 

MO). F Closeup fsedcat Shne102,M)G.Se(oser287,U)H.Cnarnnaelgn 

F ostr 

Via 333 - Osed E cl br:A , G ,0. m ,F,00 m


...'{ 

. a 

PIP -', . 

MM"'.:.ls 

FIG. 8. SEM photomicrographs of seeds of Pogonopus, Picardaea, and Pinckneya. A. Pogonopus speciosus var. sandwithianus 

(Delprete & Apreza 6359, TEX), cross section of capsule. B. Pogonopus exsertus (Gentry 6720, NY), seed. C. Pogonopus 

speciosus var. speciosus (Pittier 11672, US), seed. D. Pogonopus speciosus var. sandwithianus (Delprete & Apreza 6359, TEX), 

seed. E. Pogonopus tubulosus (Knrukoff 10167, A), seed. F. Picardaea cubensis (Liogier & Liogier 26174, NY), seed. G, H. 

Pinckneya bracteata (Duncan 2703, TEX). G. Seed. H. Close-up of seed coat. Scale bars: A, G, 3 mm; B-F, H, 0.3 un.


species and two varieties have been measured as fol- 

low: P. exsertus (Fig. 8B) 0.38-0.58 x 0.23-0.35 mm; 

P. speciosus var. speciosus (Fig. 8C) 0.6-0.67 x 0.31- 

0.49 mm; P. speciosus var. sandwithianus (Fig. 8D) 

0.44-0.64 x 0.35-0.42; P. tubulosus (Fig. 8E) 0.51- 

0.75 x 0.29-0.47. The seeds of this genus are typical 

in their irregular cells with undulate margins and 

densely granular interspace. 

Chimarrhis (Figs. 9A-H, 10A-F). Seeds with very 

irregular outline, with deeply fringed concentric wing 

and hilum in central position, peltately attached to a 

central placenta and vertical within the capsule, 0.49- 

2.38 x 0.33-1.5 mm; testa deeply reticulate, cells of 

irregular shape in the central portion and tendentiously 

elongated in the outer portion (forming a narrow concentric 

wing) and radially oriented; interspaces 

smooth or with foveolate thickening. 

Chimarrhis is exceptional within the Rondeletieae 

s.l. because of its inflorescence position, flower, fruit, 

and seed morphology. Chimarrhis duckeana (Figs. 

10B, 10F), C. turbinata (Fig. 10D), C. barbata, and 

exceptionally C. gentryana (Fig. 10E) have seed wings 

that tend to elongate in opposite directions, vaguely 

resembling the bipolar wings of the genera of the 

Cinchoneae (sensu Andersson & Persson, 1991). 

The thickenings present within the interspaces of 

the primary reticulation are not consistent within the 

species, being variable even among seeds of the same 

capsule, and cannot be used as a reliable taxonomic 

character. Nevertheless there is a tendency in C. latifolia 

(Fig. 9B), C. barbata, and C. turbinata (Fig. 10D) 

to develop cell thickenings largely parallel to each other 

wings at each corer of the body, hilum in central position, 

0.5-0.95 x 0.4-0.55 mm; testa shallowly reticulate, 

cells of irregular shape and without any particular 

orientation; interspaces finely and densely reticulate. 

Dioicodendron is considered to be closely related to 

Chimarrhis, but the seeds of the former (Figs. 9A-H, 

10A-F) differ from those of the latter in being ovoid in 

general shape, with extremely narrow wings (when 

present), and finely granulate interspaces. 

Molopanthera (Fig. 1 1B). Seeds with very irregu- 

lar outlines, with deeply fringed concentric wing and 

hilum in central position (Fig. 1 B), peltately attached 

to globular placental extensions (one each locule), 1 x 

2 mm; testa shallowly reticulate, cells elongated with 

radial orientation; interspaces with foveolate-reticu- 

late thickenings. 

The seeds of Molopanthera are extremely similar 

to those of Chimarrhis (Fig. 9A-H, 10A-F) in being 

attached by a central hilum and peltate to the placenta, 

with an irregularly fringed concentric wing, and hav- 

ing cells with foveolate thickenings. The two genera, 

however, differ in their flower, fruit, and inflorescence 

morphology. 

Dolichodelphys (Fig. 12A,B). Seeds ovoid, rectangular 

to trapezoid, 3-5-angular, unwinged and with 

hilum in lateral position (Fig. 12A), laterally attached 

to the central placenta, 0.1-0.3 x 0.01-0.15 x 0.01- 

0.1 mm; testa reticulate, cells 4-5 sub-isolateral without 

particular orientation; thickening ovate in cross 

section (Fig. 12B); interspaces smooth. 

and perpendicular to the 

This dubious 

longer margins ("scalariform" 

monotypic genus has been placed in 

thickenings). In C. brevipes (Fig. 10A), C. duckeana 

the Gardenieae (Robbrecht, 1988, 1993) or confused 

(Fig. 10B,F), C. turbinata (Fig. 10D), and C. 

with Rustia 

gentry- 

(Schumann & Krause, 1908c; Standley, 

ana (Fig. 10E), the marginal thickenings have 1930b; 

spo- 

Steyermark, 1974). Based on its seed, flower, 

radic minute echinate 

and fruit 

processes. 

morphology, Dolichodelphys obviously does 

The dimensional ranges of the seeds of the 

not 

spebelong 

to the Gardenieae, and is included in the 

cies directly observed are as follows: C. 

Rondeletieae s.l. 

parviflora 

(Delprete, 1996d). 

(Fig. 9A) 0.5-0.88 x 0.38-0.5 mm; C. latifolia (Fig. 9B) 

0.49-0.78 x 0.33-0.58 mm; C. cubensis (Fig. 9C) 0.58- Parachimarrhis (Fig. 12G). Seeds hemi-elliptic in 

0.87 x 0.48-0.77 mm; C. cymosa (Fig. 9D) 0.56-0.78 x outline, dorsiventrally flattened, laterally winged, and 

0.56-0.67 mm; C. glabriflora (Fig. 9E,F) 0.48-0.68 x lateral hilum (Fig. 12G), laterally attached to a cen- 

0.33-0.45 mm; C. hookeri (Fig. 9G) 0.5-0.63 x 0.35- tral placenta and in two ranks each locule, 2.2-3.5 x 

0.5 mm; C. microcarpa 0.68-0.73 x 0.53-0.7 mm; C. 1.5-2.5 mm; testa shallowly reticulate, cells multilatspeciosa 

(Fig. 9H) 0.7-0.95 x 0.5-0.75 mm; C. turbi- eral without particular orientation (Fig. 10G), 

nata (Fig. O1D) 0.75-1.95 x 0.75-1.25 mm; C. ducke- interspaces smooth. 

ana 1-2.83 x 0.83-1.5 mm (Fig. 10B,F); C. gentryana This monotypic genus was treated as closely related 

(Fig. 10E) 0.95-1.13 x 0.58-0.88 mm; C. brevipes to Chimarrhis by Ducke (1922, hence the name) or com- 

(Fig. O1A) 0.85-1.05 x 0.63-1 mm. 

pared to both Chimarrhis and Elaeagia, but included 

in the Condamineeae because of its valvate aestivation 

Dioicodendron (Fig. 11A). Seed bodies ovoid-sub- (Steyermark, 1965). The seed morphology of Paraorbicular 

and 3-5-angular, sometimes with narrow chimarrhis is similar to that of Simira (Fig. 12E,F) in


____I~ ~ ti~ 

, . 

-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

A lir I 7 5 - 

-v~it ;_;.i-\ _ 

i- >; w f s o w 

t fF h # ft~~~~~~~~"i 

_t ig,. , w t _ 

o. S~h*8t 

~~~~~?- 't- 's e * . >= 

.I 

O _ w 

F 

_ }j.~~~~~~~~~~ 

:' .i:- f ,''k} !t 

FIG. 9. SEM photomicrographs of seeds of Chimarrhis. A. C. parviplora (Jimenez 3418, F), seed. B. C. tatifolia (Allen 

5762, GH), seed. C. C. cubensis (Wright 1622, BR), seed. D. C. cymosa (Wilbur 8137, TEX), seed. E, F. C. glabriplora 

(Schunke 10653, F). E. Seed. F. Close-up of seed coat. G. C. hookeri (Ancuash 504, MO), seed. H. C. speciosa (Steyermark 

95211, NY), seed. Scale bars: A-E, G. H. 0.3 mm; F. 0.02 mm. 

Pt_ } 

F 

39


? . 

, I[~ 

'-.~ ~.-i- ~~ .i 

. 

.~ ,~.., ~'. 

,,.~' t~ l,,1 "1'"."', 

5.~.~ ~:,~.. ...~.:a . ,a. u 

-._ 

-.- ~ ~ ~ ~ ~ ~ ~ .. 

,..i~..:..-,:.,?, 

ii e. --~ 

~.. 

(Vquez Arevalo 8969, TEX). C. duckeana (Nascimento et al INPAWWF .302, NY). Scale bars: A-F, 0.3 . 

FIG. 10. SEM photomicrographs of seeds of Chimarrhis. A. C. brevipes (Maguire d& Politi 27971, US). B. C. duckeana 

(Fr6es 22596, NY). C. C. turbinata (Museum personnel 9 713, P). D. C. turbinata (Silva & Rosario 3 732, NY). E. C. gentryana 

S 

:. ~~:.. ~,..-. .~. 

;.*L.i ___I .~,-~.....' ~?-~I ~,~ 

':'"''"i' 'c 

FIG. 10. SEM p ~ hotmirogr ofseso ph Ciari. A .beie Mgie&Plt 77,U) ..dce


~ag ti '""~: ~t 

..?-.. ... . -. --- -- 

dendron - S colombianum Y ~ (Sanchez a et?al. 6,M, _ _ s ...S ,? . E,. 

dendron colombianum (Sanchez et 1668, MO). E. Seed. F. Close-up of seed coat Scale bars: A-C, E, 0.3 mm; D, F, 0. . 

.. 

~ ' :"~"Y''"::~'"' ......~~~~~~f 

totype), seed. C, D. Keriantherapreclara (Bisby et:a! P18128, NY). C. Seed. D. Close-up of seed coat. E, F. Cuatrecasasio- 

A.Docdendron doomioicum ( Wrancke 1383, UC). , seed. B. Molos-panthsera pncult a t. ucheli b (Warming3 E s. n.n , F, lee-mm 

' 

. ' 'f... 

?' , 

FIG~ %~, ~~~~,, SMpooirgah ? 

iioedo,* .....~her, fseso 

:oooreo 

n uteasidro.. 

A. Dioicoendron diicum ( Wudock 1383 UC), see. B, Moloonthero l:. pnicltlotovar. bttrhellii ( orming s~., F, 

its lateral orbicular wing and extensive lateral hilum, 

and I think the two genera are closely related. 

[Pinckneya (Fig. 8G,H). Seeds with a broad thin lunate-orbicular 

wing (Fig. 8G), dorsiventrally flattened, 

body of seed 2-3 mm long, the wing 1.5-6.4 x 2.5- 

10.3 mm; orbicular at distal end; testa irregularly reticulate, 

cells mostly elongate without particular orientation, 

interspaces secondarily reticulate (Fig. 8H) 

with thickenings parallel to each other and perpendicu- 

41


lar to the two longer margins. This genus is closely 

related to Pogonopus (Fig. 8A-E), but the seed mor- 

phology of each of these two genera is very distinct.] 

[Kerianthera 

(Fig. 11C,D). Seeds with very irregu- 

lar outline (Fig. 11C), with deeply fringed bilateral 

orbicular wing, dorsiventrally flattened, 3.8-5.8 x 

1.3-3.5 mm; testa shallowly irregularly reticulate, 

cells slightly elongate without particular orientation, 

interspaces secondarily irregularly reticulate, with 

subcircular secondary interspaces (Fig. 11D).] 

POLLEN MORPHOLOGY 

Aiello (1979) published a general overview of the 

pollen for most genera of the Condamineeae (sensu 

Hooker), and several genera assigned to the Cinchoneae 

and Rondeletieae. Aiello divided the pollen 

into three main sculpture types: type I, with echinate 

exine; type II, with foveolate exine; and type III, with 

reticulate exine. The genera of the Condamineeae 

(sensu Hooker) with echinate exine (subtr. Portlandiinae) 

have been transferred to the Catesbaeeae (sensu 

Delprete, 1996d). Pollen of types II and III display 

various kinds of intergradation, as the exine of some 

taxa ranges from densely reticulate to foveolate, and 

the distinction between the two types does not hold 

up. For this reason, no particular pollen sculpturing 

categories have been proposed (or maintained) in the 

present study, and the pollen is described below for 

each genus. 

The pollen grains of the genera studied were all 

[Cuatrecasasiodendron (Fig. 11E,F). Seeds tetragonal 

in outline, dorsiventrally flattened, central body 

circular (0.1-0.2 mm diam.), with subentire concentric 

square wing and hilum in lateral position (Fig. 

1 1E), laterally attached to the central placenta, 0.25- 

0.45 x 0.23-0.45 mm; testa shallowly reticulate, cells 

elongated with peltate orientation departing from the tricolporate, and with reticulate-foveolate exine. None 

hilum; interspaces with foveolate thickenings (Fig. of the genera observed has dimorphic pollen, because 

1 1F). This genus has been placed in either Condami- none of them are heterostylous. 

neeae (sensu Hooker) or Rondeletieae. In my opin- As reported in previous studies (Muller, 1979; 

ion this genus is related to Rondeletia (s.l.), and is Blackmore & Barnes, 1986; Pacini, 1986, 1990), the 

therefore not included in the present study.] pollen grains of flowering plants are subjected to 

changes in size and shape due to mechanical properties 

[Kajewskiella (Fig. 12C,D). Seeds ovoid commonly called harmomegathy (Wodehouse, 

(sometimes 

angular), unwinged and with hilum in lateral 

1935). The pollen grains were usually prolate in shape 

at release and became 

position (Fig. 12C), laterally attached to the central 

spheroid upon contact with 

placenta, 0.35-0.6 x 

moisture 

0.25-0.35 mm 

(or a stigmatic surface). The colpi of the 

(K. trichantha); 

testa shallowly reticulate, cells multilateral without pollen grains were invaginated at the release stage (as 

normally released from mature anthers), and turned 

particular orientation; interspaces densely granularinside 

out 

foveolate and interconnected in parallel rows upon contact with moisture and during ger- 

(Fig. 12D). 

mination. 

This dubious genus was included in the 

Also, the exine reticulation at the polar and 

present 

study because it was previously compared to Xantho- equatorial regions was more expanded when the pollen 

was 

phytum (Hedyotideae), Dolicholobium spheroid than when it was prolate. For the 

(Cinchoneae), 

above reasons the 

or Bikkia (Condamineeae s.l.) by Merrill and equatorial measurements in previ- 

Perry 

ous literature are 

(1947), included in the Condamineeae 

variable, depending on the stages of 

(sensu Hooker) 

the 

by Jansen (1979), and tentatively maintained pollen grains (and their laboratory preparation, i.e., 

by 

Robbrecht (1988, 1993). Because of its seed acetolysis) when measured. Because the process of 

morphology 

and presence of raphides, Tange (1995) trans- acetolysis also causes the pollen grains to expand 

ferred Kajewskiella to the Hedyotideae, a decision equatorially and assume a spheroid shape, the equatorial 

measurements were made at full 

with which I expansion, in 

concur.] 

the spheroid stage. 

Pollen-connecting threads were observed in sev- 

[Simira (Fig. 12E,F). Seeds hemi-elliptic in out- eral species of Rustia and Tresanthera. The nature of 

line, dorsiventrally flattened, laterally winged and the threads was not investigated, and their origin is 

with hilum in lateral position (Fig. 12E,F), laterally uncertain; nevertheless, because they were still present 

attached to a central placenta and in two ranks each after acetolysis, it is suspected that they are filiform 

locule; testa finely shallowly reticulate, cells multi- sporopollenin (as reported in Ericaceae and Onalateral 

without particular orientation; interspaces graceae; see Hesse, 1984, 1986; Skvarla et al., 1978; 

smooth (Fig. 12F). Seeds of Simira are included in the Waha, 1984). Pollen-connecting threads have usually 

present study because of their similarity to those of Para- been correlated (Hesse, 1986) with zoophilous pollichimarrhis, 

from which they differ by being larger.] nation. Several species of Rustia tend to be dispersed


!-_ wM 

B 

A ~A 

dp ci.or.o ( .. .Cardenas 

. 

A?.: Seeds. B. Ce of seed cot C, D K 

D, 0 .01~- ? '~ 

.~ 

imE 1 ; 2.. , 

?IG 1,~. ,aesila SMpooirgah ~,:.: Paa.~.rri. 'L..~.~"o 

fseso~lcoep, 

.fia 

A, 

'~.. ~lrcae '.~ B-~:;T-u ?ee c?'i. (aie ~:,: '~!siel &Crea IO, CL. .ed. 

o 

D, 

rihn 

(aes 16 A.i :~' ~'~:'~ i' D. seu ofse ~:',FSiialnioi SeynaR&Epnz 169,F.E 

F.Clseupofsed oa. . M),sed.Sc"'bas:A'"'!'!. 

archmarhs nn;B,0'5 reilba(Rda ia. 28 

m 

(Kajewshz 1667, A). C. Seed. D. Close-up of seed coat. E, F. Simira longifolia (Steyermark & Espinoza 106898, F). E. Seed. 

F. Close-up of seed coat. G. Parachimarrhis breviloba (Rudas et l. 2289, MO), seed. Scale bars: A, C, F, 0.3 mm; B 0.05 mm; 

D, 0.01 mm; E, 10 mm; G, 2 mm. 

43


in clusters due to connecting threads, some species of cies the exine is foveolate (non-columellate, and os 

this genus being pollinated by bees (buzz-pollination without annular thickening). The reticulate exine in 

and lap-pollination) and others by hummingbirds (see the above three species may be related to their spe- 

Pollination and Other Plant-Animal Interactions). cialized pollination syndrome, buzz-pollination by bees 

In all genera pollen is released as single grains, ex- (pers. obs.). Rustia schunkeana (Fig. 13E), R. thibaudicept 

for a single collection of one species of Rustia oides (Fig. 13F), and R. simpsonii (Fig. 14G,H) have 

(R. costaricensis, Gomez-Laurito 11698), which con- minute echinate processes. The presence of minutely 

sistently releases its pollen in tetrads (Fig. 13B). The echinate exine in these species maybe related to their 

pollen of this collection was collected from herbarium pollination syndrome, probably hummingbird pollispecimens 

and I believe that it is teratological, and nation (pers. obs.). 

should be considered so until its viability has been tested. Two photographs of pollen grains of R. rubra (Fig. 

13G,H) were taken to show the different shapes of the 

Materials and Methods 

pollen grains when freshly released from the anthers 

Mature anthers were removed from herbarium (Fig. 13H) and expanded when in contact with moisspecimens, 

and acetolyzed according to a modified ture (Fig. 13G). 

version of Adams and Morton, 1972. Mature anthers In Rustia the pollen is released as single grains, 

were boiled in distilled water (in a water bath) for 10 except for a single collection (Gomez-Laurito 11698) 

minutes, and macerated through a fine filter. The fil- of R. costaricensis for which the pollen grains were 

tered water was then centrifuged and decanted. Pol- constantly released in tetrads (Fig. 13B); in other collen 

and debris were treated with glacial acetic acid for lections of the same species the pollen is released as 

five minutes, centrifuged, and decanted. Pollen was single grains (Fig. 13A). The species directly observed 

then acetolyzed with a freshly prepared mixture of are listed below, with the measurements of the dimen- 

90% acetic anhydride and 10% sulfuric acid (adding 

sional ranges: R. costaricensis [Skutch 4777, US] (Fig. 

acid to anhydride), and boiled in a water bath for 10 13A) 18-19 x 20-21 gpm; R. costaricensis tetrads 

minutes. The acetolyzed pollen was then centrifuged [Gomez-Laurito 11698, F] (Fig, 13B) 34-38 x 41-43 pm 

and decanted in distilled water three times. The pol- (each grain 19-22 x 20-21 lm); R. occidentalis (Fig. 

len suspended in distilled water was then evenly dis- 13C) 19-20 x 20-21 mr; R. dressleri 19-20 x 20-21 jlm; 

persed in petri dishes, and left to evaporate on a warm R. alba (Fig. 13D) 19-21 x 20-21 gim; R. schunkeana 

surface for three days. The petri dishes were left dry- (Fig. 13E) 19-21 x 20-22 gim; R. thibaudioides (Fig. 

ing for this period in order to omit the laborious pro- 13F) 19-21 x 20-22 gm; R. viridiflora (Fig. 14A,B) 

cess of critical-point drying of the specimens. Pollen 19-21 x 20-22 grm; R. gracilis (Fig. 14C,D) 19-21 x 

grains were transferred by pressing the SEM stubs, 20-22 gm; R. angustifolia (Fig. 14F) 19-21 x 20-22 gLm; 

with a small portion of double-stick tape, into the petri 

R. formosa (Fig. 14E) 19-21 x 20-22 tm; R. rubra 

dishes. The mounted samples were then gold-coated (Fig. 13G-H) 19-20 x 20-25 gm; R. simpsonii (Fig. 

in a Ladd sputter-coater, and studied, measured, and 14G,H) 25-26 x 25-26 pim. 

photographed using a Phillips 515 SEM at the Cell 

Research Center of the University of Texas at Austin. Tresanthera (Fig. 15A,B). Pollen grains tricolpor- 

In the descriptions and dimensions reported below ate, 23-27 x 24-28 im; colpi acute at ends, membrane 

(which were obtained by studying at least 50 pollen granular, pores with regular margin with annular 

grains of each species), the first range refers to equa- thickening; exine foveolate. In the variety condaminetorial 

diameters, and the second range to polar lengths. 

oides (Fig. 15B) the exine is regularly foveolate, while 

in the variety thyrsiflora (Fig. 13A) the exine is fo- 

Rustia. Pollen grains tricolporate, 18-26 x 20-26 veolate with irregular pattern and with holes variably 

gm; colpi acute at ends (except in R. simpsonii, Fig. small to minute (Fig. 15A,B). The varieties of this 

14G,H), membrane smooth to minutely granular, monotypic genus have been measured as follows: T. 

pores with irregular margin with (in reticulate exine) condamineoides var. condamineoides (Fig. 15B) 24or 

without (in foveolate exine) annular thickening; 27 x 26-28 im; T. condamineoides var. thyrsiflora 

exine reticulate to foveolate; in some species with (Fig. 15A) 23-25 x 24-26 jum. 

minute echinate processes (in foveolate exine). In R. 

costaricensis (Fig. 13A,B), R. occidentalis (Fig. 13C), Condaminea (Fig. 15C-F). Pollen grains 

and R. dressleri the exine is reticulate (and columel- tricolporate, 24-26 x 26-28 jlm; colpi acute at ends, 

late) both in the apocolpium and mesocolpium (os membrane granular, pores with regular margin with 

with annular thickening), while in the rest of the spe- small annular thickening; exine reticulate (and col-


''~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~,. 

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panded grain. Scale bars: IO pm. ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ bh ' 4in 

panded grain. Scale bars' 10 ~m.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

46 

..~r 

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FI. 4.SE potmirorahsofpole gais f usia , . . irdiloa Luo 67, B) A Euaoralviw 

B. Poa iw ,D .gaii Ocin 65 S.C oa ve.D ls-p .R oms Hrne ta.65,U) 

eqaoilvew .R nusioi Gaio 43,K, oa iw .~ H.R impoi (Gaiu66,P).Eutra 

view. H. Polar view. Scale bars: AX, E-H, 10 1lm; D, 2 pm 

FIG. 14. SEM photomicrographs of pollen grains of Rustia. A, B. R. viridiflora (Lugo 3674, GB). A. Equatorial view. 

B. Polar view. C, D. R. gracilis (O??hioni 655, US). C. Polar view. !. Close-up. E. R. formosa (Heringer et al. 6157, US), 

equatorial view. F. R. angustifolia (Glaziou 14937, K), polar view. G, H. R. simpsonii (Glaziou 6564, P). G. Equatorial 

view. Hi. Polar view. Scale bars: A-C, E-H, IO gm; D, 2 gm.


~ ' . ?.41, 

%;!w" ' 

-. ? 

I 

"i'r" ~~~~~;E~~~T~~i~~C~~~~1?;~~~'~~r?S 

I ~i 

V1.~~~~~~~~~~~~~~~1 

P 

. 

i_-Fw -- 

_. . .- v 

Mi~~~~~~ 

. ' 

~~" - 'j' 

FIG. 15. SEM photomicrographs of' pollen grains of' Tresnthera, Condaminea, and/Picardaea. A. T7resanthera 

NY). G. Polar view. H. Equatorial view. Scale bars: l10 !pm. 

4V~~~~~~~~~~~~~~~~~~~F~L(~~ 

?* IC?- ~ It f*PM 

FIG 1. SM hotmirogaps o plle gain o Tesathra,Codamne, ad icadaa. . resnter 

conamieodesvar tyrsfloa Berard 529,K),poar ie. B T onamieoiesvar codaineide (roawa 

306 G) eutoil iw.CF Cnamna oymoa C , eprt &Wbse 60,TE;E,F Bee 51,N) 

C,E oa iw . qaoilve.F.Eutra iwofuepne ri. ,H ade ubni Sae 80 

NY) G.Polr vew.H. quaoral iew Scle ars IOpm 

, 

_


umellate) in the mesocolpi and foveolate in the 

apocolpi. Many formal names have been proposed for 

C. corymbosa or its segregates, because of its vestiture 

and leaf morphology variation; nevertheless its flower 

and pollen (and seed) morphology is constant through- 

out its range. Condaminea measurements are as fol- 

low: C. corymbosa (Fig. 15C,D) 24-26 x 26-28 gum. 

The dubious C. elegans, tentatively placed in this 

genus, did not have pollen material available to as- 

certain its generic placement. 

Picardaea (Fig. 15G,H). Pollen grains tricolporate, 

23-25 x 23-24 um; colpi rounded at ends, membrane 

granular, pores with irregular margin with marked 

annular thickening (Fig. 15H); exine reticulate (and 

columellate) in the mesocolpi and foveolate in the 

apocolpi. The two previously recognized species are 

here reduced to synonymy under one species (P. 

cubensis). 

Pogonopus (Fig. 16A-D). Pollen grains tricolporate, 

16-18 x 17-18 jm; colpi obtusely rounded at 

ends, membrane granular, pores with regular margin 

with marked annular thickening; exine reticulate (and 

columellate) in the mesocolpi and foveolate in the 

apocolpi. The three species and two varieties have 

practically identical pollen grains, and their measurements 

are as follows: P. exsertus (Fig. 16A,B) 17-18 x 

17-18 jm; P. speciosus var. sandwithianus (Fig. 16C) 

16-18 x 17-18 um; P. tubulosus (Fig. 16D) 16-17 x 

17-18 uim. 

[Pinckneya (Fig. 16E,F). Pollen grains tricolporate, 

24-25 x 26-27 gm; colpi acutely rounded at ends, 

membrane granular, pores with irregular margin with 

thin annular thickening; exine reticulate (and columellate) 

in the mesocolpi and foveolate in the apocolpi. 

This genus is closely related to Pogonopus, which is 

confirmed by their similarity in pollen morphology]. 

[Kerianthera 

(Fig. 16G,H). Pollen grains pentacol- 

porate (Fig. 16G), 28-30 x 25-26 gm; colpi small and 

widely acute at ends, membrane densely granular, 

pores very small with irregular margin without annu- 

lar thickening; exine minutely foveolate throughout]. 

Chimarrhis (Figs. 17A-H, 18A-E). Pollen grains 

tricolporate, 14-17 x 15-18 im; colpi irregularly 

acute at ends, membrane granular, pores with regular 

margin with annular thickening; exine thickly foveo- 

late-reticulate (and columellate) throughout. All the 

species have similar pollen grains, and differences 

among species are minimal. Two photographs of 

grains of C. parviflora were taken to show the different 

shapes when released from anthers (Fig. 17B) and 

as expanded during germination (Fig. 17A). Pollen 

grain measurements were the following: C. parviflora 

(Fig. 17A,B) 14-17 x 17-18 jm; C. latifolia (Fig. 

17C,D) 16-17 x 17-18 Lgm; C. cubensis (Fig. 17E) 

14-16 x 16-17 gum; C. jamaicensis (Fig. 17F) 14-15 x 

15-16 gim; C. cymosa (Fig. 17G-H) 15-16 x 16-17 gm; 

C. speciosa (Fig. 18A,B) 14-15 x 15-16 gm; C microcarpa 

14-15 x 15-16 gm; C. hookeri (Fig. 18C) 16- 

17 x 17-18 gm; C. turbinata (Fig. 18D) 16-17 x 17- 

18 gm; C. barbata 16-17 x 16-17 pm. 

Parachimarrhis (Fig. 18F). Pollen grains tricolporate, 

23-25 x 29-31 jm; colpi acute at ends, membrane 

granular, pores with regular margin and annular 

thickening; exine irregularly reticulate (and 

columellate) throughout (Fig. 18F). The pollen of this 

monotypic genus (P. breviloba) is easily distinguishable 

from other genera because of its irregular reticulation 

with randomly missing links and its slightly 

larger dimensions. 

Dioicodendron (Fig. 18G,H). Pollen grains tricolporate, 

13-15 x 13-15 gm; colpi acute at ends 

(markedly evident), membrane sparsely minutegranular, 

pores with regular margin with thick annular 

thickening; exine thinly reticulate (and columellate) 

throughout, the reticulation decreasing in size 

toward the colpi. This monotypic genus (D. dioicum) 

is related to Chimarrhis, to which it is palynologically 

similar in its small grains, but differs in having thinly 

reticulate exine and well-demarcated colpi. 

Dolichodelphys (Fig. 19A,B). Pollen grains tricolporate, 

21-23 x 22-23 gm; colpi acute at ends (Fig. 

19B), membrane smooth around the pores and 

sparsely minute-granular at ends, pores with regular 

margin with thick annular thickening; exine irregularly 

reticulate (and columellate) in mesocolpi and 

foveolate at apocolpi. The pollen surface of this dubiously 

placed monotypic genus (D. chlorocrater) is 

ornamented with a widely irregular reticulation (Fig. 

19A,B). 

[Kajewskiella (Fig. 19C,D). Pollen grains 

tricolporate, 17-18 x 19-20 gm (K. polyantha); colpi 

round-truncate at ends (Fig. 19D), membrane densely 

and evidently granular, pores with irregular margin 

with irregular and granular thickening; exine regularly 

foveolate (and columellate) lumina decreasing in size 

toward the apocolpi].


4; 

lo. 

rs: 

H. qutoiali Scl Ft. b w 

~~~~~~~~~~~~~~~~~~~~~~,H. 

Equatorial view. Scale bars: 10 gm." 

P~~, 

"? 

,rlra 

~r-.-lrr ' 

*i 

11 

~ ' 

lW 

', 

? 

,N~>. D~.... 

~fi ._....1 

,, . 

I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

10 

FIG. 16. SEM photomicrographs ofpollen grains of Pogonopus, Pinckneya, and Kerianthera. A, B. Pogonopus exsertus 

(Croat 12895, MO). A. Equatorial view. B. Polar view. C. Pogonopus speciosus var. sandwithianus (Deiprete & Apreza 

6359), equatorial view. D. Pogonopus tubulosus ( Vargas 1719, GH), equatorial view. E, F. Pinckneya bracteata (Osborn 

s.n., LL). E. Polar view. F. Equatorial view. G, H. Kerianthera preclara (Santos & Co~iho 759, K). G. Polar view. 

~


..~~~~~~~~~~~~~~~~~~~~: 

.~ ..~ :~.!.~.... 

_n 

-iS v .......................... 9 

- . :A _. B 

- ' _ 

?S 

* i~ .. ~ 

-@. 

.E . ..."...M., B 

~, 

't. 

_ 

' " 

ee 1'S_~'?? 

* ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 

_~~~a E96 4i m G_ 

Tilted view. B. E r o ...... 

. . T 

^ . ' ' _ f f i 1 

ds. ' * .~~~~~~~~~~~~~~~~$u ... 

I g;$-): 

CI.1? cuesis (Wihotomi26o2aphso polae ran vf?iewrhi. FI. C. jaaicnisf(Pocto 

view (Bena 1066, P). H. Equatorial view (E1cnan H338-16, K). Scale bars: IO gm. 

.;]I' :..~.. 

lii 

AS]~~~~~~~~~i 

2 * a _ _ 2 . % * * - _ y JP};...Z>~~~~~~~~~~~~~~~~~~ 

_7 *W RJ_ @ _ # 

_~~~~~~~~~~~~~~~~~~~~~~~~~j. ;w*\w 

l~~lB~~I~ 

; _s_* ,*s> 

FIG. 17. SEMphotomicrogrphs of polle grains ofC imarris A, B.C priloa(are &Blo 87 F.A 

Tite viw .Eutra 

C.cbni Wih 22 

viw(Bn 16, ) H 

iw fuepne ri.C,D .Itfla(len60,F.C 

) oa iw.F .jmiess(rco 91,TX)plrve.GH 

qutria view, (E1unani~: H381,K.Sal as 01 

itdviw .Plrve.E 

.cms.G oa 

r 

19818 TEX Bepola view G, F). CyoA. Goa


nor? 

JiA m... 

_ - x..,,:... :_ 

4,", 

A 

.

ass 

52 


I 

*:J ~Zl:, ,Mt ... A ............ .. +_ B 

..9, 

I _ ' 

FI.1.SM htmcorah a Ka n eskiela foln A, riso olcoepy 

oihdlhsclrcae 

896 qao S.A ial~ temsolium viwo 

theclpu.C, .Eqaoiliwo I.Kewila 

polyanth (Lvrk&Rddl 1346,: A. polrviwD,eutria cl iw as - C, 0Im 

FIG. 19. SEM photomicrographs ofvpollen grains ofDolichodelphys and Kajewskiella. A, B.Dolichodelphys chiorocrater 

(Cuatrecasas 8926, US). A. Equatorial view of the mesocolpium. B. Equatorial view of the colpium. C, D. Kajewskiella 

polyantha (Lavarak & Ridsdale 31346, A). C, polar view. D, equatorial view. Scale bars: A-F = 10 gtm. 

WOOD ANATOMY 

There is little information regarding the wood 

anatomy of the Rubiaceae. Solereder (1908), in his 

Systematic Anatomy ofthe Dicotyledons, reported that 

the members of the Condamineeae s.l. (among many 

other tribes) have sclerenchyma [as "prosenchyma"] 

with bordered pitting. Metcalfe and Chalk (1950), in 

their extensive Anatomy of the Dicotyledons, merely 

listed the genera under general categories and described 

the wood of the Condamineeae (sensu Hooker) 

without very distinctly bordered pits in Pinckneya. 

I found scanty information about the development 

of the cork cambium, which Solereder (1908) observed 

in Pinckneya to originate from a subepidermal 

layer of cells. In an addenda Solereder (1908) reported 

that Pitard [?] observed that in Chimarrhis (among 

many other genera) "the pericycle contains isolated 

bundles of bast-fibers." 

Accurate work on the wood anatomy of the family 

was presented by Koek-Noorman and colleagues 

as follows: with vessels relatively small (


Koek-Normann (1970, 1972), Koek-Noorman and transferred to the Condamineeae by Andersson and 

Hogeweg (1974), and Koek-Noorman and Puff(1983) Persson (1991). Coutarea has recently been transreported 

that the tribes Gardenieae, Ixoreae, Vangueri- ferred to the Catesbaeeae (Delprete, 1995a, 1996d), and 

eae, Isertieae, Naucleae, Mussaendeae, Hedyotideae, Exostema was treated as a closely related taxon (Delprete, 

Anthospermeae, and Rubieae are all homogeneous in 1995a, 1996d; McDowell, 1996, and pers. comm.). 

having only fiber-tracheids, while Psychotrieae, Pae- Koek-Noorman and Hogeweg (1974) performed 

derieae, Morindeae, Guettardeae, Hillieae, and Hame- a cluster analysis using 125 wood anatomical characlieae 

are homogeneous in having only libriform fibers. ters, but a complete data matrix of such characters was 

Koek-Noorman (1969b), in her review of the Ru- not published. Three dendrograms for the Cinchoneae, 

biaceae of Suriname, discussed the wood anatomy and Rondeletieae, and Condamineeae were produced, and 

taxonomy of Chimarrhis as follows: "Schumann was they all displayed a basal "partition in two groups, one 

not sure whether C. turbinata was rightly placed in of which shows libriform fibers, the other one fiber 

this genus. Ducke (1922) referred it to a new genus, tracheids." In the Condamineeae they reported that 

viz. Pseudochimarrhis. According to Bremekamp "the parenchyma pattern seems to be less connected 

(1934) and Steyermark (1965) Chimarrhis and with the fiber type, the arrangement of the vessels, and 

Pseudochimarrhis are not separable. Record and Hess the presence of crystal sand." Bikkia, Morierina, and 

(1943) report that the wood.of C. turbinata greatly Portlandia [which were transferred to the Catesbaeeae 

resembles C. longistipulata [= C. microcarpa], the by Delprete (1996d)] formed one cluster. Bikkia camplace 

of which is not in doubt. They differ only in panulata is reported as "dissimilar in the frequency 

minor features." In the same article Koek-Noorman of radial vessel multiples, relatively large intervascu- 

(1969b) described the wood of Chimarrhis as having lar pits, uniseriate rays, relatively large ray-vessel 

septate fibers, narrowly vasicentric parenchyma, with pitting, presence of crystal sand, and vague growth 

6-12 vessels per mm2 and up to 150 pm wide, upright rings." But this set of characters could be influenced 

ray cells up to 50 um, without sheath cells nor broad by environmental factors. 

rays. As a conclusion she states that the wood of The monotypic Pinckneya was pointed out by 

Chimarrhis resembles that of the Cinchoneae (of Koek-Noorman and Hogeweg (1974) as exceptional 

which she studied Capirona, Ferdinandusa, and Cin- to the Condamineeae (sensu Hooker) in having semichona) 

more than that of the Rondeletieae (of which ring-porous wood, with tangential pore chains and 

she studied only Elaeagia and Warszewiczia). concentric parenchyma bands. They did not mention 

Koek-Noorman and Hogeweg (1974) presented a that this is the only non-tropical genus analyzed (ensurvey 

of some representative taxa of the tribes Van- demic to Georgia and South Carolina, North America) 

guerieae, Cinchoneae, Condamineeae, and Rondeletieae and that its wood anatomy is certainly influenced by 

(all sensu Bremekamp, 1966), which is the only study the seasonal climate to which this genus is exposed. 

in which representative genera of the Condamineeae Considering the dry-seasonal forests to which Pogonwere 

analyzed. The genera of the Condamineeae opus is endemic, I would also expect its wood to be 

(sensu Bremekamp, 1966) analyzed by Koek-Noorman ring-porous (not reported by Koek-Noorman). 

and Hogeweg (1974) were Condaminea (C. corym- Pogonopus and Rustia (with Calycophyllum and 

bosa), Rustia (R. formosa), Chimarrhis (C. cymosa, Mussaendopsis) were reported to have fibers clearly 

C. microcarpa [as C. longistipulata], C. turbinata, C. septate and with "scanty" pits on their tangential walls. 

hookeri), Pogonopus (P. speciosus), Pinckneya (P. bracteata 

[as P. pubens]), Bikkia (B. campanulata, B. palauensis), 

Portlandia (P. grandiflora), and Morierina (M. 

montana). They reported that the Cinchoneae, Rondeletieae, 

and Condamineeae were rather heterogeneous 

Finally, Koek-Noorman and Hogeweg (1974) performed 

two cluster analyses of the Vanguerieae, 

Cinchoneae, Rondeletieae, and Condamineeae combined, 

one with equal character weighting and the 

other with preferential character weighting. Both 

in having both libriform fibers and fiber-tracheids. analyses also showed a basal bipartition so that gen- 

This finding is probably due to the poor delimitation era of various tribes were scattered on the dendroof 

the three tribes at that time. Unfortunately, the au- grams, and sometimes species of the same genus were 

thors discussed only the characteristics of the genera even placed on remote branches. According to these 

exceptional to the Condamineeae, and the descriptions two analyses, the authors concluded that none of the 

of wood anatomical characters for most genera stud- characters studied has reliable diagnostic value, and 

ied (Rustia, Condaminea, Pogonopus) are only frag- they were persuaded "not to pursue the attempt to 

mentary. Koek-Noormann and Hogeweg (1974) re- define the taxonomic tribes under discussion [Ronported 

Exostema and Coutarea to be exceptional for deletieae, Condamineeae, and Cinchoneae as defined 

the tribe Cinchoneae. The two genera were subsequently at that time] on the base of wood anatomy."


My opinion is that wood anatomy can certainly 

constitute an important set oftaxonomical characters 

(mainly when tempered with environmental considerations 

and geographical distribution) in attempting 

to reconstruct a sound evolutionary study of the Rubiaceae. 

No anatomical studies of the Cinchoneae- 

Rondeletieae-Condamineeae complex has been performed 

since the works ofKoek-Noorman, and I strongly 

encourage future workers to undertake such studies. 

PRESENCE OF CALCIUM OXALATE CRYSTALS 

Solereder (1908), in his anatomical overview of 

dicotyledons, listed the presence of calcium oxalate 

crystals in many rubiaceous genera. He reported the 

Condamineeae (sensu Hooker) as having "Crystalsand, 

occasionally together with clustered crystals," 

which he observed in Bikkia, Chimarrhis, Condaminea, 

Pinckneya, Pogonopus, Portlandia, and Rustia. Of the 

buds were collected and preserved in 4:3:1 (chloro- 

form:95% ethanol:acetic acid). Young anthers were 

extracted from the preserved floral buds, squashed, 

and stained with aceto-orcein. Only three successful 

counts were obtained: Rustia occidentalis (Delprete 

6329, TEX) n = 11, Condaminea corymbosa (Delprete 

6049, TEX) n = 11, and Pogonopus speciosus var. sand- 

withianus (Delprete & Apreza 6359, TEX) n = 11. In 

the three genera observed, the chromosomes were very 

small and stained darkly. 

The counts of Condaminea, Rustia, and Pogon- 

opus support the transfer of the subtribes Condami- 

neeinae and Pinckneyinae (the genera of the group 

here studied) to the Rondeletieae, since this tribe has 

chromosome numbers on a base of x = 9, 10, 11, and 

of small size (Kiehn, 1995, and pers. comm.). 

genera of the Condamineeae (sensu Hooker), Metcalfe 

POLLINATION AND OTHER 

PLANT-ANIMAL INTERACTIONS 

and Chalk (1950) reported the presence of crystalsand, 

either alone or accompanied by clustered crystals, 

in the same genera previously reported by 

Solereder (1908). None of the genera of the Condamineeae 

(sensu Hooker) has ever been reported to 

have raphides (and they are probably absent). 

Bremekamp (1966) was the first worker to consider 

the presence of calcium oxalate crystals as an 

important taxonomic character in the Rubiaceae, using 

it to define subfamilies, tribes, and genera. 

Verdcourt (1958) examined thousands of specimens 

Birds. Rustia has attained a high degree of floral 

diversity along with a variety of pollination syndromes 

(lap- and buzz-pollination by bees, hummingbird pollination, 

butterfly-pollination/visited). The pendulousflowered 

species of Rustia are probably pollinated by 

hummingbirds (at least this appears to be so for Rustia 

thibaudioides; pers. obs.); they have tubular pinkwhite 

corollas (R. thibaudioides), red with green lobes 

(R. rubra), green (R. viridiflora), and yellow-green (R. 

schunkeana). 

in order to confirm the validity of this character and 

declared it to be very important (but not at the 

subfamilial level), because of the universal presence 

of raphides in some tribes, and their consistent absence 

in some others. Verdcourt (1958) reported the Condamineeae 

(which he included in the Rondeletieae) 

as having crystal-sand, as observed by all previous 

workers. Koek-Noorman and Hogeweg (1974) used 

Bees. Rustia occidentalis is buzz-pollinated by 

bees of the genera Euglossa and Melipona (see details 

under R. occidentalis); this is the first report of 

buzz-pollination in the Rubiaceae. The fleshy-flowered 

R. bilsana, R. dressleri, and Tresanthera, also 

having anthers only half-exserted and convex toward 

the center, are probably also buzz-pollinated by bees, 

but this has not been observed. Rustia thibaudioides 

this character (classified as raphides, rhombic crystals, 

elongated crystals, druses, and crystal sand) in 

their anatomical analysis of the Cinchoneae-Condais 

visited by bees of the genera Melipona, Ceratina, 

and Trigona (pers. obs.), but their effectiveness as pollinators 

has not been tested. The minute white flowmineeae-Rondeletieae 

(all have crystal-sand but lack 

raphides), but did not publish their observations. 

I agree with Halle (1961, 1967) and Robbrecht 

(1988) that presence and kinds of raphides evolved 

ers of R. costaricensis are presumably lap-pollinated 

by small bees, which at least are frequent visitors. In 

Condaminea, at anthesis, the filaments are deflexed 

toward the lower side of the corolla tube and the anseveral 

times in the Rubiaceae. Nevertheless, the presence, 

quality, and quantity of raphides remains an 

important character for phylogenetic purposes, and in 

defining tribal and generic groups in the family. 

thers are set parallel to each other and opening upward 

(Fig. 46A-F), forming a landing area for large 

bees that pollinate (lap-pollination) the flowers. The 

minute, sweet-scented flowers of Chimarrhis are visited 

and pollinated by small bees. Reports of visita- 

KARYOLOGY 

For genera included here a few counts were obtained 

with the following methodology. Young floral 

tion by bees were obtained from herbarium labels of 

C. latifolia and personally observed in C. glabriflora 

and C. hookeri.


Wasps. Rustia thibaudioides has been observed to 

be visited (but not pollinated), and robbed laterally, 

by unidentified wasps, followed by ants entering 

through their incisions. Flower buds ofPogonopus speciosus 

var. sandwithianus have also been observed to 

be robbed primarily by wasps and secondarily by ants. 

Butterflies. Flowers of Rustia thibaudioides and 

Pogonopus speciosus var. sandwithianus are visited 

(and probably pollinated) by unidentified butterflies. 

Specimen labels further report that the flowers of 

Chimarrhis latifolia are visited by butterflies (but 

probably not pollinated by them). 

Ants. Some individuals of Rustia thibaudioides are 

guarded by unidentified small ants; the same ants were 

also observed to feed on the floral nectaries after en- 

tering the floral buds through lateral incisions previ- 

ously made by wasps. Several individuals of Con- 

daminea corymbosa were observed to be guarded by 

ants, but I consider them to be only casual visitors. 

Ants were seen robbing closed flower buds ofR. thi- 

baudioides and P. speciosus var. sandwithianus 

through lateral incisions made by wasps. 

Mites. Leaf domatia are usually associated with 

mites. All of the species of Rustia, Picardaea, 

Chimarrhis, and Pogonopus [and Pinckneya] with 

domatia have been found to harbor several unidenti- 

fied taxa of mites, but their possible mutualism is yet 

to be shown. 

Fruitgalls. The ovaries of Chimarrhis and Rustia 

can be infested by insects that cause fruit galls. Spe- 

cifically, fruit galls have been observed in Chimarrhis 

barbata, C. turbinata, C. glabriflora, C. hookeri, and 

Rustia schunkeana. The insects that cause these galls 

have not been identified, but they probably belong to 

the Coleoptera. 

HABITAT AND ECOLOGY 

Most genera studied in the present treatment occur 

in a broad range of habitats and are represented in 

several forest types at several levels, often occupying 

specific ecological niches. Rustia occurs in several 

kinds of habitats and is mostly represented by 

medium-tall shrubs that occur in evergreen-seasonal 

forests, at low elevations (300-600 m), and usually 

in proximity to running water. Several species, however, 

are medium-size trees occurring in tropical cloud 

forests and rain forests, at medium elevations (500- 

2000 m). Rustia occidentalis is exceptional, occurring 

as a small shrub in swamplands and slow-moving 

fresh water at low coastal elevations (near sea level). 

Tresanthera is the sister genus of Rustia, and is represented 

by medium-sized trees endemic to coastal 

Caribbean rain forests. 

Condaminea is most commonly represented by 

shrubs. It is very widespread and easily adaptable to 

diverse environments (in Central and South America), 

occupying dry-seasonal forests, cloud forests, rain 

forests, and being especially common in disturbed 

secondary forests and on roadsides of recently built 

Andean roads (see ecology of C. corymbosa). Its tolerance 

to elevational gradients is also remarkable, 

ranging from coastal forests (near sea level), to 

Andean slopes (up to 2000 m), and down to Amazonian 

rain forests. 

Picardaea is a rare shrub restricted to limestone 

cliffs of seasonal coastal forests of the Caribbean islands 

(Cuba and Hispaniola). 

Pogonopus is mostly represented by tall shrubs 

occurring on slopes in dry-seasonal, deciduous, transitional 

tropical forests at medium elevations (500- 

1400 m), and rarely in cloud forests (P. tubulosus) of 

Central and South America, at medium-high elevations 

(1600-2800 m). It is usually encountered in 

proximity to creeks and running water. 

Chimarrhis is represented by large trees with mature 

individuals present at the canopy levels of Caribbean 

seasonal forests, riverine forests, rain forests 

and cloud forests, to lowland non-inundated Amazon 

forests (terra firme), from low to medium elevations 

(300-2000 m). 

Dioicodendron is a rare tall shrub (to small tree) 

present in the understory of seasonal-dry Andean forests 

and cloud forests at relatively high elevations 

(900-3500 m). 

Molopanthera is a tree occurring in seasonal Atlantic 

forests of southern Brazil, at medium elevations 

(300-700 m). 

Dolichodelphys is another rare shrub (to small 

tree), but generally widespread, occurring in Andean 

cloud forests and rain forests, at medium elevations 

(400-1400 m). 

Parachimarrhis is a rare tree known from a few, 

isolated collections from lowland, seasonally inundated 

Amazonian basin. 

USES 

Members of this group have medicinal, economic, 

and ornamental uses. Pogonopus has been used as a 

secondary source of quinine to cure malaria and intermittent 

fevers. Pogonopus exsertus is used as a local 

antimalarial remedy in Guatemala (Standley 79549, 

label data). The Colombian local names "quina" and

56 Flora Neotropi 

"quina morada" (Delprete & Apreza 6359) for P. speciosus 

are suggestive of the local medicinal use of this 

species. The single species most often used to cure 

malaria is P. tubulosus (as "quina" or "sacha quina"), 

which was extensively used by Peruvian, Bolivian, 

and Argentinean indigenous tribes as a remedy for 

malaria and was later adopted by European colonists 

(see details under P. tubulosus). Weddell (1854) reported 

that the bark of P. tubulosus [named by him 

Howardiafebrifuga (Wedd.) Wedd.] was found "on 

the English market" by Elliot Howard, a contemporary 

quinologist; Howard isolated an alkaloid from its 

bark, which Weddell (1854) named "Howardine" and 

described it to have a bitter taste with a styptic aftertaste. 

The alkaloids that were later isolated from the 

bark of P. tubulosus are pogonopamine, pogonopeine, 

pogonopidine, pogonopine (Willaman & Schubert, 

1966), and tubulosine (Raffauf, 1970). Tubulosine is 

currently being studied for its antitumor properties in 

brine shrimp, crown gall tumors of potato, and human 

tumor cells (Ma, 1991; Ma et al., 1990). The bioactive 

properties of tubulosine have also been studied for 

effectively inhibiting the process of peptide chain elongation 

in eukaryotic polysomes (Carrasco et al., 1976). 

Because of its very attractive calycophylls, 

Pogonopus has been cultivated as an ornamental shrub 

throughout its natural range and elsewhere. Cultivated 

individuals of Pogonopus exsertus are reported from 

botanical gardens in the United States (Florida and 

Hawaii), Honduras, Cuba, Puerto Rico, and Brazil 

(see Cultivated Specimens Examined of this species). 

Pogonopus tubulosus is the species with the largest 

calycophylls (to 15 cm long and 13 cm wide!), and is 

widely cultivated in botanical and home gardens in 

Peru, Bolivia, and Argentina. 

The large trees of Chimarrhis offer a much-appreciated 

pale yellow hardwood used for construction 

purposes. The woods of this genus are reported as 

follows: Chimarrhis parviflora and C. latifolia as 

valuable hardwoods in Costa Rica and Panama, C. 

cubensis as dark yellow hardwood in Cuba, C. 

jamaicensis as good construction wood in Jamaica, 

and C. cymosa as a much sought-after construction 

wood in the Lesser Antilles. Chimarrhis glabriflora 

and C. hookeri offer excellent timbers used by Ecuadorian 

and Peruvian indigenous tribes for construction 

of houses and boats; these two species are also 

reported (Duke & Vasquez, 1994) to be used for establishing 

living fences and as sources of good firewood. 

In Venezuela the wood of C. microcarpa is 

used to make wheel-spokes, beams, and posts. The 

pale yellow hardwood of C. turbinata and C. barbata 

are used by indigenous tribes of French Guiana and 

Brazil for construction of houses. Plank-wood from 

the large buttresses of these two Amazonian fort 

giants is commonly used by indigenous tribes to ma 

canoe paddles, and is locally called "pau de remo" 

Brazil and "bois pagaie" in French Guiana. The 

brous bark of C. glabriflora, C. hookeri, and C. turi 

nata is used by indigenous people of Amazonian E 

uador, Peru, French Guiana, and Brazil to make rope 

Ducke (1922) reported that the wood of Par 

chimarrhis is brownish yellow and of good quality 

SPECIES CONCEPTS 

Throughout this study I have used as a worki 

hypothesis the taxonomic species concept of Dan 

and Heywood (1963): "assemblages of individu; 

with morphological features in common and separal 

from other such assemblages by correlated morph 

logical discontinuities in a number of features." I ha 

tried to reevaluate with a critical approach many ml 

phological characters that have been historically treat 

as fundamental to the macroclassification of the R 

biaceae (e.g., valvate corolla aestivation, fruit flesi 

ness, placentation, vestiture, leaf shapes, stipul 

etc.). At the same time, I tried to find new charactc 

that have not previously been used in generic a 

specific delimitations (e.g., leaf domatia, lea 

branchlet succulence, stipule persistence and morph 

ogy, stipular colleters, inflorescence architectu] 

anther morphology, etc.). Leaf shape and gene] 

vestiture have been treated as useful secondary ch; 

acters for the delimitation of species, but I conclud 

that they are better used at most at the varietal lev 

Throughout my treatment only one infraspeci 

rank has been recognized, the variety. Varietal ra 

was assigned when morphogeographic population 

elements were discerned, especially when the ch; 

acters that marked them tended to intergrade in regic 

of contiguity. This follows the practice of many cc 

temporary practitioners, including Cronquist (pe 

comm.), Fryxell (pers. comm.), Gentry (pers. comrr 

Takhtajan (pers. comm.), Turner (pers. comm.), a 

Webster (pers. comm.), among numerous others. 

SYSTEMATIC TREATMENT 01 

THE GENERA STUDIED 

RUSTIA 

1. Rustia Klotzsch in Hayne, Getreue Darstell. Ge 

14: tabs. 14, 15, 555. 1846; Chamisso 

Schlechtendal, Linnaea 4: 179-181. 1829; 

Candolle, Prodr. 4: 360-361. Sep 1830; Karste 

Linnaea 30: 150-151. 1859; J. D. Hooker 

Bentham & Hooker, Gen. PI. 2: 45. 187 

Bentham, Bot. Voy. Sulph. 104. 1844; Baill

Systematic Treatment of the Genera Studied 57 

Hist. PI. 7: 471. 1880; Hemsley, Biol. Centr.- 

Amer. Bot. 2: 14. 1881; Humboldt & Bonpland, 

PI. Aequin. 1: 138. 1808; Karsten in Engler, Bot. 

Jahrb. Syst. 8: 354-360. 1887; Schumanm in 

Martius, Fl. Bras. 6(6): 260-264. 1889; Schumanm 

in Engler & Prantl, Nat. Pflanzenfam. 4(4): 18. 

1891; Standley, N. Amer. Fl. 32: 7. 1918; Standley, 

in each locule, horizontally inserted; immature fruits 

green to red, carnose. Capsules coriaceous to woody, 

globular, turbinate, or obovate; dehiscing 

loculicidally; disk septicidal dehiscence present in old 

capsules of some species. Seeds small, pointed at the 

ends, angular or with minute marginal wings. 

Publ. Field Columbian Mus., Bot. Ser. 7: 26. 1930; 

Steyermark in Lasser, Fl. Venez. 9: 261. 1974; 

Dwyer, Ann. Missouri Bot. Gard. 67: 482-485. 

1980; D. Simpson, Phytologia 33: 4-8. 1976; 

Delprete, Novon 5: 133-139. 1995; Delprete, 

Brittonia 48: 29-34. 1996; Delprete, Novon 8: 

Rustia is unique in the Rubiaceae in having poricidal 

anthers opening by two distinct apical pores. The 

closely related Tresanthera also has anthers pointed 

at apex, but its dehiscence is by one lateral triangular 

pore (below the apex); it also has larger fruits, and 

flattened and truncate seeds. Both Rustia and 

231-234. 1998; Delprete in Harling & Andersson, Tresanthera are also unique in the Rubiaceae in hav- 

Fl. Ecuador (in press). 1999. Type species. Rustia ing leaves with "pellucid glands." These two genera 

formosa (Chamisso & Schlechtendal ex de are in turn closely related to Condaminea, with which 

Candolle) Klotzsch. 

they share filaments attached at base of corolla tube 

and multiseeded loculicidal 

Henlea H. Karsten, Linnaea 30: 151. 1859. capsules with minute 

Type spe- seeds 

cies. Henlea thibaudioides H. Karsten [= Rustia thi- horizontally inserted. Condaminea differs from 

baudioides (H. Karsten) Delprete] non Henlea Grise- Rustia and Tresanthera in having four foliose stipules 

bach, nom. illeg., Abh. K6nigl. Ges. Wiss. G6ttingen per node (vs. two per node), anthers that open by lon- 

9: 37. 1860 [Apr 1861] (Malpighiaceae). gitudinal slits, deciduous calyx, sessile to subpetiolate 

Stomandra Standley, Publ. Field Mus. Nat. Hist., Bot. leaves, and leaf blades without pellucid glands (as in 

Ser. 23: 247. 1947. Type species. Stomandra costa- the vast majority of Rubiaceae). 

ricensis Standley [= Rustia costaricensis (Standley) The flowers of Rustia consistently have valvate 

D. H. Lorence]. 

aestivation (sometimes valvate-reduplicate), are 

protandrous (the stigma is receptive only after the 

Shrubs to trees, sometimes with small buttresses; anthers are dried up), and vary considerably in size, 

bark grayish-pale brown, smooth. Stipules interpetiolar, 

below petiole attachments, free at base, entire, 

narrowly triangular, readily caducous. Leaves petiolate, 

lanceolate, elliptic to ovate, usually acuminate 

shape, and habit. In R. occidentalis, R. dressleri, and 

R. bilsana the flowers are fleshy and usually erect, 

with anthers convex toward the center and closely 

united, and buzz-pollinated by bees. In R. thibaudiat 

apex, pellucid-punctate; petioles thickened at base, 

rarely pulvinate; domatia absent or a tuft of sparse hairs. 

Inflorescences terminal, solitary, paniculate, racemoid 

secundiflorous or thyrsoid secundiflorous; lateral 

branches (when present) opposite, decussate, usually 

oides, R. rubra, R. viridiflora, and R. schunkeana the 

flowers are non-fleshy, pendulous, brightly colored 

(red-green), with anthers included or partially exserted 

(consistent within the species), and commonly visited 

by hummingbirds, butterflies, and bees. In R. alba, 

monochasial. Flowers protandrous; hypanthium narrowly 

obconical, turbinate to cupulate. Calyx cupular 

or extremely reduced, with barely visible lobes or truncate, 

persistent. Corolla tubular, cupular to campanulate, 

with more or less spreading lobes, thin to fleshy, 

white, red, green, or yellow-green; tube short to long 

tubular, glabrous to puberulent inside; lobes (4-)5 

(-6), deltoid to narrowly triangular; aestivation valvate 

to valvate-reduplicate with contact zone. Stamens 

(4-)5, partially or completely exserted, attached 

near the base of the corolla tube; anthers dorsifixed 

near the base, opening by 2 apical pores. Pollen 

tricolporate, exine reticulate to foveolate (sometimes 

minutely echinate). Style exserted; style branches 

short, thick, ovate to oblong. Ovary 2-celled, 

obconical to turbinate; placentation axile; ovules many 

R. formosa, R. simpsonii, and R. gracilis the flowers 

are non-fleshy (thin), white, either erect or nodding, 

with anthers exserted well beyond the corolla, and 

commonly visited by bees. None of the species has 

been reported to have flowers with any particular fragrance, 

except for R. formosa and R. alba (pers. obs.). 

The seeds of Rustia are minute, horizontally inserted, 

non-winged, and irregularly shaped, varying 

from apically truncate to uncinate, and 3-4-sided. 

Rustia is encountered as small shrubs, tall shrubs, to 

medium-size trees. At least one species (R. alba, pers. 

obs.) has been found with small buttresses. The wood 

is usually soft and of no particular use, and the bark 

is usually smooth to rugose, and pale brown to grayish. 

Rustia has 14 species, ranging from Nicaragua to 

southern coastal Brazil. The center of diversity of


Rustia is in South America. Ecuador has the largest species (R. secundiflora [= R. thibaudioides], R. 

number of species (six), three of which are endemic. angustifolia, and R. gracilis) from South America, and 

Four other species are endemic to southern Brazil, synonymized R. pohliana and R. sellowiana under R. 

mostly in the states of Slo Paulo and Rio de Janeiro. formosa. He correctly included Henlea under Rustia 

but erroneously transferred Henlea rosea [= Laden- 

Taxonomic History 

bergia muzonensis (Goudot) Standl.] to Rustia 

The type species of Rustia was first presented by (Schumann, 1891). 

Chamisso and Schlechtendal (1829) as Exostema Urban (1931) described Rustia haitiensis from 

formosum but without description, therefore as nomen specimens without flowers and with old capsules withnudum 

(for detailed information about the nomencla- out seeds. This species remains dubious because only 

ture of this species see Rustiaformosa). De Candolle sterile material was available, and it almost certainly 

(1830) validated Exostemaformosum Cham. & Schltdl. does not belong to Rustia (see doubtful species). 

ex DC., as part of Exostemma sect. Pseudostemma; Standley (1930a) transferred Henlea splendens to 

after a brief description, he stated, "Species Brasili- Rustia, apparently without noticing the obvious simianae. 

An Genus proprium?" Under sect. Pseudo- larity to R. secundiflora [= R. thibaudioides] and to 

stemma he included E. cuspidatum St.-Hil. [= Bathysa H. thibaudioides [= R. thibaudioides]. He also maincuspidata 

(St.-Hil.) J. D. Hook.], E. australe St.-Hil. tained R. rosea [= Ladenbergia muzonensis (Goudot) 

[= Bathysa autralis (St.-Hil.) J. D. Hook], E. for- Standl.; see excluded species] and described R. ferrumosum 

[= Rustiaformosa], and E. souzanum Martius ginea [=Bathysa sp.] (Standley, 1916), R. iserii [=Aliex 

DC. [Bathysa sp.?]. 

bertia isernii] (Standley, 1940), R. longifolia [=Dolicho- 

Rustia was founded by Johann Friedrich Klotzsch delphys chlorocrater] (Standley, 1930b) and R. 

(1846), who dedicated its name to Dr. Rust ("Dem venezuelensis (Standley & Steyermark, 1953). 

Andeken des verbosteren Prasidenten Dr. Rust Standley (1947) established the genus Stomandra 

gewidmet"). Under the genus Rustia Klotzsch (1846) [= Rustia] which he described as different from Rustia 

placed three species: R. formosa, R. sellowiana (vali- in having smaller flowers and corolla tube shorter than 

dated by direct reference to Exostemmaformosum var. the lobes. Both of these characters are not sufficient to 

leprosum Cham. & Schltdl.), and R. pohliana (which separate the two genera, and Lorence (in Burger & Tayhe 

described). 

lor, 1993) recently transferred S. costaricensis to Rustia. 

Karsten (1859) established the genus Henlea with Donald Simpson (1976) published a short revision 

H. thibaudioides, later adding (Karsten, 1861) H. of the Rustia species occurring in western South 

splendens [= R. thibaudioides]. Henlea has been America, and validated Standley's unpublished R. 

treated as a dubious taxon (Robbrecht, 1988, 1993). rubra. In addition, he correctly transferred R. isernii 

In the present treatment all the species of Henlea have to Alibertia, erroneously treated R. longifolia [= Dolibeen 

treated as synonymous with either Ladenbergia chodelphys chlorocrater] as synonymous with Tresor 

Rustia. Henlea thibaudioides (the type species of anthera condamineoides, and declared that the short 

the genus) is here transferred to Rustia, and Henlea description of Henlea thibaudioides [= R. thibaudiis 

treated as synonymous with Rustia (see discussion oides] "is insufficient to permit any taxonomic deciunder 

R. thibaudioides, and rejected species below). sion on its status" and that "it may be something other 

Baillon (1880: 471-472) included both Tres- than Rustia." 

anthera and Henlea in Rustia, stating that Rustia has Dwyer (1980) described Rustia panamensis [= R. 

"poricidal or shortly rimose [anthers], enclosed or costaricensis (Standl.) Lorence], without noticing its 

exserted" and that it is closely related to Condaminea. close resemblance to Stomandra costaricensis. 

Hemsley (1881) transferred Exostemma (sect. Delprete (1995, 1996, 1998) recently described 

Pseudostemma) occidentale Benth. to Rustia (see dis- five new species of Rustia from Panama and western 

cussion under R. occidentalis) without comment. South America (R. dressleri, R. alba, R. viridiflora, 

Schumann (1889) described three additional Rustia R. schunckeana, and R. bilsana). 

Key to the species of Rustia 

1. Corollas 1 cm long (assuming that R. venezuelensis has flowers >1 cm long).


3. Flower buds clavate, rounded at apex; corollas 14 mm long (South America). 

5. Leaf blades broadly elliptic, to broadly obovate; width >/2 the length. 

6. Corolla lobes /3 of corolla length; calyces cupular and membranous; leaves with 16-21 

secondary veins each side (mountains of N Ecuador) .................................................................... 4. R. alba 

6. Corolla lobes '/3-/2 of corolla length; calyces much reduced; leaves with 12-16 secondary 

veins each side (central and coastal Brazil) ........................................ .......................... 11. R. formosa 

5. Leaves narrowly elliptic to oblanceolate to lanceolate; width

60 

,~~~ 

?? , . 

? c~~~~~~~~~~~~~? 


FIG. 21. Distribution of Rustia costaricensis (closed circles) and R. dressleri (closed triangles). 

below; pellucid punctate; primary and secondary veins puberulent hairs (ca. 0.05 mm long) 0.5-0.6 mm from 

glabrous, prominent below; secondary veins 12-20 the base, at the same point as filament attachments; 

each side; tertiary veins starting subparallel and openly lobes 4-6, ca. /3 of corolla length, 1-1.5 mm x ca. 1.5 

reticulate toward the center; petioles 2.5-4.5(-6) cm mm, ovate to triangular, glabrous, minutely papillose 

long, 2-3 mm thick, terete or adaxially flattened; pul- at tip of margins inside. Stamens (4-)5, partly exvinus 

absent but the petiole thickened, becoming serted because of spreading lobes, subequal, attached 

corky at base; domatia absent. Inflorescences pyra- 0.5-0.6 mm from the base of the tube; filaments 1- 

midal, densely to laxly paniculate with opposite or 1.2 mm long, barbate at base, with a minute tuft of 

subopposite decussate branches, departing almost white-puberulent hairs (hairs 0.1-0.3 mm long); anperpendicularly 

from the rachis; 20-33 x 16-29 cm, thers convex toward center, flattened adaxially, yellateral 

branches 8-11 pairs, basal portion of axis not low, banana-shaped, 2-3 x 1-2 mm, dorsifixed near 

branched 3-7.5 cm long; rachis terete or decussately the base, base rounded, microscopically (40x) papilcompressed, 

rachis and branches glabrous to minute- lose throughout. Pollen exine reticulate. Style expuberulent; 

flowers alternate on distal branches; dis- serted, 5-6 mm long, glabrous; style branches ovate, 

tal bracts subtending primary branches 0.8-2.5 x 0.8- 0.6-0.7 mm long, stigmatic surface smooth. Capsules 

1.2 mm, narrowly triangular to deltoid; bracteoles globular to widely obovate, apex hemispherical, 4- 

subtending flowers 0.5-1.3 x ca. 1 mm, deltoid. Flow- 5 x 4-5 mm, dark brown with small lenticels, glaers 

sessile to pedicellate, pedicels to 2.5 mm long, brous throughout; disk black. Seeds 0.75-1 x 0.25glabrous 

to puberulent; hypanthium turbinate, 1-1.5 0.4 mm wide. 

x ca. 1 mm, glabrous. Calyx reduced to an undulate 

margin, truncate or with barely distinguishable lobes, Distribution (Fig. 21) and ecology. Evergreen- 

0.5 mm x ca. 1 mm; lobes (when evident) shallowly seasonal forests of Pacific slopes of Costa Rica, and 

deltoid, to 0.5 mm long. Corolla cupular with spread- Pacific and Atlantic slopes of Panama, at 300-900 m. 

ing lobes, 3-4 mm long, color white to cream-white; Flowering specimens were collected in February, 

tube narrow-cupular, 1-1.5 mm x ca. 1.5 mm, gla- March, and August. Fruiting specimens were collected 

brous outside, glabrous inside, with a ring of white- in February and November.


Reproductive biology. The collections of this bark grayish. Leafy branchlets glabrous, dark green, 

species, from coastal seasonal forests, suggest that it terete; older branches rugose, grayish; lenticels many, 

might have two flowering periods (cf. Lorence in punctiform to linear, 0.5-4 mm long. Stipules nar- 

Wilbur & Taylor, 1993): one after the dry season and rowly triangular, acuminate, glabrous outside, glaone 

after the rainy season. 

brous with basal colleters inside, 10-15(-30) x 25- 

30 mm, dark green, readily caducous, leaving a 

Specimens examined. COSTA RICA. PUNTARENAS: 

white-grayish linear scar. Leaves 17-28(-40) 

Hilly slopes above Rio Naranjo, near Londres and Villa 

Nueva, 300 m, 84028'W, 09?28'N, 10-12 Feb 1988 (fl), 

Burger et al. 12314 (CR, F). SAN Jost: Tarrazu, slopes of 

Cerro Nara, between Esquipulas and Villa Nueva, 84?03'W, 

09?29'N, 12 Feb 1988 (fl), G6mez-Laurito 11698 (F). 

PANAMA. CHIRIQUI: Burica Peninsula, San Bartolo 

Limite, 19-21 km W of Puerto Armelles, 400 m, 83?00'W, 

08?17'N, 19 Feb 1973 (fr), Busey 464 (MO[3], US), 24 Feb 

1973 (fr), Croat 22191 (MO[2], US). VERAGUAS: Valley of 

Rio Dos Bocas, 15.6 km NW of Santa Fe, 450-550 m, 31 

Aug 1974 (fr), Croat 27709 (MO); Valley of Rio Dos Bocas, 

11 km from Escuela Agricola Alto de Piedra, on rd. to 

Calovebora, 450 m, 30 Aug 1974 (fl), Croat 27551 (MO), 

14 Nov 1974 (fr), Mori & Kallunki 3086 (MO). 

Rustia costaricensis is unique in the genus in having 

minute flowers (3-5 mm long), small usually 

globular capsules (4-5 mm long), and inflorescences 

openly paniculate with delicate, sometimes scandent, 

secondary branches. 

Dwyer (1980) described Rustia panamensis on the 

basis of fruiting material, but without referring to 

Stomandra costaricensis. The two taxa are obviously 

synonymous and were treated accordingly by Lorence 

(cf. Burger & Taylor, 1993). 

Two herbarium specimens of Rustia costaricensis 

(Busey 464, MO) were annotated by Dwyer as "R. 

veraguensis," but this epithet remained unpublished. 

The type of Rustia costaricensis is atypical in having 

inflorescences puberulent throughout, with 

branches slightly geniculate and widely zig-zagging, 

while the other collections have inflorescences openly 

paniculate, glabrous, with secondary branches thin and 

delicate and not geniculate. In combination this set of 

characters suggests that these plants deserve formal 

recognition, but more collections are needed. 

2. Rustia occidentalis (Bentham) Hemsley, Biol. 

Centr.-Amer., Bot. 2: 14. 1881. Exostemma (sect. 

Pseudostemma) occidentale Bentham, Bot. Voy. 

Sulph. 104. 1844. Type. Colombia. Nariiio: Isla 

Gorgona, 1841 (fl), Hinds 354 (lectotype, K, selected 

by Delprete, 1999b; isolectotype, BM). 

Figs. 3A, 4A, 5E, 13C, 22, 23 

Shrub 2-5 m tall, exceptionally a tree to 8 m tall, 

to 30 cm dbh, much-branched, rarely single-stemmed; 

x 4-12 

cm, L/W 2.5:1 to 4:1; narrowly elliptic, lanceolate to 

oblanceolate, cuneate to acute (BA = 18-25?) at base, 

acute-caudate at apex, tapering to a long acuminate 

apex, the acumen often falcate, 1.5-3 cm long; dark 

green above, pale green below, foliaceous; drying 

grayish olive-green, stiff-chartaceous; glabrous above 

and below; pellucid punctate; primary and secondary 

veins glabrous, slightly depressed above, prominent 

below, secondary veins 11-16 each side; tertiary veins 

starting subparallel and reticulate in the center; petioles 

20-40 mm long, 1-2.5 mm thick, adaxially concave 

to flattened; pulvinus evident in young leaves, 

becoming corky in mature leaves; domatia absent. 

Inflorescences reduced-paniculate to racemoid, opposite 

to subopposite decussate lateral branches; 4- 

1(-15) x 2-10(-13) cm, lateral branches to 4 pairs, 

basal portion of axis not branched to 3 cm long; rachis 

decussately compressed to terete, rachis and 

branches glabrous; flowers alternate on distal branches 

or in sparse cymules; distal bracts 1-2 x ca. 1 mm, 

deltoid; bracteoles subtending flowers ca. 1 x 0.7-1 

mm, deltoid, glabrous. Flowers pedicellate, pedicels 

4-15 mm long, glabrous; hypanthium narrowly turbinate, 

3-5 x 2-3 mm, glabrous; flower buds clavate. 

Calyx reduced to an undulate margin, to 3 mm long, 

sometimes ciliolate. Corolla tubular, medially faintly 

constricted with perpendicularly spreading lobes, 1.1- 

1.4 cm long, purple to flesh-red outside, white to pearlwhite 

inside, carnose to coriaceous when fresh; tube 

subcylindrical, 7-9 mm long, 2.5-3.5 mm wide at base 

and 4-5 mm wide at the orifice, glabrous outside, glabrous 

inside, without ring of hairs; lobes 5 (rarely 4), 

/2 of corolla length, 5-6 x 1.7-2 mm, triangular, glabrous 

outside and inside. Stamens 5, exserted because 

of spreading lobes, subequal, attached 4-5 mm from 

the base of the tube; filaments ca. 2 mm long, glabrous, 

distally terete, basally flattened-adnate to tube, 

ca. 0.8 mm at base; anthers convex toward the center, 

yellow, banana-shaped, 5-7 x 1.8-2 mm, dorsifixed 

near the base, base rounded, smooth throughout. Pollen 

exine reticulate. Style exserted, 10-14 mm long, 

glabrous; style branches ovate, 0.8-1.2 x ca. 0.9 mm, 

stigmatic surface smooth. Capsules obovoid to widely 

obovoid (rarely globular), obtuse to rounded at base, 

apex hemispherical, 7-12 x 7-9 mm, dark brown to 

black at maturity, with minute lenticels, glabrous 

throughout; disk septicidal dehiscence present of old 

capsules. Seeds 1.06-1.26 x 0.67-0.83 mm.


? 

l: 6mm .":, . 

=*' ??, ,I ~ ~ ~ 

?"?i?r?~ 

FIG 22 Rutiaoccdenali (rofrsmaeiladDlet63,TE.A.Hbtfinoesnewthaue 

FIG. 22. Rustia occidentalis (from fresh material and Delprete 6329, TEX). A. Habit of inflorescence with mature 

leaves. B. Mature capsule. C. Flower bud. D. Longitudinal section of flower bud. E. Flower in later stage of anthesis (note 

style protruded and receptive). F. Detail of receptive style branches. G. Stamen, dorsal view. H. Stamen, lateral view. 

rr~:?' 

D,-

64 


thesis. C. Flower buzz-pollinated by Euglossa 

7z 

sp. D. Flower buzz-pollinated by Melipona sp. 

I~. 3._.: ?: 

....... 

.. 

.. F 

:"' 

... . .'.. '.. 

.... ........ 

?:'?? :~~ :'. .: 

F: 

FIG. 23. Rustia occidentalis. A. Coastal environment typical of R. occidentalis (Choc6, Colombia). B. Flower in an- 

Distribution (see Fig. 29) and ecology. Mostly in 

swampy coastal areas (Fig. 23A), near the coast but 

not influenced by salt water, in proximity of streams 

in lowland costal forests, and in coastal areas of some 

islands, 0-300 m, in Nicaragua, Guatemala, Costa 

Rica, Panama, Colombia, and N Ecuador. Flowering 

specimens were collected in all months of the year 

except January. Fruiting specimens were collected in 

February, March, April, July, August, and December. 

Pollination biology. I observed Rustia occiden- 

talis in coastal swampy sites of Choc6, Colombia, 

where in some areas this species is in almost pure 

stands. The flowers of this species are fleshy and 

purple-red to flesh-red, and never pendulous. Rustia 

occidentalis is mainly visited by bees (Euglossa sp. 

and Melipona sp.; see Fig. 23C,D), which perform a 

rather characteristic buzz-pollination. During late- 

morning hours, if one listens carefully in a large popu- 

lation of this species, it is possible to hear the scat- 

tered buzzing of bees visiting the flowers. This is the 

first species of the entire Rubiaceae that has been 

observed and reported to be buzz-pollinated. 

Representative specimens examined. GUATEMALA. 

IZABAL: San Juan de Nicaragua, 1841, Friedrichsthal s.n. (F) 

[dubious label data-specimen probably collected in Guatemala]. 

*


NICARAGUA. ZELAYA: Area of Bahia de Bluefields, 

Rio Escondido, near Rio Yanten, 0-30 m, 2 Apr 1949 (fr), 

Molina 2085 (F). 

COSTA RICA. Cocos ISLAND: Valley behind Bahia 

Iglesias, 9-11 Apr 1979 (fr), Foster 4133 (CAS, F, US), Mar 

1970 (fl), G6mez 3307 (F); Wafer Bay, 13 Apr 1965 (fl), 

Jimenez 3173 (F, NY); along brook flowing into Wafer Bay, 

18 Apr 1930 (fl), Svenson 331 (F, UC), 3-13 Sep (fl), Stewart 

315 (CAS, GH, US), 5 Dec 1959 (fl), Klawe 1492 (US). 

LIM6N: Near edge of waterway, 3 km N of Moin, 5 m, 13 

Feb 1965 (fl), Lent 359 (F); Cafio de Moin, 13 Feb 1965 (fl), 

Jimenez 2898 (F). PUNTARENAS: Peninsula de Osa, Parque 

Nacional Corcovado, Estaci6n San Pedrillo, 10-200 m, 23 

Jul 1991 (fr), Delprete et al. 5267 (CR, MO, TEX). 

PANAMA. BOCAS DEL TORO: Old Bank Island, 17 Feb 

1941 (fl-fr), Wedel 2114 (F, MO); Laguna of Chiriqui, Nov- 

Dec 1885 (fl), Hart 109 (US). COL6N: Corr. Crist6bal, near 

mouth of Rio Chagres, 11 Mar 1935 (fr), Allen 860 (BR, G, 

MO, P); Corr. Crist6bal, Chagres, Feb-Mar 1850 (fl-fr), 

Fendler 289 (F, K[2], MO[3], US); Corr. Crist6bal, Ft. 

Sherman, 1.5 km SW of Ft. San Lorenzo, near sea level, 20 

Sep 1979 (fl), Antonio 1935 (MO); Corr. Crist6bal, W of 

Limon Bay, Gatun lake, 30 Mar 1956 (fr), Johnston 1745 

(MO); Miguel de la Borda, 20 Apr 1970 (fr), Croat 9828 

(F, MO, NY); along Cafio Rey, near Cocle del Norte, 18 Aug 

1972 (fl), Dressier 4207 (MO); along Rio Culebra, above 

Santa Isabel, 10 Aug 1911 (fl), Pittier 4166 (NY, US). 

DARIEN: Puerto St. Dorothea, 21 Jul 1962 (fl), Dwyer 2290 

(F[2], MO, US); Punta Guayabo Grande, 0-50 m, 20 Apr 

1980 (fr), Antonio & Hahn 4240 (MO); vic. of Pifias, 2 Mar 

1967 (fr), Duke 10567 (ECON), 10605 (MO). 

COLOMBIA. CHoc6: Upstream of Pizzarro, Rio 

Baud6, 2 Aug 1973 (fr), White & Warner 82 (GH, MO); near 

Baud6, right bank of Rio Baud6, ca. 5 m, 2 Feb 1967 (fr), 

Fuchs & Zanella 21703 (US); Mecana, N of Bahia Solano, 

Rio San Juan, 77?21'W, 06?16'N, 31 Dec 1983 (fr), Juncosa 

1589 (MO); Rio San Juan, 80 m, May 1853 (fl), Triana 1784 

(3262.3) (BR, F); Corr. Pangui, trail toward swamps, 17 Dec 

1992 (fl-fr), Delprete 6319 (CHOCO, COL, F, MO, NY, 

TEX), 6329 (NY [4]); Utria bay, trail toward mtns., 18 Dec 

1992 (fl-fr), Delprete 6330 (CHOCO, COL, F, MO, NY, 

TEX); Mun. Nuqui, Corr. Termales, between quebrada Piedra 

Piedra and Rio Terco, 0-25 m, 5 Sep 1994 (fr) Acevedo- 

Rodriguez et al. 6774 (US), 9 Sep 1994 (fr) Acevedo- 

Rodriguez et al. 6900 (US). NARIFIO: Isla de Gallos, near 

Tumaco, Feb 1943 (fr), Dryander 2583 (US); Gorgona Island, 

NE side, 5-20 m, 17 Mar 1975 (fl-fr), Cabrera 3264 

(MO, NY, QCA); Gorgona Island, 18 Oct 1924 (fl), St. 

George Expedition 624 (US). VALLE: La Cuarantena, Buenaventura 

Bay, 9 Jun 1944 (fl), Killip & Cuatrecasas 38970 

(US); Pacific Coast, Rio Naya, Puerto Merizalde, 1 Mar 1943 

(fl), Cuatrecasas 14317 (US); Malaga Bay, mouth of Quebrada 

Sierpe, 17 Feb 1983 (fl-fr), Gentry et al. 40418A (MO). 

ECUADOR. ESMERALDAS: Estero Batia, 22 Jul 1967 (fr), 

Jdtiva & Epling 771 (NY[3], S[2], US[3], VEN); near 

Borb6n, 1967 (fr), Jdtiva & Epling 2201 (NY[2], S[2], 

US[2]); Limones-Borb6n, 5 km before Borb6n, 10 m, 

79?00'W, 01?07'N, 5 Sep 1980 (fl-fr), Holm-Nielsen et al. 

26039 (AAU); Borb6n, between Punto de Piedra and 

Empalme, 78?58'W, 0104'N, 11 Feb 1993(fl-fr), Jaramillo 

14990 (QCA); San Lorenzo, Rio San Antonio, Finca El Horizonte, 

12 Aug 1967 (fl), Sparre 18300 (S); San Lorenzo, Rio 

San Antonio, El Chorro, 22 Aug 1967 (fl), Sparre 18340(S). 

Bentham (1844) first described this species as 

Exostemma (sect. Pseudostemma) occidentale from 

specimens collected from the Gorgona Island, near the 

Pacific coast of Colombia. In his protologue Bentham 

did not refer to specific collections, but nevertheless 

one herbarium specimen preserved at Kew bears the 

label: "Exostemma occidentale Benth., Voy. Sulph. 

104, a large shrub, Isle Gorgona, Hinds 1841." This 

is probably one of the specimens that Bentham saw 

when he described the species, and I selected this as 

lectotype (Delprete, 1999b). Hemsley (1881) later 

transferred it to Rustia, citing three specimens: "Gua- 

temala (Friedrichstal); Panama (Fendler 289); Co- 

lombia Hb. Kew." The last specimen is probably the 

one seen by Bentham when he described this species. 

Rustia occidentalis is the most widespread species 

of the genus, usually encountered as much-branched 

shrubs 2-4 m tall, often with roots submerged in wa- 

ter. This species is easily recognized by its carnose 

flesh-red to purple flowers, deep-yellow convex ("ba- 

nana-like") anthers, and clavate flower buds. 

Rustia occidentalis is most similar to R. dressleri 

(from Panama), but the latter has white flowers with 

yellow lobes, young leafy branchlets semi-succulent, 

and leaf blades 30-45 cm long with 24-30 secondary 

veins (exceptionally to 30 cm long with 11-16 sec- 

ondary veins in R. occidentalis). 

3. Rustia dressleri Delprete, Novon 5(2): 137, fig. 

3. 1995. Type. Panama. Panama: El Llano-Carti 

Hwy., 8-10 km N of El Llano, 31 Aug 1974 (fl), 

Dressier 4703 (holotype, MO). Fig. 24 

Shrub to tree to 7 m tall. Leafy branchlets 7-9 mm 

wide, semi-succulent, grayish, glabrous, terete; older 

branches rugose, pale brown-grayish; lenticels not 

found. Stipules not seen (readily caducous), leaving 

a linear scar ca. 1 mm wide, of the same color as the 

stem. Leaves 30-54 x 9.5-13 cm, L/W 3:1 to 4:1; 

narrowly elliptic to oblanceolate, cuneate (BA = 20- 

25?) at base, acute or acuminate at apex, the acumen 

ca. 1 cm long; shiny grass-green above, pale grayish 

green below, subcoriaceous; drying olive-green, semi- 

coriaceous; glabrous above and below; evidently pel- 

lucid punctate; primary and secondary veins glabrous, 

prominent below, secondary veins 24-30 each side, 

subparallel; tertiary veins starting subparallel and 

openly reticulate in the center, very evident above,


Sn~ 

.r ," AS ^p .\ . .. . . . ,' 

' 

'," 

A 

FIG. 24. Rustia dressleri (A-D from Dressier 4703, MO, holotype; E, F from Herrera 1663, TEX). A. Habit of inflorescence 

with mature leaf. B. Abaxial view of blade showing pellucid punctation. C. Detail of open corolla from flower 

bud, showing callous base and descending pilose tufts. D. Flower bud (clavate). E. Detail of corolla lobe tip, abaxial view. 

F. Stamen with open anther. (Reproduced with permission from Novon 5, fig. 3. 1995.)


faintly evident below; petioles 28-35 mm long, ca. 3 mm Jul 1994 (fl), Herrera 1663 (CR, GB, MEXU, MO, PMA, 

thick, adaxially concave to flattened, glabrous; pul- STRI, TEX). 

vinus not evident but thickened, becoming corky at 

base; domatia absent or a tuft of few hairs. Inflores- Rustia dressleri is similar to R. occidentalis, from 

cences reduced panicles with opposite decussate which it differs in having white flowers with brownbranches; 

(12-)18-30 cm long, basal branches 6.5- ish yellow lobes (flesh-red in R. occidentalis), fila- 

13 cm long, lateral branches 4-7 pairs, basal ments with a basal tuft of retrorse 

portion 

white-pilose hairs 

of axis not branched 0.9-2 cm long; rachis decussately (puberulent in R. occidentalis), larger inflorescences 

compressed, terete at base, rachis and branches semi- (to 11 cm long in R. occidentalis), young branchlets 

succulent, 

thick and 

glabrous throughout, moderately puberu- semi-succulent, the whole plant glabrous, 

lent at nodes; flowers in opposite cymules on rachis and leaf blades 30-45 cm long (exceptionally to 30 

and lateral branches; distal bracts subtending lateral cm in R. occidentalis) with 24-30 secondary veins 

branches 2.5-3 x 3-4 mm, deltoid, glabrous; along each side (11-16 in R. occidentalis). 

bracteoles subtending flowers 1-1.5 x 1-2 mm, deltoid, 

glabrous. Flowers subsessile to pedicellate, 4. Rustia alba Delprete, Novon 5(2): 133, fig. 1. 1995. 

pedicels 0.5-4 mm long, glabrous; hypanthium 

obconical, 2.5-3.5 x Type. Ecuador. Carchi: Environs of Maldonado, 

2-2.5 mm, glabrous; flower buds 

wet montane forest, 1450-1650 m, 31 May 1978 

clavate, 2.5 mm at base and 3.5-4 mm at bulge. Ca- 

(fl), Madison, Plowmann & Besse 4808 (holotype, 

lyx extremely reduced, with a wavy margin or with 

AAU; isotypes, F, QCA, SEL-n.v., US). 

barely distinguishable lobes, 0.5 mm long, sometimes 

Figs. 3B, 6F, 13D, 25A-E, 26A,B 

ciliolate. Corolla tubular with spreading lobes, 1.2- 

1.4 cm long, white with yellow lobes, semi-fleshy 

when fresh; tube 7-8 mm long, 2.5 mm wide at base Shrub to tree to 12 m tall, to 40 cm dbh, muchand 

3.5-4 mm wide at the orifice, glabrous outside, branched tree, with straight trunk, with small butglabrous 

inside, with a ring of white-pilose descend- tresses; bark grayish. Leafy branchlets glabrous, tering 

to erect hairs, 5-6 mm from the base, at the same ete; older branches rugose, grayish to pale brown; 

point of the filament attachments at base of lobes; lenticels sparse. Stipules narrowly triangular, glalobes 

5, ca. /2 of corolla length, 5-6 mm x ca. 3 mm, brous outside, glabrous and with few colleters inside, 

triangular, semi-fleshy, glabrous outside, white-stri- 28-37 x 7-9 mm, reddish green, readily caducous, 

gose at base, margins microscopically (40x) papillose leaving a linear scar, ca. 0.5 mm wide, of the same 

inside. Stamens 5, partially exserted because of nar- color of the stem. Leaves 25-31 x 13-16 cm, L/W 

rowly spreading lobes, subequal, attached 5-6 mm 1.6:1 to 1.7:1; broadly elliptic to broadly obovate, 

from the base of the tube; filaments ca. 2 mm long, acute to rounded (BA 

distally terete, basally flattened-adnate to tube, barbate 

at base, with a tuft ofretrorse white-pilose hairs (0.1- 

0.2 mm long); anthers convex toward the center, yellow, 

banana-shaped, 4-5 x 1.8-2 mm, dorsifixed near 

the base, bases of thecae rounded, subequal, microscopically 

(40x) papillose-echinate throughout. Pollen 

exine reticulate. Style exserted, 13-16 mm long, 

glabrous; style branches ovate to narrowly ovate, ca. 

1.5 mm long, stigmatic surface smooth. Immature 

capsules green and semi-fleshy when fresh; mature 

capsules not seen. Seeds not seen. 

Distribution (Fig. 21) and ecology. Known only 

from three collections made in the provinces of Panama 

and San Bias (Panama). The specimens were collected 

in flowering stage in July, August, and September. 

Additional specimens examined. PANAMA. PANAMA: 

El Llano-Carti Hwy., 10-12 kmN of El Llano, 12 Sep 1974 

(fl), Dressier 4749 (MO). SAN BLAS: Rio Playon, Chico, 

Campamento Neba Dummat, 09 15'N, 78?15'W, 50 m, 10 

= 30-45?) at base, obtuse and 

short-acuminate at apex; the acumen to 1 cm long; 

blade pale green above, yellowish green below, semicoriaceous; 

drying pale green, semi-leathery; glabrous 

above and below; pellucid punctate; primary and secondary 

veins glabrous, prominent below; secondary 

veins 16-21 each side; tertiary veins starting subparallel 

and openly reticulate in the center, faintly 

evident above and below; petioles 2-3 cm long, 3- 

4.5 mm thick, adaxially narrow-concave to flattened, 

glabrous, basally thickened but not pulvinated; domatia 

absent. Inflorescences laxly paniculate, pyramidal, 

lateral branches decussate; 30-46 cm long, 

basal branches 22-30 cm long, lateral branches 6-8 

pairs, basal portion of axis not branched 7-11 cm long; 

rachis terete, rachis and branches glabrous to minutely 

hirtellous; distal bracts 4-5 x 4-5 mm, deltoid; 

bracteoles subtending flowers 2-3 x 2-3 mm, deltoid, 

glabrous. Flowers pedicellate, pedicels 4-6 mm long, 

glabrous to minutely puberulent; hypanthium narrowly 

obconical, 4-6 x 2-3 mm, glabrous; flower buds 

cylindrical, pointed at apex. Calyx cupular with small

68 

..............p.. U 

E(;I F 

' 

. 

.....1 . 


view of blade showing pellucid punctation. C. Flower bud (acute at apex) showing cupular calyx. D. Stamen. E. Detail of 

anther external surface. F. Receptive style. (Reproduced with permission from Novon 5, fig. 1. 1995.) 

.. 

i


4 ,~~~~~~~~6 

~ '.q:" ...,v~,:.~'~.~;~:. 

:~.i iJiii': ............. 

tct 

t. 

'cP,,;t7~lyii 

a ' 

r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

~? ' ::!~::'~.:".:';!:.d :~. .......~,.,:.'":~:~Z~ 2..:,: g. 

.:.,.. 

..W 

-,ZV 

t?3~~~~~~~~~~~~~~~~~~~~~~~? 

. .... . 

?_: ~e~ ,e '??. i~~~~~~~~~~~~~~~~~~~~~~~~~~.... 

...... i 

i![[l-; 

i~ 

'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG. 26. Rustia alba. A. Mature tree, ca. 15 m tall, within dense vegetation (arrows indicate tree canopy). B. Inflores- 

cence with immature fruits. 

lobes, membranous, glabrous, greenish white, 1.5-3 

mm long; lobes 5, 1-2.5 x 1.5-2 mm, ciliolate. Corolla 

tubular with a basal constriction and reversed 

lobes, 1.5-1.9 cm long, white, semi-fleshy when fresh; 

tube subcylindrical with basal constriction, 5-6 x 2 mm, 

3.5-4 mm wide at the orifice; glabrous, minutely verrucate 

outside; microscopically (40x) granulate inside, 

without a ring of hairs; lobes 5, /3 of corolla length, 

9-11 x ca. 3 mm, narrowly triangular, reversed, glabrous, 

minutely verrucose outside, densely ascending 

short-strigose at base, minutely papillose throughout, 

Pollen exine densely foveolate (non-echinate). Style 

exserted, 16-20 mm long, glabrous; style branches 

ovate, 3-3.5 x 1-1.3 mm, stigmatic surface papillose. 

Capsules obovoid, apex obtusely hemispherical, 8- 

12 x 7-9 mm, without lenticels, glabrous throughout, 

dark-brown; dehiscing loculicidally, disk septicidal 

dehiscence absent even in old capsules. Seeds 2.25- 

2.9 x 0.65-0.75 mm. 

Distribution (see Fig. 29) and ecology. Known 

only from the cloud forest of Carchi province, NW 

microscopically (40x) papillose at margins inside. Ecuador, at 1500-1700 m. An extremely rare and 

Stamens 5, fully exserted, subequal, attached 4-5 mm threatened species, because the area where it occurs 

from the base of the tube; filaments ca. 3 mm is 

long, highly disturbed and primarily dedicated to cow 

flattened, widened at base, with a tuft of ascending pastures. Flowering specimens were collected in Febstrigose 

hairs (0.3-0.5 mm long); anthers cylindrical, ruary, May, and August. 

subsagittate at base, rectilinear to slightly curved (not Specimens examined. ECUADOR. CARCHI: El 

convex), 6.5-7 x ca. 1.5 mm, dorsifixed near the base, Carmen, N of Gualchan, 78?13'W, 00?49'N, 1500 m, 9 Feb 

the thecae subequal at base, papillose throughout. 1992 (fl-fr), Palacios et al. 9663 (MO, QCNE, TEX); Mun.


Maldonado, rd. Maldonado-Chical, 1500-1700 m, 14 Aug 

1994 (fl-fr), Delprete & Verduga 6414 (AAU, MO, NY, 

QCA, QCNE, TEX, UPS), 6419 (NY, TEX), 6420 (NY, 

QCA, TEX). 

Rustia alba is similar to R. formosa but differs in 

having a larger inflorescence (exceptionally to 36 cm 

long in R. formosa); calyx cupular, more expanded, 

membranous, and greenish white; smaller corollas; 

and shorter styles with longer, membranous style 

branches (ca. 1 mm long in R. formosa and ca. 3 mm 

long in R. alba). Rustia alba and R. formosa are trees 

commonly reaching 12 m tall and with coriaceous to 

semi-coriaceous leaves, but R. formosa is encountered 

only in southern Brazil. 

Rustia alba differs from R. bilsana in having larger 

flowers, inflorescences, and capsules, expanded mem- 

branous (vs. much-reduced) calices, widely obovate 

(vs. oblanceolate) leaves, and white (vs. purple) co- 

rollas (see discussion under R. bilsana). 

The type material of Rustia alba was collected 

from an individual with several stump sprouts ca. 6 m 

tall. Near the type locality, among a few young trees 

(see Fig. 26A) I encountered one living stump, with 

sprouts 4-5 m long, that was ca. 40 cm dbh. 

5. Rustia bilsana Delprete, Novon 8: 234, figs. 1, 2. 

1998. Type. Ecuador. Esmeraldas: Quinind6, Bilsa 

Biological Station, Mache Mtns., 35 km W of Qui- 

ninde, 5 km W of Santa Isabel, along Cube River, 

premontane wet forest, 00?21'N, 79044'W, 500 m, 

2 Oct 1996, Clark 2979 (holotype, QCNE; iso- 

types, AAU, CAS, COL, DAV, F, G, GB, GH, K, 

MO, NY[2], P, QCA, QCNE, SEL, TEX, US, 

VEN). Figs. 27A-E, 28A-D 

Tree to 19 m tall, to 47.5 cm diam. Leafy branchlets 

glabrous. Stipules narrowly triangular, glabrous 

outside, glabrous and with a few colleters inside, 23- 

27 x 8-10 mm, readily caducous, leaving a linear scar 

ca. 0.5 mm wide. Leaves 33-55 x 13-21 cm, oblanceolate, 

L/W 2.5, acute-decurrent at base, rounded and 

short-acuminate at apex, thick-chartaceous, glabrous 

above and below; secondary veins 15-18 pairs, faintly 

evident above and below; petioles 2-2.5 cm long; 

domatia absent. Inflorescences laxly paniculate, pyramidal, 

16-26 cm long, basal branches 9-12 cm long, 

lateral branches 4-5 pairs. Flower pedicillate, pedicels 

1-2 mm long, glabrous; hypanthium obconical, 3-5 

x ca. 3 mm, glabrous; flower buds cylindrical, pointed 

at apex. Calyx reduced to a wavy margin with barely 

distinguishable lobes, 0.5-1 mm long. Corolla tubu- 

lar with a basal constriction and reversed lobes, 0.8- 

0.9 cm long, purple, semi-fleshy; tube 3.5-4 mm long, 

glabrous outside and inside; lobes 5, 2/3 of corolla 

length, 5-5.5 mm long, narrowly triangular, reflexed, 

glabrous outside, microscopically (40x) papillose at 

margins inside. Stamens 5, fully exserted, subequal, 

attached ca. 4 mm from the base of the tube; filaments 

2 mm long, flattened, widened at base, with a basal 

tuft of antrorse, retrorse strigose hairs; anthers convex 

toward the center and rounded at base, 4.1-4.4 x 

1.2-1.3 mm, dorsifixed near the base, the thecae equal 

at base. Pollen not seen. Style exserted, 9-11 mm 

long, glabrous; style branches ca. 1.5 mm long, stigmatic 

surface papillose. Capsules widely obovoid, 

apex obtusely hemispherical, 7-8 x 6-6.5 mm, without 

lenticels, glabrous throughout, dark brown; disk 

septicidal dehiscence absent even in old capsules. 

Seeds 0.6-1.2 x 0.3-0.5 mm. 


from two collections from the premontane wet forest 

of Esmeralda province, in the Mache Mountains, Bilsa 

Biological Station, in N Ecuador, at 400-600 m. The 

specimens have been collected in late flowering-early 

fruiting stage in October. 

Specimens examined. ECUADOR. ESMERALDAS: Quininde, 

Bilsa Biological Station, Mache Mtns., 35 km W of 

Quininde, 5 km W of Santa Isabel, premontane wet forest, 

00?21'N, 79?44'W, 400-600 m, 3 Oct 1994, Clark, Bass, 

Pitman 139 (MO, QCA, QCNE, TEX). 

Rustia bilsana is similar to R. alba, from the cloud 

forests of northwestern Ecuador, in having floral buds 

pointed at apex and widely obovoid capsules. The two 

differ in the former having leaves oblanceolate (vs. 

widely obovate), acute-decurrent [BA = 20-25?] at base 

(vs. acute-rounded [30-45?]); inflorescence ca. 16 cm 

long (vs. 30-46 cm long); calyx reduced, with a wavy 

margin (vs. cupular and small lobes); corolla purple 

and 8-9 mm long (vs. white and 15-19 mm long); 

corolla lobes 5-5.5 mm long (vs. 9-11 mm long), 

glabrous inside (vs. densely ascending short-strigose 

at base); anthers half-exserted (vs. fully exserted and 

pendulous); style 9-11 mm long (vs. 16-20 mm long), 

style branches 1.5 mm long (vs. 3-3.5 mm long); capsules 

7-8 mm long (vs. 8-12 mm long); and seeds 0.6- 

1.2 x 0.3-0.5 mm (vs. 2.25-2.9 x 0.65-0.95 mm). 

This species differs from Rustia occidentalis in 

being tall trees growing at medium elevations (vs. 

shrubs growing in swampy or wet areas), having 

smaller flowers, flower buds pointed (vs. clavate), 

bigger and more branched inflorescences, and smaller 

capsules.


FIG. 27. Rustia bilsana (Clark et al. 139, TEX, holotype). A. Habit of inflorescence with mature leaf. B. Flower bud. 

C. Open flower. D. Stamen, dorso-lateral 

view. E. Receptive style. (Reproduced with permission from Novon 8, fig. 1. 1998.) 

.' 

.


? N 

i A: .... 

stage of anthesis (top) and one in later stage (bottom). 

6. Rustia rubra Standley ex D. Simpson, Phytologia 

33: 4. 1976. Type. Ecuador. Pastaza: in clearing 

at La Esperanza, Rio Chingual, near jct. with Rio 

Las Ollas, ca. 1700 m, 7 Aug 1943 (fl), Steere 

8021 (holotype, F; isotypes, US, NY). 

Figs. 5F, 13G-H, 30, 31 

Shrub to tree 15-20 m tall, to 30 cm dbh, much 

branched tree, with straight trunk, sometimes with 

small buttresses; bark grayish. Leafy branchlets gla- 

brous, terete (rarely obtusely compressed); older 

branches rugulose, grayish to pale brown; lenticels 

sparse, elongate to linear. Stipules narrowly triangu-


lar, glabrous outside, glabrous and with few colleters 

inside, 20-35 x 4-6 mm, reddish green, readily caducous, 

leaving a linear scar ca. 0.5 mm wide, of the 

same color of the stem. Leaves 22-37(-46) x 8.5- 

14(-17.5) cm, L/W 2:1 to 3:1, elliptic, oblanceolate 

to obovate, acute to rounded (BA = 23-32 mm long, glabrousstyle branches ovate, 0.5- 

0.7 mm long, stigmatic surface smooth; immature 

fruits red and semi-carose. Capsules obovoid to 

oblong-elliptic, apex subtruncate to obtusely hemispherical, 

(9-)11-20 

25-45?), acute to 

obtuse-acuminate at apex; the acumen (when present) 

deltoid, to 1 cm long; blade dark green above, yellowish 

below, chartaceous to semi-coriaceous; drying 

olive-green or pale brown, stiff-chartaceous to 

semi-leathery; glabrous above, glabrous to minutely 

puberulent below; pellucid punctate; primary and secondary 

veins glabrous to minutely puberulent, prominent 

below; secondary veins 14-20 each side; tertiary 

veins starting subparallel and openly reticulate in the 

center, faintly evident above, very evident below; 

petioles 2-5.5(-7) cm long, 2-3(-4) mm thick, terete 

to adaxially flattened, glabrous; basally thickened but 

not pulvinate; domatia absent. Inflorescences densely 

paniculate, pyramidal, twice to thrice compound, lateral 

branches decussate, often secundiflorous, sometimes 

racemoid; rachis, branches, pedicels, calyx, 

corollas, and juvenile fruits deep red; 20-90 x 12-50 

cm, lateral branches 4-12 pairs, basal portion of axis 

not branched 3-9 cm long; rachis terete to decussately 

compressed, rachis and branches glabrous to minutely 

puberulent; flowers alternate on distal branches; distal 

bracts subtending lateral branches 1-4 x 2-3 mm, 

deltoid; bracteoles subtending flowers 0.5-0.8 x 0.5- 

0.8 mm, deltoid, glabrous. Flowers pedicellate, 

pedicels 3-15 mm long, glabrous to minutely puberulent; 

hypanthium narrowly obconical, 3-4 x 1.5-2 mm, 

glabrous to puberulent; flower buds faintly clavate. 

Calyx reduced to a wavy margin or with minute lobes, 

ca. 0.5 mm long; lobes 5 (when distinguishable), 0.1- 

0.5 x 1.5-2 mm. Corolla tubular, 1.8-2.2 cm long, 

deep red with green to yellow-green lobes outside, 

deep red inside, semi-fleshy when fresh; tube subcylindrical, 

striate, 1.6-1.9 cm long, 2-2.5 mm wide at 

base and 2-3 mm wide at the orifice, glabrous outside; 

ascending long-strigose at base inside, without 

a ring of hairs, the remaining medial and superior 

zones glabrous; lobes 5, ca. 1/10 of corolla length, 2- 

3.5 x 1.5-2 mm, deltoid, glabrous outside, glabrous, 

minutely papillose (40x) at margins inside. Stamens 

5, included or partially exserted, subequal, attached 

5-6 mm from the base of the tube; filaments 10-15 

mm long, sparsely pilose at base; anthers subcylindrical, 

sagittate to caudate at base, rectilinear (never 

convex), (4-)5-8 mm long, 1-2.5 mm wide at apex, 

dorsifixed near the base, base sagittate to caudate, the 

thecae sometimes very unequal (3 and 7 mm long on 

same filament), smooth throughout. Pollen exine 

densely foveolate, minutely echinate. Style exserted, 

x 5-8 mm, black, without (rarely 

with few) lenticels, glabrous to puberulent below the 

disk; disk black, glabrous; disk septicidal dehiscence 

absent even in old capsules. Seeds 1.16-1.5 x 0.33- 

0.53 mm wide. 

Distribution (Fig. 29) and ecology. Primary and 

secondary rain forests and cloud forests, and relative 

remnants, in low to medium elevations, 450-1900 m, 

of Amazonian Ecuador and Peru. Flowering specimens 

were collected in February, March, June, and 

July. Fruiting specimens were collected in January, 

May, and June. 

Specimens examined. ECUADOR. NAPO: Rd. Baeza- 

Tena, S slope of Cordillera de Guacamayos, 1800 m, 9 Feb 

1980 (fl), Harling & Andersson 16423 (GB[3], TEX[3]); zone 

of Sofia, valley of Rio Chingal, 15 Jun 1944 (fr), Steere 8339 

(F, NY); Cant6n Archidona, S slope of Vulcan Sumaco, rd. 

Hollin-Loreto, km 31, 00?43'S, 77?36'W, 1200 m, 8-17 Jan 

1989 (fr), Alvarado 233 (AAU, NY, QCNE); Parque Nacional 

Sumaco Napo-Galeras, block 19, 00?53'S, 77?31'W, 1200 m, 

22 Apr 1996 (fr), Freire & Cerda 461 (QCNE); Parque 

Nacional Galeras, 1 km SW of Comunidad Santa Rosa de Arpino, 

block 19, 00?51'S, 77?31'W, 1050 m, 3 Apr 1996 (fr), 

Vargas & Grefa 908 (QCNE); rd. El Chaco-El Reventador, 

San Rafael waterfall, 1500 m, 77?35'W, 00?06'S, 16 Dec 1993 

(fl), Freire-Fierro & Yanez 2697 (NY, QCNE). PASTAZA: 

Mera, 25 Mar 1940 (fl-fr), Lugo 115 (S[2]); Mera, 1100 m, 

27 Feb 1972 (fl), Harling 11076 (GB[2], TEX[2], US); 1-5 km 

NW ofMera, 7 Mar 1980 (fl),Harling & Andersson 17003 (GB); 

Mera, Rio Pastaza, Hac. La Gloria above El Topo, 1900 m, 

2-9 May 1944 (fr), Steere & Camp 8228 (F[2], NY, US); confluence 

of Rio Quijos with Rio Salado, below Baeza, 1930 m, 2 

Mar 1944 (fl), Ownbey 2682 (F, IAN, US). SucuMBios: San 

Rafael, on rd. El Chaco-Lumbaqui, 1300-1400 m, 77?35'W, 

00?06'S, 20 Jun 1996 (fr), StAhl et al. 2800 (AAU, QCA). 

TUNGURAHUA: Forest of Volcan Tungurahua, Sodiro s.n. (P). 

PERU. HUANUCO: Prov. Pachitea, Pucallpa, W part of 

Sira Mtns., 24 km SE of Puerto Inca, 09?28'S, 74?47'W, 29 

Jul 1988 (fl), WallnOfer 113-29788 (K[2], LZ, NY, USM, 

W). PAsco: Prov. Oxapampa, W side of Cordillera de San 

Matias, between Iscozacin and summit, 75?12'W, 10?1 'S, 

450 m, 22 Jun 1982 (fl), Smith 2075 (MO). 

Dubious collection: Peru. Huanuco: W of Tingo 

Maria, 685 m, 19 Sep 1962 (fl), J. Schunke Vigo 6162 

(F, MO, US). The label data report these specimens 

as being collected from a tree 7-8 m tall, with "moderate 

purplish-pink (2.5 RP 7/8)" flowers. These specimens 

have semi-leathery leaves, 28-32 x 7.5-10.5 cm 

wide; inflorescences sparsely and thinly branched, 36- 

41 cm long; capsules 4-5 mm long and 2-3 mm wide.


101 

Al? 

o~~~~~~~~~~~b 

AI 

10~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

i~~~~~~~~~~~~~? 

I e ' ' i- i i i ' 7 a 1 

FIG. 29. Distribution ofRustia occidentalis (solid circles), R. venezuelensis (open star), R. alba (solid star), R. thibaudi- 

oides (solid squares), R. schunkeana (solid triangles), R. rubra (open circles), R. bilsana (open square) and R. viridiflora 

(open triangles). 

I


S cmn 

.I f 

C' 

t ~ ~ ~ : Flt 

FIG. 30. Rustia rubra (A-D from Harling 11076, US; E from Steere 8021, F, holotype). A. Habit of inflorescence 

(frondose). B. Flower at anthesis. C. Open anther. D. Detail of style. E. Mature capsule.


'':?" ~ 

...... . .. ~~~~..... I 1 _ _ I . 

- JI :,...........: .......... .... 

...': : ."... .:.... ?..................................... .... 

:.:: . : i 

.. : 

*::: _ . '.i; i U l |l ll 

~ ~ ~ ~ ~ ~ ~ ~ ~ ,Pj 

',. :. : : 

FIG.31Rstiarbra inforscec.. wit. foeS i ateI and _mtr .? e,, * fut(poobBeil 'J.,=ns . ' : . ..".. ! ' .ig;'; ....:... .Nx '. i Shl. | I | I I I | 

, ................. 

_ 

_ ~~~~~~~~~~~~~~?:C^',#^?' i ' 

. . .. i _:i : : > 3 -. ! .. . .. . ........................................ 

- . :: :~~~~~~~~~~~~~~~~~~~~~~~ 

? 

~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ':i?i :?. 

* . .. .: ...:'. i. :~ ~ ~ ~ ~~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~.?i~ 

??. i:: 

. .. ..~ .....~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

~~ ~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG. 31.?? Ru rubrairsee s t ia~I~ 

fru it .h 

* i ..i.-s.:'....o.X.'.. }...: .. ii. ; .i' > . ^'.,: i .-,xF }-..::;- . ~ 

FIG. 31. Rustia rubra inflorescence, with flowers in anthesis and StChl).i?ie immature fiuits (photo by Bertil 

The capsules are much smaller than the normal size 

of R. rubra, and probably undersized because infested 

by insects, and with circular exit hole on the 

capsule wall. Because of its semi-leathery leaves, 

sparsely and thinly branched inflorescences, and its 

collection locality, I tentatively attribute these specimens 

to R. rubra. Nevertheless, future collections might 

prove that this collection represents an undescribed 

species of Rustia. 

Simpson (1976) used for this species the name that 

was handwritten by Standley on herbarium specimens. 

It has pendulous flowers and occurs in the Ecuadorian 

and Peruvian Amazon. At the end of the description 

of R. rubra, he stated: "Both R. rubra and R. 

splendens [= R. thibaudioides] have short deltoid corolla 

lobes that are about as long as wide. Rustia splendens 

[= R. thibaudioides] has typically short petioles 

(mostly 1-2 cm long) while in R. rubra they generally 

exceed 2.5 cm, and the corolla tube of the former 

is usually longitudinally striate, longer, and with larger 

diameter than the smaller, non-striate tube of the latter. 

The most obvious character separating these two 

species is the markedly obtuse leaf base of R. rubra 

which contrasts with the broadly acute base of R. 

splendens [= R. thibaudioides]." This set of charac- 

ters is in agreement with my definition of these two 

species, and reliably distinguishes them. 

Rustia rubra differs from R. schunkeana in hav- 

ing the rachis of the inflorescences, flowers, and im- 

mature capsules deep red, corolla lobes '/lo of the co- 

rolla length, inflorescences densely paniculate with 

5-10 lateral branches (vs. racemoid-thyrsoid, un- 

branched to 1-3 branches in R. schunkeana). 

7. Rustia viridiflora Delprete, Novon 5(2): 135, fig. 

2. 1995. Type. Ecuador. Morona-Santiago: Achut- 

za, Jibaro settlement in the vicinity of Macuma, 

ca. 50 km N of Macas, 23 Mar 1973 (fl), Lugo 

3674 (holotype, GB). Figs. 6E, 14A,B, 32A-F 

Tree to 20 m tall, to 20 cm at dbh. Leafy branch- 

lets glabrous to minutely puberulent, terete; older 

branches glabrate, rugulose, grayish to pale brown; 

lenticels sparse, punctiform to linear, to 1 cm long. 

Stipules not seen (readily caducous), leaving a linear


pA 

C 

:^\ .A, 

:1I 

't/.-a --. ; i ' 

' 

C 

FIG. 32. Rustia viridiflora (A-E from Lugo 3674, GB, holotype; E from Prieto CHUP-20). A. Habit of inflorescence with 

mature leaf. B. Abaxial view of blade showing pellucid punctation. C. Flower bud showing bulbose base. D. Open anther. 

E. Detail of receptive style. F. Mature capsule showing conical disk. (Reproduced with permission from Novon 5, fig. 2. 1995.) 

I


scar 1-1.5 mm wide, of the same color of stem. Leaves 

38-69x-9.5-20.5 cm, L/W 3:1 to 3.5:1; lanceolate to 

elliptic, acute to obtuse (35-65?) at base, acute to attenuate 

at apex, sometimes tapering to an acumen 2- 

3 cm long; blade dark green above, dull green below, 

stiff-foliaceous; drying pale brown, stiff-chartaceous; 

glabrous above and below, pellucid punctate; primary 

and secondary veins glabrous, prominent below; secondary 

veins 19-25 each side; tertiary veins starting 

subparallel and openly reticulate in the center, faintly 

evident above, fairly evident below; petioles 4.5-7 cm 

long, 3-5 mm thick, subterete, glabrous; basally thickened 

but not pulvinate; domatia absent. Inflorescences 

laxly paniculate, pyramidal, with decussate 

racemoid lateral branches; 22-54 x 9-22 cm, lateral 

branches 3-7 pairs, basal portion of axis not branched 

10-20 cm long; rachis terete to decussately obtusecompressed, 

rachis and branches glabrous to minutely 

puberulent; flowers on distal racemoid branches or on 

racemoid-cymules; bracts subtending lateral branches 

1-3 x 2-3 mm, deltoid; bracteoles subtending flowers 

0.5-1 x 0.7-1 mm, deltoid, glabrous. Flowers 

pedicellate, pedicels 6-9 mm long, glabrous to minutely 

puberulent; hypanthium narrowly obconical, 4- 

6 x 2-4 mm, glabrous to minutely golden-hirtellous; 

flower buds cylindrical. Calyx reduced to a wavy 

margin with barely distinguishable lobes, 0.3-0.5 mm 

long. Corolla tubular with bulbous base and small 

constriction just above, 1.9-2.2 cm long, green 

throughout (rarely reddish with green lobes), semifleshy 

when fresh; tube subcylindrical, striate, 1.4- 

1.8 cm long, 4-5 mm wide at base and 2-2.5 mm wide 

at the orifice, glabrous or golden-hirtellous outside; 

antrorse-strigose at base inside, without a ring of hairs, 

the remaining medial and superior zones glabrous or 

with sparse hairs; lobes 5, ?-4/5 of corolla length, 3- 

5 x ca. 2.5 mm, triangular to ovate, glabrous outside, 

glabrous, with minute glandular hairs (hairs to 0.5 mm 

long) at margins inside. Stamens 5, included (only 

the very tip exserted), subequal, attached 3.5-5.5 mm 

from the base of the tube; filaments 6-7 mm long, 

sparsely ascending-pilose, with a tuft of golden-pilose 

hairs at base; anthers elongated, rectilinear (never 

convex), 11-12 x 1.2-1.3 mm, dorsifixed near base, 

base subcaudate, with sharp ridges, papillose-echinate 

to ascending-strigulose, descending glandular-pilose 

at base. Pollen exine densely foveolate, sparsely minutely 

echinate. Style exserted, 22-28 mm long, glabrous 

and minutely verrucose throughout; style 

branches ovate, acute at apex, 1-1.3 mm long, stigmatic 

surface smooth to minutely papillose; immature 

fruits green to reddish. Capsules very narrowly 

obconical, apex conical, (14-)18-23 x dehiscence absent even in old capsules. Seeds 0.87- 

1.27 x 0.33-0.83 mm. 


disturbed forests, from low to medium elevations, 

(300-)800-1200 m, of S Amazonian Ecuador. Flowering 

specimens were collected in March, October, 

and November. Fruiting specimens were collected in 

March and November. Its pollination biology has not 

been studied, but its flowers are likely to be pollinated 

by hummingbirds. 

Specimens examined. ECUADOR. MORONA- 

SANTIAGO: Near Lim6n (General Plaza Gutierrez), 

roadside in remnant forest, 1200 m, 27 Mar 1981 (fr), 

Berg 1227 (QCA); rd. Bella Union-Limon, km 30, 

78?17'W, 02?48'S, 8 Nov 1991 (fl), Persson et al. 43 

(QCA); low hills W of Rio Upano, along Rio Chupiangas, 

800-1050 m, 20-22 Nov 1944 (fr), Prieto 

CHUP-20 (NY[2], US). PASTAZA: Cushillo Urco, ca. 

8 km N of Puerto Sarayacu, 6 Oct 1974 (fl), Lugo 3946 

(GB, US); Chapeton on Rio Bobonaza, ca. 8 km ESE 

of Canelos, 400-450 m, 77?40'W, 01?38'S, 2-5 Nov 

1995 (fl), St&hl et al. 1822 (AAU, QCA); trail between 

Chapet6n and Canelos, 400-450 m, 77?43'W, 01 ?35'S, 

6 Nov 1995 (fl-fr), Stahl et al. 1866 (AAU, QCA). 

Rustia viridiflora is unique in the genus in its long 

narrowly obconical capsules with conical disk, corollas 

basally bulbose, and long rectilinear anthers. Its 

corollas are commonly entirely green (rarely red tube 

with green lobes). This is one of the four species of 

Rustia with pendulous flowers (Delprete, 1996a), all 

of which have long-narrow corollas and occur in western 

South America. A key to the pendulous-flowered 

species of Rustia was published by Delprete (1996a). 

8. Rustia schunkeana Delprete, Brittonia 48: 29, fig. 

1. 1996. Type. Peru. San Martin: Prov. Mariscal 

Cacheres, Tocache Nuevo, NW of the nursery of 

Instituto Agropecuario de Tocache, 20 Apr 1970 

(fl), J. Schunke Vigo 3952 (holotype, COL; 

isotypes, F, G, NY, US). Figs. 6A, 13E, 33A-D 

Shrub 3-5 m tall, much branched, with slender 

branches, outer branches often scandent. Leafy branchlets 

glabrous, dark green, terete; older branches pale 

brown to grayish; lenticels sparse, punctiform to linear. 

Stipules present only in bud, narrowly triangular, 

glabrous outside, glabrous, with few colleters in- 

4-6 mm, black, side, 30-40 x 5-6 mm, dark green, readily caducous, 

without lenticels, glabrous throughout; disk septicidal leaving a linear whitish scar, ca. 0.5 mm wide. Leaves


n t 

E 

FIG. 33. Rustia 

schunkeana (A-C from Schunke Vigo 3952, COL, holotype; D from Schunke Vigo 6696, F). A. Habit 

of inflorescence with mature leaf. B. Open flower and flower bud. C. Open anther. D. Mature capsule. (Reproduced with 

permission from Brittonia 

48, fig. 1. 1. 1996.) 

30-45(-60) x 8.5-12 cm, L/W 3:1 to 3.5:1; narrowly 

lanceolate, oblanceolate to narrowly elliptic, acute 

(BA = 20-30?) at base, acute at apex, sometimes tapering 

to an acumen 2-3 cm long, dark green above, 

pale green below, subcoriaceous; drying brown, stiff- 

chartaceous; glabrous above, glabrous to hirsutulous 

below, pellucid punctate; primary and secondary veins 

glabrous to hirsutulous, prominent below; secondary


veins 15-25 each side; tertiary veins starting subpar- 

allel and openly reticulate in the center, evident be- 

low; petioles 2-7.5 cm long, 2.5-4.5 mm thick, 

subterete, glabrous to hirsutulous, basally thickened 

but not pulvinate; domatia absent. Inflorescences 

racemoid secundiflorous panicles to thyrsoid- 

secundiflorous with opposite or subopposite decus- 

sate branches, sometimes without lateral branches, 

25-60 cm long, to 17 cm wide, basal branches to 20 

cm long, lateral branches usually ascending and to 4 

pairs, basal portion of axis not branched 2-15 cm long; 

rachis terete, rachis and branches glabrous (rarely hir- 

sutulous); flowers alternate on distal branches or 

rarely in cymules, usually hanging downward, secund 

(pedicel attachments opposite or alternate); distal 

bracts 1-3 x 2-3 mm, deltoid; bracteoles subtending 

flowers 0.5-0.8 x 1-2 mm, deltoid, glabrous. Flow- 

ers pedicellate, pedicels 4-25 mm long, glabrous to 

hirtellous; hypanthium narrowly obconical, 4-6 x 2- 

2.5 mm, glabrous to hirtellous; flower buds cylindri- 

cal. Calyx reduced to a wavy margin with barely dis- 

tinguishable lobes, ca. 1 mm long. Corolla tubular 

with small erect lobes (not spreading), 2.5-4 cm long, 

greenish yellow (rarely pinkish yellow), semi-fleshy 

when fresh; tube subcylindrical, faintly striate, 1.5- 

2.5 cm long, 2.5-4 mm wide at base and 3.5-5 mm 

wide at the orifice, glabrous outside; antrorse-strigose 

at base inside, without a ring of hairs, the remaining 

medial and superior zones glabrous; lobes 5, 4-'/3 of 

corolla length, 9-13 x 1.5-3 mm, triangular to ovate, 

glabrous outside, glabrous, with minute conical-bullate 

hairs (hairs 0.1-0.3 mm long) inside. Stamens 5, 

included (only the very tip exserted), subequal, attached 

4.5-5.5 mm from the base of the tube; filaments 

6-7 mm long, flattened at base, microscopically 

(40x) papillose throughout, minutely barbate at base 

with white puberulent hairs; anthers cylindrical, rectilinear 

(never convex), 9-12 x 1.5-2 mm, dorsifixed 

near the base, base elliptic, smooth throughout. Pollen 

exine densely foveolate (non-echinate). Style exserted, 

30-40 mm long, glabrous, microscopically 

(40x) papillose; style branches ovate, 1-1.5 mm long, 

stigmatic surface smooth; immature fruits reddish 

green, semi-fleshy. Capsules obovoid to oblong, apex 

shallowly hemispherical, (10-)15-20 x 7-9 mm, without 

lenticels, dark brown, glabrous throughout; disk 

black, obviously exceeding the calyx; disk septicidal 

dehiscence absent even in old capsules. Seeds 1.17- 

1.67 x 1-1.33 mm. 


secondary lowland rain forests (200-350 m) to pre- 

montane wet forests (to 2000 m), often on river banks, 

on the W Andean slopes of Amazonian Ecuador and 

Peru. Found in association with Grias, Couepia, 

Clusia, Ficus, Guarea, Geonoma, Mabea, and Ardisia 

(Jaramillo 14167,14290). Flowering specimens were 

collected in March, May, and June. Fruiting specimens 

were collected in February, June, August, September, 

and December. 

Specimens examined. ECUADOR. NAPO: Nor-Oriente, 

Nuevo Rocafuerte, SW of village, forest toward Rio Braga, 

200 m, 27 Feb 1981 (fr), Jaramillo & Coello 4322 (QCA); 

Archidona-Tena region, W of Tena, along creek between Rio 

Lupi-yacu and Rio Pano, near foot of cordillera de 

Guacamayos, 800 m, 25 Mar 1944 (fl), Ownbey 2735 (F[2], 

IAN, MO[2], US[2]); Cerro Antisana, 2 km SW of Tena, 

00?30'S, 78?W, 700 m, 27 Aug 1960 (fl-fr), Grubb et al. 1485 

(K, NY[2]). ZAMORA-CHINCHIPE: Canton Nangaritza, Rio 

Nangaritza, between Pachicutza and Miazi, 78?38'W, 

04?12'S, 1000 m, 12 Dec 1990 (fr), Neill & Palacios 9718 

(MO, QCNE, TEX[2]); 19 Dec 1990 (fr), Palacios 6616 

(MO); left margin of Rio Nangaritza, near Miasi military 

camp, 04?20'S, 78?40'W, 900 m, 20 Oct 1991 (fr), Jaramillo 

14167 (QCA), 950 m, 22 Oct 1991 (fr), Jaramillo 14290 

(NY, QCA). 

PERU. HUANUCO: Villa Isabel, Rio Cuchara, 21 Sep 

1961 (fr), Schunke Vigo 5683 (F[2]). SAN MARTIN: Prov. 

Mariscal Caceres, Dtto. Tocache Nuevo, trail to Santa Rosa, 

Rio Mishollo, 350 m, 4 Aug 1973 (fr), Schunke Vigo 6696 

(F, MO, NY[2]); Prov. Mariscal Caceres, Dtto. Tocache 

Nuevo, Prov. Mariscal Cacheres, Santa Rosa de Mishollo, 

4 km from Puerto Pizana, 8 May 1971 (fl), Schunke Vigo 

4877 (COL, F, G, NY, US); Prov. Mariscal Caceres, Dtto. 

Tocache Nuevo, Bambamarca, 20 Jun 1978 (fr), Schunke 

Vigo 10270 (F, MO, NY); Prov. Lamas, 2-4 km N of San 

Antonio, along Rio Cumbasa, 400 m, 2 Oct-4 Nov 1937 (flfr), 

Belshaw 3514 (F, GH, LL, MO, NY, UC, US); Dtto. 

Uchiza, 500 m, 23 Jun 1969 (fl), Schunke Vigo 3205 (COL, 

F, G, NY, US, VEN). 

Local names. Peru: cascarilla masha (San Martin, 

Schunke 3952), sacha cascarilla (Huanuco, Schunke 

Vigo 5683), cascarillita (San Martin, Schunke Vigo 

6696, 10270). 

Rustia schunkeana is similar to the recently described 

R. viridiflora (Delprete, 1995) in having greenish 

corollas, the former differing from the latter in 

having thyrsoid (vs. paniculate) inflorescences, cylindrical 

flowers 25-40 mm long (vs. basally bulbose, 

19-22 mm long), and capsules ovoid to oblong (vs. 

narrowly obconical). 

Rustia schunkeana has yellow-green flowers 

(rarely with pinkish base), while R. thibaudioides has 

pink-red flowers with white apices, and R. rubra has 

deep-red flowers sometimes with green apices. Rustia 

schunkeana also differs from R. thibaudioides in hav- 

ing corolla lobes l/4--13 of the corolla length (vs. '/o-'/8), 

oblong-obovoid capsules 15-20 mm long (vs. obo-


void, 8-12 mm long), and disk exceeding the calyx 

(vs. not exceeding the calyx). 

The specimens of Jaramillo 14290 (NY, QCNE) 

have been observed to have large, fleshy fruit galls. 

The galls develop from the base of the ovary, and 

enlarge laterally into an irregular-obovoid shape, 1.5- 

2.2 x 0.9-1.5 cm. When sectioned, the galls have longitudinal 

burrows of minute larvae of unknown identity 

(Coleoptera?). 

9. Rustia thibaudioides (H. Karsten) Delprete, comb. 

nov. Henlea thibaudioides H. Karsten, Linnaea 30: 

151. 1859. Type. Colombia. Meta: Mun. San Juan 

de Arama, Reserva Nacional Serra de la Macarena, 

2 km before La Danta research station, 6 Jan 1993 

(fl-fr) Delprete & Fagua 6378 (neotype, NY, selected 

here; isoneotypes, COL, F, MO, NY[2], 

TEX). Figs. 4C, 6C-D, 13F, 34A-E, 35A-C 

Rustia splendens (H. Karsten) Standley, Publ. Field. 

Columbian Mus., Bot. Ser. 7: 26. 1930. Henlea 

splendens H. Karsten, Fl. Columb. 1: 157, pl. 78. 

1861. Type. Colombia. Meta: Llano de San Martin, 

Jiramene, vic. of Rio Meta, Karsten s.n. (lectotype, 

W, selected by Delprete, 1996a; isolectotype, LE?). 

According to Tryon (1963), the first set of Karsten's 

collections is preserved at LE, but no material has 

been seen from this institution. Also, plate N. 78 in 

Flora Columbiae is a beautiful color illustration of 

the type. 

Rustia secundiflora K. Schumann in Martius, Fl. Bras. 

6(6): 261. 1889. Type. Colombia. Caqueta: Upper 

Amazonas, at base of Mt. Araracoara, Jan-Feb (fl), 

Martius 3139 (B*; lectotype, M, selected by 

Delprete, 1996a; photo-M at F, MO, VEN). 

Shrub 5-6 m tall or tree 8-10(-12) m tall, to 25 cm 

dbh, much-branched shrub to much-branched singlestemmed 

tree; bark yellowish-grayish. Leafy branchlets 

glabrous to golden-pubescent, dark green, terete; 

older branches glabrate, rugose, grayish to pale brown; 

lenticels sparse, small. Stipules narrowly triangular, 

glabrous to minutely puberulent to hirtellous outside, 

glabrous with few colleters inside, 20-30 x 4-5 mm, 

dark green, readily caducous, leaving a grayish scar 

encircling the stem, evident in older branchlets. 

Leaves 25-50(-65) x (6-)10-17 cm, L/W 2:1 to 3.5:1; 

lanceolate to oblanceolate to obovate, acute to rounded 

(BA = tellous (rarely hirsute) below; pellucid punctate; primary 

and secondary veins glabrous to hirtellous 

(rarely hirsute), prominent below; secondary veins 

11-18 each side; tertiary veins starting subparallel and 

openly reticulate in the center, faintly evident below; 

petioles 1.5-6 cm long, 1.5-4.5 mm thick, terete to 

adaxially concave to flattened, pulvinus present in 

young leaves, becoming corky in older leaves; 

domatia absent. Inflorescences openly to densely 

paniculate to thyrsoid-secundiflorous with opposite or 

subopposite decussate branches; (12-)25-70(-153!) 

cm long, 5-25(-45) cm wide at base, lateral branches 

to 15 pairs, basal portion of axis not branched 5-25 

cm long; rachis terete to decussately obtuse-compressed, 

rachis and branches glabrous or puberulent 

to hirtellous; flowers alternate on distal branches or 

rarely in cymules, usually pointing downward, giving 

the false appearance of secundiflorous position 

(pedicel attachment opposite and not secundiflorous); 

distal bracts 1-4 x 2-4 mm, deltoid; bracteoles subtending 

flowers 0.5-1 x 0.5-1 mm, deltoid, glabrous 

to minutely puberulent. Flowers pedicellate, pedicels 

3-15 mm long, glabrous, puberulent to hirtellous, 

often curved downward; hypanthium narrowly 

obconical, 2-5 x 1.5-2.5 mm, glabrous to puberulent; 

flower buds cylindrical. Calyx extremely reduced, 

truncate or with minutely wavy margin, 0.3-1 x 2-4 mm, 

glabrous to puberulent. Corolla tubular, 2-2.5(-3) cm 

long, white apically and pink-reddish at base outside, 

white inside, semi-fleshy when fresh; tube subcylindrical, 

striate, 1.9-2.2 cm long, 2.5-3.5 mm wide at 

base and 2.5-3.5 mm wide at the orifice, glabrous to 

puberulent to hirtellous outside; antrorse-strigose to 

strigulose at basal 2-3/4 of the tube inside, without a 

ring of hairs; lobes 5, '/,o-'/ of corolla length, 1.5-4.5 

x 1.5-2.5 mm, deltoid, glabrous to minutely puberulent 

outside, glabrous, minutely papillose (40x) at 

margins inside. Stamens 5, included or only the tips 

exserted, subequal, attached 5-7.5 mm from the base 

of the tube; filaments 7-8.5 mm long, minutely (40x) 

ascending strigulose to papillose at lower half portion; 

anthers cylindrical, narrowly oblong, rectilinear or 

slightly convex, 5-6.5 x 0.9-1.4 mm, dorsifixed near 

the base, base rounded, microscopically (40x) papillose-echinate 

(rarely smooth) throughout, dehiscing 

by two pores at apex. Pollen exine foveolate, minutely 

echinate. Style exserted, 30-35 mm long, glabrous, 

subtetragonous; style branches ovate, 0.4-1 mm long, 

stigmatic surface smooth. Capsules obovoid to widely 

20-40?) at base, acute to obtuse-acuminate at obovoid, acute at base, apex subtruncate to obtusely 

apex, the acumen, when present, 1-2 cm long, dark hemispherical, 8-12 x 6-7(-10) mm, dark brown 

green above, pale olive-green below, foliaceous to without lenticels, glabrous to puberulent below the 

semi-coriaceous; drying green-brownish, stiff-chart- disk; disk glabrous; disk septicidal dehiscence absent 

aceous; glabrous above; glabrous, puberulent to hir- even in old capsules. Seeds 1.4-1.5 x 0.75-1.25 mm.


l I l 

'4i ~A 

C CA 

FIG. 34. Rustia thibaudioides (from fresh material and Delprete & Fagua 6378, TEX). A. Habit of inflorescence with 

mature leaves. B. Flower, lateral view and inside view. C. Flower bud. D. Open anther. E. Mature capsule. (Reproduced 

with permission from Brittonia 48, fig. 2. 1996 [as R. splendens].)


:., ...... .,,,,~! 1 

F -? 6..o,2'?C~ 

. 

...' ....~ 

.I~~~~~~~~~~~~~~~~~~~~~~~~~N 

? .: 

? , '": .. ... 

_ :.f 

'' ,-' ;' 

. 

' 

; ,............. .. 

''~~~~~~~~~~~~~:;:,? i.? : 

8 ...'"....~., .. ,, ~:'* :';, t.';..~....: ... :?:, ~ 

.. ~!._ , ,7 . ~,..>...:i, ......:...,. 

.~ ~ = ,~ ,'. 4 , ................ ..... 

~~??Xl. _ d? _ .:~~?~ . , 

~.~~~~~~~~~~~~~~~~~g :i:.r': 

|~~~~~~~~~~~~~~~~~~~~~~~~~::? I~ 

?? 

. 35.: Rd . A. Sal (o_, a mI .U..': c,i 

'' .;r r iBsFI ~ 

I 11 '&Q i' 

anhsi, C.' Flwri"nhsssenfo otmuwad(oeicue ope'n anthers.. 

: -: 

i?-. 

r 

;' " 

. 

. .... 

, 

I"' .. . ..:_ ii .. @ " . _: __. 

'~~~l ,'~' :_i;,,, ,?-_4-_'"". 

,._~~~ o -. 

fiF: .-- 

. _ 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?...L|. 

.. '1:"~~' , 

:..~ie 

FIG. 3S. Rustia thibaudioides. A. Sh--b F5 m tall (arrow indicates main trunk). B. Inflorescen.c ca. I m long, in 

anthesis. C. Flower in anthesis seen f.om bottom upward (note included open anthers). 

r 

.


Distribution (Fig. 29) and ecology. Gallery forests 

of central (states of Boyaca, Cundinamarca, Santander, 

Tolima) and Amazonian Colombia, to 1400 

m, in lateritic soils. A single specimen was collected 

in Venezuela (state of Tachira), and a few specimens 

were reportedly collected in Brazil and their labels 

read: "Martius Iter Brasiliense, Rio Negro, Brazil." 

This species has long inflorescences with up to a few 

hundred flowers, and the blooming period of a single 

individual could last up to two months. Flowering 

material was collected in January, February, March, 

July, October, November, and December. Specimens 

in early fruiting stage have been collected in, January 

and December. 

Near Rio Negro, E of Prado, 10 Feb 1944 (fl), Little 7186 

(COL, US). VAUPtS: Environs of Mitfi, trail to Cerro Miti, 

29 Oct 1976 (fl), Davis 184 (COL, F, GH); cabeceras of Rio 

Cuduyary, 10 Dec 1943 (fl), Alien 3217 (COL, G, MO, US); 

Rio Vaupes, Miti, 9 Oct 1966 (fl), Schultes 24356 (COL). 

VENEZUELA. TACHIRA: Ca. 35 km SSE of San Crist6bal, 

La Buenafia, 6-12 km W of Quebrada Colorado, 600- 

1200 m, 07?28'N, 72?09'W, 20-21 Mar 1981 (fr), Liesner 

& Gonzdles 10940 (VEN). 

BRAZIL. AMAZONAS: Rio Negro, Martius Iter Brasiliense 

2607 (M), 2608 (M), s.n. (M). 

Local name. Colombia: doncello (Meta, Fosberg 

19440). 

Pollination biology. This species was observed in 

Rustia thibaudioides is unique in the genus in having 

apically white and basally pink corollas, included 

bloom in the Serrania de La Macarena (Department anthers, deltoid corolla lobes 1/lo-'/8 of the corolla 

of Meta). The flowers of R. thibaudioides are mainly length, and obovoid capsules 8-12 mm long with disk 

pendulous, sometimes as many as several hundred per not exceeding the calyx (Fig. 34B-E); the closely reinflorescence, 

pinkish red at base and white at the lated R. schunkeana has deltoid corolla lobes ?4-'/3 

medio-distal portion, and the anthers remain included of the corolla, anthers excluded at anthesis, and obin 

the corolla tube at anthesis. They are frequently long-obovoid capsules 15-20 mm long (Fig. 33B-D). 

visited by hummingbirds, bees, wasps, and butterflies. 

The bees most frequently visiting the flowers are of 

the genera Melipona, Ceratina, and Trigona (2 spp.) 

(pers. obs.). No detailed pollination biology studies 

were performed, but I suspect that this species is pollinated 

primarily by hummingbirds and secondarily 

by bees. Some flowers were observed to be robbed 

by wasps through lateral incisions. Many individuals 

were also guarded by ants, which are probably feeding 

on the easily accessible floral nectaries. 

Karsten (1859) cited Henlea thibaudioides, the 

type species of Henlea, as "Crescit in planitie ad orientalem 

Cordillerae Bogotensis ad littora fluminis 

Meta" [Colombia, Cordillera Oriental, Rio Meta]. No 

types of this species were found at W, and I had no 

access to the LE specimens. Because of this, I am 

selecting as the neotype for H. thibaudioides a specimen 

ofDelprete & Fagua 6378 (NY) that corresponds 

to its original description and was collected close to 

the locality cited by Karsten (Serrania de la Macarena, 

near the riverwash of Rio Meta). Henlea thibaudioides 

Representative specimens examined. COLOMBIA. is here transferred to Rustia, and Rustia and Henlea 

ANTIOQUIA: San Rafael, 29 Nov 1958, Daniel 5533 (US). are congeneric. 

BOYACA: Llano de San Martin, Bogoti, 300 m, [1851-1857], A discussion of the lectotypification and synonymy 

Triana 1785 (3262.4) (P); Llano de San Martin, Jiramena, of Rustia splendens and R. secundiflora was presented 

220 m, Jan 1856 (fl), Triana s.n. (3262) (F, COL). CAQUETA: in 

Rd. Puerto Rico-Florencia, left 

Delprete, 1996a. The 

side of Rio Guayas, 350lectotype 

ofHenlea splendens 

400 m, 7 Oct 1975 (fl), Cabrera 3701 (F); confluence of Rio (= R. thibaudioides) is a specimen preserved at W, 

Mesay with Rio Cufiar6, 1-6 Mar 1980 (fl), Pabdn 960 

with embossed "Dr. Karsten," and labeled in his hand- 

(COL[2]). CUNDINAMARCA: Near Guayabetal, trail to Manza- writing "Henlea splendens - Llano de St. Martin." The 

nares, Valley of Rio Negro, Western Cordillera, 1400 m, 3 lectotype of R. secundiflora is a specimen preserved 

Jan 1956 (fl-fr), Vogel 54 (VEN). GUAVIARE: Cafio Grande, at M, with its label stating "Mart. Obs. 3139 - ad M. 

between Calamar and San Jose del Guaviare, 240 m, 1 Nov Araracoara - Dr. Martius Iter Brasil: Jan-Feb." On a 

1939 (fl), Cuatrecasas 7370 (F). META: Uribe, Cordillera small label glued to the herbarium sheet, Martius an- 

Oriental, 770 m, 20 Dec 1942 (fl-fr), Fosberg 19440 (NY[2], notated this specimen as "Hamelia albopurpurea," an 

UC, US[2]); Mun. San Juan de Arama, Reserva Nacional 

unpublished name. 

Serra de la Macarena, 2 km before La Danta research sta- 

Rustia warscewicziana Klotzsch is another 

tion, 6 Jan 1993 (fl-fr) Delprete & Fagua 6381 

unpub- 

(COL, F, MO, 

lished name for this taxon. The 

TEX), 6382 (COL, NY); Caflo Lozada, riverside of Rio 

corresponding her- 

Guayabero, 350 m, 30 Jan 1959 (fl), Pinto et al. 370 

barium 

(COL, specimens were preserved at Berlin (B) and 

P). PUTUMAYO: Rio San Miguel, left affluent Quebrada La were destroyed. Fortunately the specimens were pho- 

Hormiga, 290 m, 15 Dec 1940 (fl), Cuatrecasas 11076 tographed by Macbride (negatives N. 10 and 11) and 

(COL, F, US). SANTANDER: Region around Landazuri, 70 km the photographs distributed to several herbaria (F, 

N of Velez, 1 Jul 1944 (fl), Fassett 25427 (US). TOLIMA: MO, NY, US). These specimens were collected by


Warscewicz in Colombia ("Neu Grenada"), in the tending primary branches 1-3 x 1-2 mm, deltoid; 

southern Cordillera Occidental ("cordillera mer. bracteoles subtending flowers ca. 0.5 mm long, deloccident."), 

an area very close to the type locality of toid. Flowers not seen. Pollen not seen. Capsules 

Henlea splendens (= R. thibaudioides). Simpson subglobose to obovoid, acute at base, apex hemi- 

(1976) studied the two photographs, and referred to spherical, 6-9 x 6-6.5 mm, dark brown to black with- 

R. warscewicziana as follows: "It is certainly a Rustia out lenticels, glabrous throughout; disk black, exceedbut 

the panicle seems much larger and more branched ing the calyx; pedicels below mature capsules 9-15 

that any I have seen otherwise." After close exami- mm long. Seeds 0.93-1 x 0.4-0.57 mm. 

nation of the photographs I concluded that the destroyed 

specimens pertain to R. thibaudioides. Distribution (Fig. 29) and ecology. This species 

Extensive field research has been carried out in the is known only from the type specimens, reportedly 

area where the three types were obtained, and I have collected from a tree 10-12 m tall growing on dry 

collected R. thibaudioides at several localities. This canyon slopes, ca. 1550 m, in the state of Merida, 

species is a very beautiful shrub, usually 6-7(-10) m Venezuela. The single fruiting specimen was collected 

tall, with very attractive inflorescences that vary in in May. Flowering specimens are still unknown. 

size and shape, sometimes to 1.5 m long, most of the 

time horizontal and sometimes ascendent. In the in- This is the only species of Rustia collected in Venflorescences 

of R. thibaudioides the flowers are pen- ezuela. Steyermark (1974) also reported R. longifolia 

dulous, which in herbarium specimens have been er- Standl., but this taxon is instead Dolichodelphys 

roneously interpreted as secund. 

chlorocrater. 

Rustia venezuelensis is similar to R. formosa, of 

southern Brazil; the former is presently kept distinct 

from the latter because of its smaller sparsely flow- 

10. Rustia venezuelensis Standley & Steyermark, ered inflorescence and its very isolated geographical 

Fieldiana, Bot. 28: 616. 1953. Type. Venezuela. distribution. Future collections of flowering speci- 

Merida: Between Mucuchachi and Canagua, be- mens will be crucial in establishing the relationships 

low El Bao, 1570 m, 6 May 1944 (fr), Steyermark of this species. 

56333 (holotype, F; isotype, VEN). Fig. 6G,H 

Tree 10-12 m tall, much branched; bark grayish. 

Leafy branchlets glabrous, pale brown-grayish, terete; 

older branches rugose, white-grayish. Stipules not 

seen (readily caducous), leaving a linear scar ca. 0.5 

mm wide, of the same color as stem. Leaves 20-37 x 

8-14 cm, L/W 2:1 to 2.5:1; widely obovate, acute (BA 

?30?) at base, narrowly obtuse, acuminate at apex; the 

acumen deltoid, 1-1.5 cm long; olive-green above and 

pale green below, subcoriaceous; drying olive-green, 

semi-coriaceous; glabrous above and below; pellucid 

punctate; primary and secondary veins glabrous, 

prominent below, secondary veins 12-15 each side; 

tertiary veins starting subparallel and openly reticulate 

toward the center, evident only above; petioles 

15-35 mm long, 2-3 mm thick, subterete or adaxially 

flattened, pulvinus absent but thickened becoming 

corky at base, glabrous; domatia absent. Inflorescences 

reduced paniculate, with opposite decussate 

ascending branches; 17-28 x 13-24 cm, basal primary 

branches 8-15 cm long, lateral branches 3-6 pairs, 

basal portion of axis not branched ca. 7 cm long; rachis 

terete or decussately obtuse-compressed, rachis 

and branches glabrous; flowers alternate on distal 

branches or in terminal lax cymules; distal bracts sub- 

11. Rustia formosa (Chamisso & Schlechtendal ex 

de Candolle) Klotzsch in Hayne, Getr. Darstell. 

Gew. 14: tab. 15. 1846. Exostema formosum 

Chamisso & Schlechtendal ex de Candolle, Prodr. 

4(4): 361. 1830. Exostema formosum Chamisso 

& Schlechtendal var. fi laeve Chamisso & 

Schlechtendal ex de Candolle, nom. inval., Prodr. 

4(4): 361. 1830. Exostema formosum Chamisso 

& Schlechtendal, nom. nud., Linnaea 4:179. 1829. 

Exostema formosum var. fi laeve Chamisso & 

Schlechtendal, nom. inval., Linnaea 4:179. 1829. 

Type. Brazil. Sao Paulo: Estagao Biologica Alto 

da Serra, 19 Oct 1931 (fl-fr), C. Lemos 28678 (neotype, 

NY, selected here; isoneotypes, F, GH, US). 

Figs. 3C, 5C, 14E, 36A-E 

Rustia sellowiana Klotzsch in Hayne, Getr. Darstell. 

Gew. 14: tab. 15. 1846 (validated by direct refer- 

ence to description of Exostema formosum var. 

leprosum Chamisso & Schlechtendal). Exostema 

formosum Chamisso & Schlechtendal var. leprosum 

Chamisso & Schlechtendal ex de Candolle, nom. 

inval., Prodr. 4(4): 361. 1830. Exostemaformosum 

var. leprosum Chamisso & Schlechtendal, nom.


inval., Linnaea 4: 179. 1829. Type. Brazil. Rio de 

Janeiro: Corcovado, Caminho de Larangeiros, Jan 

1834 (fl), Luschnath 179 [Martius H. Fl. Bras. 111] 

(neotype, BR, here selected). Martius H. Fl. Bras. 

111 (B*, BR, frag-F, G, K, M, NY, P) are dubious 

isoneotype specimens because their labels report 

various collectors, localities, and dates. In Herbarium 

Florae Brasiliensis, Martius (1837) reports N. 111 

as "Exostemma formosum Cham. et Schlecht. 

Linnaea IV. p. 179. Quina do Rio de Janeiro: 

Brasiliens. Arbor 25-pedalis. - Est forma leprosum 

Cham. et Schlecht. 1. c. In monte Corcovado et in 

Caminho de Larangeiras: fl. Jan. Dryas." 

Rustia pohliana Klotzsch in Hayne, Getr. Darstell. Gew. 

14: tab. 15. 1846. Type. Brazil. Goias: Ponzo Alto a 

Caldas Nuovas, 1837 (fl), Pohl 809 (lectotype, W, 

here selected; isolectotypes, F, GH). 

Rustia mosenii K. Schumann ex Glaziou, nom. nud., Bull. 

Soc. Bot. France 56, mem. III: 337. 1909. The fourword 

description does not distinguish this from other 

species (Glaziou, 1909). Cited specimens: Glaziou 

21548a (B*, C-n.v., K). 

Shrubs 4-6 m tall, exceptionally trees to 12 m tall, 

to 30 cm dbh, much-branched, sometimes with small 

buttresses; bark yellowish grayish. Leafy branchlets 

grayish-tomentulous, tetragonous-terete; older 

branches glabrate, rugose, terete, grayish; lenticels 

punctiform, 0.5 mm wide. Stipules narrowly triangular, 

acuminate, tomentulous outside, glabrous with 

basal colleters inside, (15-)25-35 x 13-16 mm, pale 

green, readily caducous, leaving a white-grayish linear 

scar. Leaves (22-)25-38(-54) x (7-)9-17(-23) 

cm, L/W 1.6:1 to 2:1; elliptic to oblanceolate to obovate, 

acute to obtuse (BA = x 1-1.5 mm, deltoid, glabrous to minutely puberulent, 

often ciliolate. Flowers pedicellate, pedicels 3-5 mm 

long, glabrous (rarely minute puberulent), fragrant; 

hypanthium narrowly obconical, 3-5 x 1.5-2.5 mm, 

glabrous; flower buds cylindrical. Calyx smallcupular, 

truncate or with barely distinguishable lobes, 

3-4 mm wide; lobes 5, 0.2-0.4 x 0.9-1.2 mm, shallowly 

deltoid, ciliolate. Corolla tubular with spreading-reversed 

lobes, 1.5-1.8 cm long, white to creamwhite, 

rarely pink-reddish at base outside, white 

inside, semi-fleshy when fresh; tube narrowly funnelform, 

9-12 mm long, 2-2.2 mm at base and ca. 2.5 

mm wide at the orifice, glabrous to minutely puberulent 

outside, glabrous inside, with a ring of white-sericeous 

hairs (0.5-1 mm long), 4-5 mm from the base, 

at the same point of the filament attachments at base 

of lobes; lobes 5, '/3-l of corolla length, 5-7 x 0.9- 

1.2 mm, narrowly triangular, glabrous to minutely 

puberulent outside, white-strigose at base (hairs 0.5- 

1 mm long), margins densely papillose inside. Stamens 

5, exserted because of spreading lobes, subequal, 

attached 4-5 mm from the base of the tube; filaments 

4-5 mm long, very thin, pendulous, glabrous, barbate 

at base, with a tuft of white sericeous hairs (0.8-1 mm 

long); anthers cylindrical, narrowly oblong, 6-8 x 1- 

1.5 mm, dorsifixed near the base, base rounded, mi- 

30-50?) at base, rounded 

at apex; dark green above, pale green-grayish below, 

semi-coriaceous; drying grayish olive-green, chartacroscopically 

(40x) papillose-echinate throughout. 

Pollen exine densely foveolate (non-echinate). Style 

exserted, 20-25 mm long, glabrous; style branches 

oblong, 0.9-1.1 mm long, stigmatic surface glabrous 

to microscopically (40x) papillose. Capsules obovoid 

to subglobose, acute at base, apex subtruncate, 5-7 

(-8) 

ceous; glabrous above, glabrous to golden-pilose below; 

pellucid punctate; primary and secondary veins 

glabrous to golden-pilose, prominent below, secondary 

veins 12-16 each side, subparallel; tertiary veins 

openly reticulate, faintly evident; petioles (10-) 13- 

18(-25) mm long, 2.5-3 mm thick, adaxially concave 

to flattened, glabrous to golden-pilose, pulvinus absent; 

domatia absent or a tuft of sparse hairs 1-1.5 mm 

wide, hairs white 0.2-0.4 mm long. Inflorescences 

openly paniculate, pyramidal, with opposite or subopposite 

decussate branches; 24-30(-36) cm long, (8-) 

14-20 cm wide at base, lateral branches 4-8 pairs, 

basal portion of axis not branched (4-)6-14 cm long; 

rachis decussately compressed or quadrangular (rarely 

terete) in cross section, rachis and branches glabrous 

(rarely minute- puberulent); flowers alternate on distal 

branches; distal bracts decreasing distally from 12 

to 3 mm long, 1-1.5 mm wide, linear, narrowly triangular 

to deltoid, bracteoles subtending flowers 1-1.5 

x ca. 5 mm, dark brown with small lenticels, glabrous 

throughout. Seeds 1.4-1.6 x 0.48 mm. 

Distribution (see Fig. 38A) and ecology. Margins 

and gaps in mesophytic gallery forests and Cerrado, 

usually in proximity of creeks and fast rivers of SE 

Brazil in the states of Goias, Minas Gerais, Distrito 

Federal, Rio de Janeiro, and SAo Paulo, at 100-1000 

m. Flowering specimens were collected in all months 

of the year except December and January. Fruiting 

specimens were collected in June, August, October, 

and November. 

Specimens examined. BRAZIL. DISTRITO FEDERAL: 

Cachoeirinha, affluent of left margin of Rio Paranoa, 26 May 

1982 (fl), Pereira 270 (K); hills ca. 10 km N of Planaltina, 

975 m, 2 Oct 1965 (fl-fr), Irwin et al. 8881 (MO, NY, US, 

VEN); 50 km E of Brasilia, 700-1000 m, 20 Aug 1964 (flfr), 

Irwin & Soderstrom 5369 (F, NY, UC, US, VEN); S slope 

ofMorro da Canastra, 920 m, 15?35'S, 47?54'W, 19 Oct 1983 

(st), Kirkbride 5431 (F, NY); estrada de Sobradinho, 24 Jan 

1965 (fl), Sucre & Heringer 624 (IAN, VEN); Sumare, 23 

Feb 1959 (fl), Pereira & Duarte 4484 (RB); between


t4r 

0~~~~~~~~~~~~ 

tl 

.. 

.~~~~~~~~~ I~~~~~~~~~~~~~ 

.... 

,'.~~~~~~~~~~~~' 

FIG. 36. Rustiaformosa (A-D from Pereira 270, MO; E from Irwin et al. 8881, NY). A. Habit of inflorescence with 

view of blade showing pellucid punctation. E. Mature capsule.


Calcareo and Silva, 1 May 1963 (fl), Pires et al. 9539 (F, ofMartius 111 reports the common name "Quina do 

NY); Parque Municipal do Gama, 1000 m, 21 Mar 1966 (fl), Rio de Janeiro," probably because its bark was used 

Irwin et al. 14160 (NY); Rio Sao Bartolomeu, 22 Apr 1980 as a secondary source of quinine. Reported as 

(fl), Heringer et al. 

very 

4448 (NY, US), 12 Jun 1980 (fr), 5082 

ornamental tree with 

(MO, UB, US), 9 Feb 1981 (fl), 6157(MG, NY, US); Escola 

fragrant flowers. 

Agro-T6cnica de Brasilia, Beira do Rio, 23 Jul 1963 (fl-fr), 

Heringer 9156 (VEN); Area de Protegco Ambiental de 

Dubious collection: Brazil. "Prov. S. Pauli," 1868, 

Cafuringa, Fazenda Palestina, 4 Mar 1993 (fl), Pereira & Burchell 3181 (BR, K[3]). These specimens have very 

Mecenas 2442 (US); Area de Protenao Ambiental de large leaves (to 51 cm long and 21 cm wide!) that are 

Cafuringa, 26 Mar 1993 (fl), Pereira & Alvarenga 2503 (SP, obviously pellucid-puncated; stipules exceptionally 

US). GOIAS: Prov. Alexania, Rio Arelia, 9 Aug 1990 (fl), De large, 3.5-4.5 x 5-7 mm, long-lanceolate, entirely 

Paula 3233 (UB); Mun. Caldas Novas, Serra de Caldas [No- glabrous; inflorescences rather stout, woody, 33 cm 

vas], 10 Jul 1976(f), Hatschbach 38796 (US); Goias, 1844 

long, with opposite-decussate branches old fruits of 

(fl), Weddell 2637 (P), 2638 (P); Serra Dourada, ca. 17 km 

very small size (7-8 

S ofGoiis Velho, 6 km NE of Mossamendes, 750 m, 9 May 

1973 (fl), Anderson 9927 (NY, US); Serra Dourada, 27 May 

1965 (fl), Heringer 10468 (F, RB); Contraforte Central, NE 

of Catallo, 875 m, 23 Jan 1970 (fl), Irwin et al. 25206 (F, 

MO, NY, US, VEN). MINAS GERAIS: Mun. Corrego 

Barreiras, rd. BR-365, Pato de Minas, 26 Mar 1980 (fl), 

Hatschbach 42945 (MG, NY, UB, UC); Mun. Araguari, Rod. 

BR-40, Rib. Jordao, 13 Apr 1981 (fl), Hatschbach 43851 

(BR, F, INPA, NY, SP, US); Mun. Paracatu, Fazenda 

Acangau, 17?12'S, 47?06'W, 5 Mar 1989 (fl), Mendonca et 

al. 1292 (UB, US); Teixeira Loares, Mar 1908 (fl), Sampaio 

612 (F); Faz. do Brejao, 31 Jul 1960 (fl), Heringer 7581 

(UB); 1838, Claussen 669 (F, NY); 1839, Claussen s.n. (G); 

1816-1821, Saint-Hilaire 348 (F, P). Rio DE JANEIRO: Rio 

de Janeiro, rd. Vista Chinesa, 500 m to Parque Nacional 

Tijuca, 20 Jun 1985 (fl), Argeli & Oliveira 691 (HRB, US); 

Rio de Janeiro, Vista Chinesa, 1 Mar 1931 (fl), Brade 10639 

(F, GH); Rio de Janeiro, Campo Grande, Serra Mendanha, 

600-700 m, 4 Apr 1978 (fl), Martinelli et al. 4133 (RB); Rio 

de Janeiro, Mt. Corcovado, 1866, Glaziou Ib" (BR, F); Rio 

de Janeiro, 1833, Riedel 1236 (BR, US); Mun. Parati, 

Laranjeiras, trail to Praia do Sono, 100 m, 10 Jun 1991 (fr), 

Giordano et al. 1005 (RB); Mun. Parati, trail to Morro da 

Pedra Rolada, 300 m, 28 Jun 1995 (fr), Campos et al. 48 

(RB); Gavea, E Guanabara, 4 Mar 1947 (fl), Duarte et al. 

s.n. (RB 60854) (NY); Gavea, 4 Mar 1947 (fl), Apparicio 

59161 (F[2]); old rd. down serra to Angra dos Reis, 1 Feb 

1968 (fl), Smith & McWilliams 15364 (US); Rio de Janeiro, 

1833, Vauthier 205 (F, G, K); Rio de Janeiro, Apr 1836 (fl), 

Vauthier 15 (F, G); Rio de Janeiro, 1844 (fl), Widgren (BR, 

F, US). SAo PAULO: CubatAo, 9 Aug 1899 (fl), Edwall 23938 

(F); Cubatao, Serra de Morrao, Rio Quilombo, 5 Jun 1990 

(fr), Kirizawa & Chiea 2350 (SP); Cubatao, 15 Mar 1923 

(fl), Gehrt s.n. (SP); slope of Serra do Mar, estrada velha, 

Cubatlo, 10 Apr 1956 (fl), Kuhlmann 3769 (F, NY, TEX); 

Santos, Feb 1875 (fl), Mosen 3408 (F[3], P); Reserva 

Ecol6gica de Jur6ia, Peruibe, 11 Nov 1982 (fr), Rodriguez 

& Figuereido 15745 (UB). WITHOUr LOCALITY: Brazil, Pohl 

2681 (M); Brazil, 1839 (fl), Pohl s.n. (M); Brazil, Feb-Mar 

1823, Riedel s.n. (M). 

x 4-5 mm), already open. Because 

of its small fruits, large leaves, and collection 

locality, I tentatively attribute these specimens to R. 

formosa, but flowering collections of these individuals 

might prove to be an undescribed taxon. 

Rustia formosa is a beautiful shrub, with white, 

fragrant flowers easily distinguishable from the other 

species of Rustia by their throat and lobes (internally) 

densely pubescent. From R. alba (of the Ecuadorian 

cloud forests) this species differs in the extremely 

reduced calyx and its geographical distribution. Rustia 

simpsonii differs from R. formosa in having much 

larger, thicker corollas, short-glandular hair at the 

internal margins of the corolla lobes (vs. densely pubescent), 

and cupular-membranous calyx with obvious 

lobes (vs. extremely reduced). 

This species was first presented by Chamisso and 

Schlechtendal (1829) as Exostema formosum, with 

two varieties (var. a leprosum and var.? laeve). There 

was no description for E. formosum, but there were 

descriptions for each of the two varieties. Therefore, 

E. formosum is a nomen nudum, and the two varieties 

are invalid because they were attributed to an invalid 

Local names and uses. Brazil: caapeba (Minas 

Gerais, Sampaio 612, Hoene 11865), guapeba branca 

(Campos et al. 48), quina do Rio de Janeiro (Martius 

111), sobrasil (Sao Paulo, Edwall 23938). The label 

species. 

De Candolle (1830) validated Exostemaformosum 

Cham. & Schltdl. ex DC., under which he cited the 

two varieties a leprosum andfi laeve. The two variet- 

ies remained invalid because de Candolle did not cite 

the typical varieties of the species. 

Klotzsch (1846) separated this species from 

Exostema (as Exostemma) to establish the genus 

Rustia, under which he recognized three species. He 

validated R. pohliana by a description, and R. sel- 

lowiana by indirect reference to previously published 

description of E. formosum var. laeprosum; E. form- 

osum had already been validated by de Candolle 

(1830). Schumann (1889), in Flora Brasiliensis, 

treated these three species as synonymous, a conclu- 

sion with which I concur. 

The description of E. formosum var. laeve was 

prepared by Chamisso and Schlechtendal (1829) from


a Sellow collection from tropical Brazil. No Sellow ing-reflexed lobes, 1.8-2.2 cm long, white to creamspecimens 

of this species were encountered. There- white inside and outside, thick and fleshy when fresh; 

fore, I am here selecting a neotype for Rustiaformosa, tube faintly funnelform, 8-10 mm long, ca. 4 mm wide 

the type species of the genus. Lemos 28678 (NY) was 

selected as neotype for R. formosa because it was 

at base and ca. 6 mm wide at the orifice, glabrous 

outside, with a white glabrous basal zone inside, with 

collected from eastern tropical Brazil, and corresponds a ring of ascendent-pilose hairs 6-7 mm from the base, 

most closely to the description ofE.formosum var. laeve. at the same point of the filament attachments at base 

of lobes; lobes 5, 

12. Rustia simpsonii Delprete, sp. nov. Type. Brazil. 

Rio de Janeiro: Petropolis, 21 Feb 1873 (fl), 

Glaziou 6564 (holotype, P; isotypes, BR, K, 

frag-F). Figs. 14G,H, 37A-C 

Arbores R.formosae similes sed ab ea corollis majoribus 

18-22 mm longis, 4-6 mm latis (nec 15-18 mm longis, 2- 

2.5 mm latis), corollae lobis secus medium glabris, intus 

juxta margines pilis glandulosis biseriatis provisi (nec basi 

albo-strigosis intus juxta margines dense papillosis) antheris 

?5 mm latis (nec 1-1.5 mm) basi extensioribus sphaeroidesis 

papillosis auctis (nec nudis) diversa. 

/2-2/3 of corolla length, 12-14 x 4- 

5 mm, narrowly triangular, glabrous outside, glabrous 

in central zone with two rows of minute (40x) glandular 

hairs inside, near the margins. Stamens 5, exserted 

because of spreading lobes, subequal, attached 

ca. 6 mm from the base of the tube; filaments ca. 7 

mm long, with a compact tuft of white ascending hirsute 

hairs at base; anthers cylindrical, narrowly oblong, 

ca. 9 x ca. 5 mm, dorsifixed near the base, base 

rounded, with papillose spheroid extension at base, 

smooth throughout. Pollen exine densely foveolate, 

minutely echinate. Style exserted, 20-23 mm long, 

glabrous; style branches narrowly ovate, ca. 2 mm 

long, stigmatic surface microscopically (40x) papillose. 

Capsule not seen. Seeds not seen. 

Trees; bark grayish. Leafy branchlets glabrous, 

terete; older branches rugose, grayish; lenticels many, 

punctiform to linear, 0.5-3 mm long. Stipules not seen 

(readily caducous), leaving a white linear scar. Leaves 

34-38 x 10-11 cm, L/W 3:1; oblanceolate, cuneate 

(BA = 20-25?) at base, obtuse at apex, sometimes with 

a deltoid acumen ca. 1 cm long; dark green above, 

yellowish green below, semi-coriaceous; drying ol- 

Distribution (see Fig. 39B) and ecology. Known 

only from the surroundings of Rio de Janeiro and 

Petropolis, an environment that is now mostly destroyed. 

This species is known only from two collections 

by Glaziou made in 1867 and 1873, and is possibly 

extinct. For notes on conservation biology of this 

species, see discussion under R. gracilis. 

ive-green, subcoriaceous; glabrous above, glabrous 

below; pellucid punctate; primary and secondary veins 

glabrous, prominent below, secondary veins 15-17 

each side, subparallel; tertiary veins starting sub- 

Specimens examined. BRAZIL. Rio DE JANEIRO: Rio 

de Janeiro [Corcovado and Serra da Estrella (Glaziou, 

1909)], 1867, Glaziou 1098 (BR[2]). 

parallel and reticulate in the center; petioles 41-45 mm 

long, 2.5-3 mm thick, terete or adaxially narrowly 

concave, thickened at base, but not pulvinate; domatia 

On the herbarium sheet Glaziou 6465 preserved 

at Field (isotype fragment-F), Donald Simpson wrote: 

absent. Inflorescences openly paniculate, pyramidal, This collection [was] cited by Schumann in Fl. 

with opposite or alternate, decussate branches; ca. 36 Bras. as R. formosa (C. & S.) Kl. but the flowcm 

long, basal branches to 20 cm long, in ca. 5 pairs, ers are extraordinarily thick. The anthers debasal 

portion of axis not branched ca. 8 cm long; ra- hiscing by 2 pores at the apex confirms its 

chis decussately compressed, narrowly rhombic or placement in Rustia, and the corolla lobes benarrowly 

elliptic in cross section, rachis and branches ing at least as long as the tube and calyx 

glabrous; flowers on tertiary branches alternate or in 

terminal lax cymules; distal bracts 6-10 x 3-5 mm, 

subtruncate places it in or near R.formosa, but 

the flower is nearly twice as thick as in most 

narrowly triangular to deltoid; bracteoles subtending 

flowers 3-5 x 2-4 mm, narrowly deltoid. Flowers 

pedicellate, pedicels 9-12 mm long, glabrous; hypanthium 

obovate to obconical, 4.5-7 x 4-6 mm, glabrous; 

flower buds cylindrical, pointed at apex. Calyx 

cupular, semi-membranous, with barely distinguishable 

lobes, 2-3.5 x 6-8 mm; lobes 5, ca. 1 x ca. 3 mm, 

R. formosa, and the corolla lobes are ca. twice 

as long as the tube (i.e. 2/3 the corolla length) 

while Schumann describe R. formosa as having 

corolla lobes 1/3 to 12 corolla length (i.e. less 

than or equalling the tube). Rustia gracilis has 

corolla lobes 2/3 the corolla length but flowers 

are very slender in that species. 

shallowly deltoid. Corolla short-tubular with spread- 

- D. Simpson, 

July 1977.


., ~ 

~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~W 

,.~~~~~~~~~~~~~~~~~~~4 

.? '~~~~~~~P ~ ~ ~ ~ X 

..~~~~~~~~~~~~~~~~!Mq 

FI.37 uti ipsni(Gaiu 54 B n P yes.A Hbtofiforsecewt mtr la . pnflwr 

shwngodopnanhr adesetd eetiesyl.C.Dtilo nte bs hoigspeoi xenin


Rustia simpsonii is unique in the genus in having 

anthers with papillose spheroid extensions at base 

(Fig. 38C). It differs from the closely related R. 

formosa in having corollas 18-22 x 4-6 mm (vs. 15- 

18 x 2-2.5 mm), corolla lobes glabrous in central zone 

and with two rows of glandular hairs at margins inside 

(vs. white-strigose at base and margins densely 

papillose inside), anthers ca. 5 mm wide (vs. 1-1.5 mm) 

and with papillose spheroid extensions at base (vs. 

without such extensions), and calyx 2-3.5 x 6-8 mm 

(vs. 0.2-0.4 x 0.9-1.2 mm) (Fig. 38A-C). 

Donald Simpson has been an ephemeral, extremely 

accurate botanist, and the author of the only brief study 

specifically dedicated to Rustia. His acute observations, 

penciled on the herbarium specimens (and folders) 

at the Field Museum (F), have been very helpful 

in my revision of this genus, and I therefore honor him 

with the present appellation. 

13. Rustia angustifolia K. Schumann in Martius, Fl. 

Bras. 6(6): 262. 1889. Type. Brazil, Rio de Janeiro, 

Serra da Estrella, 1885 (fl), Glaziou 14937 (B*; 

photo-B at F, GH, MO, NY, VEN; lectotype, G, 

here selected; isolectotypes, BR, K, MA, R). The 

specimen Glaziou 14937 preserved at R has the 

date (manu Glaziou) of 25 August 1883, but on 

other duplicates the collecting date is solely 1885 

(manu Glaziou). Figs. 6B, 14F, 38B, 39A-E 

Shrubs 4-6 m tall, with slender branches usually 

terete; bark grayish-whitish. Leafy branchlets 

golden-tomentulose; older branches glabrate; lenticels 

absent. Stipules narrowly triangular but not acuminate, 

tomentulose outside, glabrous, with short 

colleters inside, 18-22 x 1.3-1.5 mm, light green, 

readily caducous, leaving a linear scar. Leaves (10-) 

11.5-33(-36.5) x section, rachis and branches minutely puberulent; 

flowers alternate on distal branches or in cymules; 

distal bracts 3-7 x 3-4 mm, narrowly lanceolate; 

bracteoles subtending flowers 1-2 x 0.5-0.7 mm, 

narrowly lanceolate, tomentulous outside, glabrous 

inside. Flowers pedicellate, pedicels 8-20 mm long, 

glabrous to minutely puberulent; hypanthium narrowly 

obconical, 2.5-5 x 2-3.5 mm, glabrous; flower 

buds cylindrical, acute at apex. Calyx small-cupular, 

truncate or with barely distinguishable lobes, 3-4 mm 

wide; lobes 5, 0.5-0.9 x 0.5-1 mm, shallowly triangular 

to deltoid, ciliolate. Corolla tubular with spreading 

lobes, 1.4-1.8 cm long, pinkish red to flesh-red 

outside, pearl-white inside, thick and fleshy when 

fresh; tube cylindrical 1.1-1.4 x 3-4 mm, glabrous 

outside, glabrous inside, with a tuft of sparse hairs at 

the sides of each filament attachment, 5-6 from the 

base (microscopically papillose); lobes 5, 1/5-l/4 of 

corolla length, 3-4 x 1-1.2 mm, ovate-triangular, glabrous, 

with the margins minutely papillose inside. 

Stamens 5, exserted because of spreading lobes, subequal, 

attached 2.5-5.5 mm from the base of the tube; 

filaments 1.8-2 mm long, barbate at base, with white 

to golden-yellow ascending hairs 0.1-0.2 mm long; 

anthers cylindrical, narrowly oblong, 5-8 x 0.6-0.9 mm, 

dorsifixed near the base, base rounded to subsagittate, 

microscopically (40x) papillose at very base. Pollen 

exine foveolate (non-echinate). Style exserted, 15-23 

mm long, glabrous; style branches narrowly triangular, 

0.9-1.1 mm long, glabrous, stigmatic surface 

smooth. Capsules obovoid to oblong-obovoid, apex 

subtruncate to shallowly hemispherical, 10-16 x 7- 

10 mm, sometimes with sparse punctiform lenticels, 

dark brown, glabrous throughout. Seeds 1.33-1.5 x 

0.4-0.73 mm. 

Distribution (Fig. 38B) and ecology. Understory 

shrubs in primary and secondary forests in the moun- 

2.5-4(-5.5) cm, L/W 3:1 to 5:1, tains surrounding Rio de Janeiro, Brazil. Flowering 

narrowly lanceolate, acute at base and at apex, some- specimens were collected from March to May. For 

times tapering to an acuminate apex to 1-1.5 cm long, conservation biology of this species, see discussion 

with a round tip; dark green above, pale green below, under R. gracilis (below). 

semi-coriaceous; drying olive-green, chartaceous; 

glabrous above and below, pellucid punctate; primary Specimens examined. BRAZIL. Rio DE JANEIRO: Rio 

and secondary veins glabrous, secondary veins (8-) de Janeiro, Serra da Estrella, old rd. Rio de Janeiro- 

10-17 each side, tertiary veins openly reticulate; pet- Petropolis, 23 May 1988 (fr), Gomes et al. 323 (HRB, 

ioles (10-)15-35 mm long, 1-2.5 mm thick, adaxial- RB[2]); Rio de Janeiro [Corcovado and Serra da Estrella 

ly concave to flattened, pulvinus present in young (Glaziou, 1909)], Mar 1872 (fl), Glaziou 2607 (K); Rio de 

leaves, 4-6 mm long, 1.5-2.5 mm Janeiro, 1891, Glaziou 18906 (K); Rio de Janeiro, Serra da 

thick, rarely absent; 

Estrella 

domatia absent. Inflorescences 

[near Mandioca (Glaziou, 1909)], 2 May 1880 (fl), 

pendulous, long-pe- Glaziou 12038 (F, G, P); in primary forest, Macah6, Maydunculate, 

paniculate with opposite decussate 

Jun 1832 (fl), Riedel & Luschnath 1062 (F, NY); Serra do 

branches; 11-20 x 8.5-16 cm, lateral branches 3-5 

Orgios, Alto Macah6, 1891, Glaziou 18306 (G); Porto Estrepairs, 

basal portion of axis not branched 6-8.5 cm lla, Dec 1823 (st), Riedel 66 (BR); Riedel 68 (BR); vic. of Manlong; 

rachis compressed, narrowly rhombic in cross- diocca, Martius 2964 (M), 2965 (M), 2966 (M), 2967 (M).


Ao 

FIG. 38. Distribution of four species of Rustia. A. R. formosa. B. R. simpsonii (stars), R. gracilis (diamonds) and R. 

angustifolia (circles). 

This species is unique in the genus in having long 

pedunculate, pendulous inflorescences; terminal fas- 

cicles of leaves at the tip of the vegetative branchlets; 

leaves narrowly lanceolate, three to five times as long 

as wide; corollas flesh-red outside and pearl-white 

inside, and corolla lobes one-fifth of the corolla length. 

Until few years ago, the last collection of R. 

angustifolia was dated 1891, and because of the in- 

creasing destruction of the natural environment in the 

surroundings of Rio de Janeiro (Dean, 1995), this 

species had been suspected to be extinct. Neverthe- 

less, the recent collection (1988) of Gomes et al. 323 

at Serra da Estrella, has proven the ongoing existence 

of this extremely rare and endangered species. 

14. Rustia gracilis K. Schumann in Martius, Fl. Bras. 

6(6): 263. 1889. Type. Brazil. Rio de Janeiro: 

Riverbanks of Rio Mandioca, Dec 1822-Jan 1823 

(fl), Riedel 424 (B*; lectotype, M, here selected; 

isolectotypes, BR[2], F, GB, GH, K, US). 

Figs. 5D, 14C,D, 38B, 40A-E 

3, 0 

Shrub 4-6 m tall, with slender branches; bark 

grayish-whitish. Leafy branchlets glabrous, slender, 

terete to tetragonous, grayish; lenticels absent. 

Stipules not seen (readily caducous), leaving a white 

linear scar. Leaves (11-)14-18 x 3.5-5.5 cm, L/W 

3:1 to 3.6:1, lanceolate to oblanceolate, acute (BA = 

18-25?) at base, acute at apex, tapering to an acuminate 

apex to 1-1.5 cm long, with a round tip; dark 

green above, pale green below, membranaceous; drying 

olive-green, chartaceous; glabrous above, glabrescent 

to sparsely pubescent below, pellucid punctate; 

primary and secondary veins glabrescent to sparsely 

pubescent, secondary veins 10-13 each side, tertiary 

veins openly reticulate; petioles 13-20(-25) mm long, 

1-1.5(-2) mm thick, subterete; pulvinus absent; 

domatia absent or a tuft of sparse hairs 0.5 mm wide, 

hairs white 0.5-0.6 mm long. Inflorescences openly 

paniculate with opposite decussate branches; 12-16 

x 11-13(-20) cm, lateral branches 3-5 pairs, basal 

portion of axis not branched 3.5-6.5 cm long; rachis 

compressed, elliptic or narrowly rhombic (rarely terete) 

in cross section, rachis and branches glabrous to 

minutely puberulent, extremely thin and delicate (in


FIG 3. Rsta ngutioli 143, ( fom laio B ,ioettp;BDfo id102 Y rmRee6) 

E.~~~~~~~~~r M"ba capsur ture l e.I 

FIG. 39. Rustia angustifolia (A from Glaziou 14937, BR, isolectotype; B-D from Riedel 1062, NY; E from Riedel 68). 

A. Habit of inflorescence with young and mature leaves. B. Flower bud. C. Stamen. D. Detail of anthers base, dorsal view. 

E. Mature capsule.


C cm 

[~~~~~~~~~~~~~~~~~~~~~~~~~5 

FIG. 40. Rustia gracilis (A-D from Riedel 424, BR; E from Glaziou 14893, BR). A. Habit of inflorescence with mature 

leaves. B. Abaxial view of blade showing pellucid punctation. C. Flower bud. D. Style. E. Mature capsule. E. Stamen, 

showing pilose base and open anther. 

w '. 

: .....:.: .......... :~ : : - . . . . . 5 : 

FI.40uti r rm cils(- R e?44 R rmGaiu183 R.A ai fnlrscc ihmtr 

showing pilos? bag~~~~~~ andopnanter


comparison to all other species); flowers alternate on 

distal branches or in cymules; distal bracts 2-3 x 1- 

1.5 mm, narrowly triangular; bracteoles subtending 

flowers 1-1.5 x 0.5-0.7 mm, narrowly triangular. 

Flowers pedicellate, pedicels 5-10 mm long, very thin 

and delicate, glabrous; hypanthium narrowly 

obconical, 2-2.5 x 1-1.5 mm, glabrous; flower buds 

clavate. Calyx narrowly cupular, truncate or with 

barely distinguishable lobes, 1.5-2.5 mm wide; lobes 

deltoid, 0.8-1 x 0.8-0.9 mm, ciliolate. Corolla tubular 

with narrowly spreading lobes, 1.2-1.8 cm long, 

white with pink-reddish base outside, white inside, 

thin and delicate when fresh; tube narrowly funnelform, 

5-7 mm long, ca. 1 mm wide at base and 2-2.5 

mm wide at the orifice, glabrous outside, glabrous and 

microscopically (40x) papillose inside, with a tuft of 

white-pilose hairs (0.8-1 mm long) at the sides of each 

filament attachment and base of lobes; lobes 5, 1'3-/2 

of corolla length, 5-7 x 0.9-1.1 mm, narrowly triangular, 

glabrous outside, with few sparse hairs and margins 

minutely papillose inside. Stamens 5, exserted 

because of spreading lobes, subequal, attached 5-6 

mm from the base of the tube; filaments 3-3.5 mm 

long, very thin and pendulous, barbate at base, with a 

tuft of white ascending hairs (0.8-1 mm long); anthers 

cylindrical, narrowly oblong, (2-)3-3.5 x 0.5-0.6 cially the threatened parks of Serra do Orgaos and 

Serra de Estrella. 

Specimens examined. BRAZIL. Rio DE JANEIRO: Rio 

de Janeiro, Serra de Estrella, Mandioca, 29 Mar 1884 (fl), 

Glaziou 14893 (BR, F, G, P); riverside close to Mandioca, 

Feb-Mar 1823 (fr), Riedel 367 (BR); E of Rio de Janeiro, 

Serra do Orgaos, Caneca, Dec 1974 (fl), Occhioni 6655 (US); 

Rio de Janeiro [Serra dos Orgaos, Alto Macahe (Glaziou, 

1909)], 1881, Glaziou 12036 (B*, BR, F, G[2], R); Rio de 

Janeiro, Glaziou 19437 (BR). 

Rustia gracilis is unique in the genus in having 

inflorescences with extremely thin and delicate 

branches (hence the specific epithet), and pedicels and 

corollas long and slender, which gives an overall frail 

appearance. This species could be confused with R. 

angustifolia, which is also an endemic of the mountain 

slopes surrounding Rio de Janeiro, but it differs 

from R. angustifolia in its openly paniculate inflorescences, 

slender whitish corollas (1-2 mm wide), and 

its membranous oblanceolate leaves. On the other 

hand, the inflorescences of R. angustifolia are rather 

stout, long-pedunculate, and densely corymbose distally; 

having thicker blood-red to flesh-red corollas 

(3-4 mm wide), and semi-coriaceous narrowly lanmm, 

ceolate leaves. 

dorsifixed at 0.2-0.3 mm from the base, base rounded, 

the base of two thecae often asymmetrical, microscopically 

(40x) papillose-echinate. Pollen exine 

Doubtful Species 

densely foveolate (non-echinate). Style exserted, 15- 

Rustia haitiensis Urban, Ark. Bot. 24A 

16(-18) mm long, glabrous, minutely papillose at dis- 

(4): 36. 1931. 

tal zone; style branches ovate, 0.4-0.5 mm 

Type. Haiti. Massif de la Hotte, western group, 

long, gla- 

Jeremie, left arm of 

brous, stigmatic surface glabrous; ovaries often 

Voldrogue River, 400 m, very 

rare, 22 Jul 1928 

parasitized by insects, causing an abnormal 

(fl-fr), Ekman H-10408 (B*; lecenlargement 

even before anthesis, becoming spherical and to 

totype, S, here selected). 

5-6 mm diam. but with no seeds inside. Capsules 

obovoid to oblong-obovoid, apex subtruncate, 6-8 x Shrub or small tree, glabrous. Leafy branchlets 

4-5 mm, dark brown, glabrous throughout. Seeds biconvex; older branches subterete; lenticels sparse. 

0.87-1.16 x 0.43-0.67 mm. 

Stipules narrowly triangular, acuminate, 6-8 mm 

long, readily caducous. Leaves petiolate, petioles 10- 

Distribution (Fig. 39B) and ecology. All the 20 mm long, 1.5-2 mm thick, adaxially concave to 

specimens examined have been collected either in flattened; blades 17-21 x 5-8 cm, L/W ca. 3:1; obo- 

Serra do Orgaos or Serra de Estrella, in the vicinity vate-elliptic, acute (BA 

of Rio de Janeiro. Flowering collections have been 

collected from December through March. Fruiting 

specimens were collected in February and March. 

Conservation status. This species is also extremely 

rare; most specimens were collected before 

1900, with a single collection from 1974 [Occhioni 

6655]. Rustia gracilis (with R. simpsonii and R. 

angustifolia) emphasize the extreme importance in 

protecting the remnants of the few patches left of 

Brazilian Atlantic forest (cf. Dean, 1995), and espe- 

= 27-32?) at base, widely 

obtuse to rounded at apex, chartaceous; glabrous 

above and below; pellucid punctate; primary and secondary 

veins glabrous, slightly depressed above, 

prominent below; secondary veins 11-14 each side; 

tertiary veins starting subparallel and reticulate in the 

center; domatia absent. Flowers, capsules, seeds, and 

pollen not seen. 

From Urban's (1931) original description: Inflorescences 

terminal, paniculate, with opposite to 

subopposite decussate lateral branches; ca. 25 cm 

long; rachis and branches glabrous, with lenticels;


distal bracts 2-3 mm long, lanceolate and acuminate, 

bracteoles subtending flowers ca. 1 mm long, deltoid, 

glabrous. Flowers pedicellate, pedicels ca. 2 mm long 

(in fruits). Capsule globose, to 3.5 mm long. 

14 Oct 1863 (fl), Isern 2121 (holotype, F) = 

Alibertia isernii (Standley) D. R. Simpson. 

Simpson (1976) transferred this species to 

Alibertia, a conclusion with which I concur. 

Distribution and ecology. Known only from the 

type, from the now-destroyed cloud forest on Massif Rustia longifolia Standley, Publ. Field Columbian 

de La Hotte in southern Haiti. From the description, Mus. Nat. Hist., Bot. Ser. 8: 159. 1930. Type. 

the specimens were collected in flowering stage in Colombia. Caqueti: Between Santa Marta and Mar- 

July, with some old capsules persisting on branches. sella, Aug 1926 (fr), Woronow & Jusepczuk 6426 

In the protologue Urban (1931) stated: "Rarissima (holotype, LE-n.v.; isotypes, F, LE-n.v.). =Dolichoet 

parca, Jul cum inflor. valde juvenilibus et capsulis delphys chlorocrater K. Schumann & K. Krause. 

vetustis. Obs. An re vera hujus generis, antherae adhuc 

ignotae demonstrabunt. Habitus et folia glandulosopunctata 

ad genus Rustiam spectant" [... Even though Rustia rosea (Klotzsch & Karsten ex Walpers) K. 

it was not possible to see the anthers, habit and pellu- Schumann in Engler & Prantl, Nat. Pflanzenfam. 

cid-punctate leaves show that it belongs to Rustia]. 4(4): 18. 1891. Basionym: Henlea rosea Klotzsch 

Only one sterile specimen (Ekman H-10408, S) of & Karsten ex Walpers, nomen., Ann. Bot. 2: 788. 

this species was found. The above description is based 1852. Type. Pro. syn. = Ladenbergia muzonensis 

on personal observations, to which I added the transla- (Goudot) Standley (1930a). 

tion of Urban's (1931) original description. The leaves 

Schumann (1891) erroneously treated H. rosea as 

pellucid punctation is not as obvious as in all the spe- a valid species and as the type species of Henlea. As 

cies of this genus, suggesting that this species prob- noted by Standley (1930a), H. rosea was never valably 

does not belong to Rustia. Until further collecidly 

described. Walpers (1852) treated this taxon as 

tions of flowering and fruiting material can be made the 

valid and listed it as synonymous with Cascarilla 

identity of this taxon is destined to remain doubtful. 

muzonensis [= Ladenbergia muzonensis], an interpretation 

with which I agree. 

Excluded Species 

Henlea muzonensis (Goudot) Klotzsch & H. Karsten 

ex Walpers, Ann. Bot. Syst. 2: 788. 1852. Cinchona 

muzonensis Goudot, Phil. Mag. New Ser. 

3: 132. 1828. Cascarilla muzonensis (Goudot) 

Weddell, Ann. Sci. Nat. 3(10): 11. 1848. Cascarilla 

[sect.] muzonia Weddell, Hist. Nat. Quinquinas 

90. 1849. Buena muzonensis (Goudot) Weddell, 

J. Linn. Soc. 11: 187. 1869. Type. Colombia. 

Santander: Near Muzo, Goudot s.n. (lectotype, P, 

selected by Andersson, 1994). = Ladenbergia 

muzonensis (Goudot) Standley (1930a). 

Rustiaferruginea Standley, Contr. U.S. Natl. Herb. 

18: 142. 1916. Type. Panama. Col6n: Along Rio 

Fat6, in forest or thickets, 10-100 m, Aug 1911 

(fl), Pittier 4201 (holotype, US). = Rustia pauciflora K. Schumann ex Solereder, nomen., 

Ber. Deutsch. Bot. Ges. 8: (99). 1890. Rustiapauciflora 

K. Schumann, in herb. Type. Tobago. Forest 

of Cremorue River, 360 m, Nov 1889 (fr), Eggers 

5812 (holotype, M; isotypes, K, US). 

Bathysa (?). 

This species does not belong to Rustia, but because 

of the depauperate conditions of the unicate specimen, 

it is impossible to attribute this species to a genus with 

certainty. 

Rustia isernii Standley, Publ. Field Mus. Nat. Hist. 

Bot. Ser. 22: 212. 1940. Type. Peru. Chachamayo, 

= Tresanthera 

condamineoides Karsten var. condamineoides. 

For further details on the taxonomic history and 

identity of this taxon see discussion under Tresanthera 

condamineoides Karsten var. condamineoides. 

TRESANTHERA 

2. Tresanthera H. Karsten, Fl. Columb. 1: 37. pl 19. 

1858; Hooker in Bentham & Hooker, Gen. P1. 2(1): 

44. 1873; Baillon, Hist. P1. 7: 471. 1880; Karsten, 

Bot. Jahrb. Syst. 8: 356. 1887; Schumann in Engler 

& Prantl, Nat. Pflanzenfam. 4(4): 19. 1891; 

Solereder, Ber. Deutsch. Bot. Ges. 8: 99. 1890; 

Solereder, Bull. Herbier Boissier 1: 280-181. 

1893; Standley, Publ. Field Columbian Mus., Bot. 

Ser. 7: 371. 1931; Steyermark, Bol. Soc. Venez. 

Cienc. Nat. 25: 248. 1964; Steyermark in Lasser & 

Steyermark, Fl. Venezuela 9(1): 256-261. 1974. Type 

species. Tresanthera condamineoides Karsten.


Shrubs to single-stemmed trees, terminal large pyramidal shrubs (rarely medium-sized trees), 

branches with soft pith in the center, with leaves clus- and has never been found to develop buttresses. Its 

tered at apex of each branch; bark grayish-pale brown. wood is usually soft and of no particular use, its bark 

Stipules interpetiolar, free at base, overlapping in bud, is smooth and grayish, and its seeds are minute, horinarrowly 

triangular, readily caducous. Leaves peti- zontally inserted, non-winged, flattened, and apically 

olate, elliptic to oblanceolate, obtuse or short-acumi- truncate (as in some species of Rustia). 

nate at apex; pellucid-punctate; petioles thickened at Tresanthera is sometimes confused with Rustia, 

base, not pulvinate, becoming corky in older leaves; but the latter has terminal poricidal anthers, anther 

domatia absent. Inflorescences terminal, solitary, connective tissue not as evident, smaller fruits, and 

paniculate or single thyrsoid axis; lateral branches variously shaped seeds. 

(when present) cymose or thyrsoid. Flowers 

protandrous; hypanthium obconical, acute at base. 


Calyx reduced to a wavy margin or with minute lobes, 

persistent. Corolla campanulate with patent to re- Tresanthera was founded by Karsten (1858), deflexed 

lobes, fleshy when fresh, yellowish orange to riving the name from the Greek tresis (perforation) 

orange-red outside, yellowish white to cream-white and antheros (anther) to refer to its porate anthers. 

inside; tube basally ventricose with medial constric- Karsten produced a very detailed description of the 

tion, glabrous; lobes 5; aestivation valvate-reduplicate genus and of T. condamineoides, the type species. This 

with contact zone. Stamens 5, alternate to corolla genus was included (with Henlea) by Hooker (1873) 

lobes, half-exserted, attached near the base of corolla in Rustia. Nevertheless, Karsten (1887) maintained 

tube; anthers convex toward the center, acute at apex, them as separate and published a brief description on 

yellow, dorsifixed near the base, dehiscing by a com- their basic morphological differences. After a series of 

mon lateral triangular pore below the apex. Pollen unpublished and invalid names, Solereder (1893) 

tricolporate, exine reticulate. Style exserted; style legitimized the taxon T. pauciflora K. Schum. ex Soler. 

branches membranous, ovate, reflexed at maturity. Steyermark (1964a) published Tresanthera 

Ovary 2-celled, obconical, glabrous, placentation thyrsiflora, basing its description only on portions of 

axile; ovules many in each locule, horizontally in- an inflorescence. Later, Steyermark (1974) stated that 

serted; immature fruits oblong-elliptic, reddish green the three species recognized by him could as well be 

and semi-carose when fresh. Capsules woody, nar- conspecific, suggesting that subsequent collections 

rowly obovoid; disk obviously exceeding the calyx; might better clarify their identity. In the present treatdehiscing 

loculicidally, disk septicidal dehiscence ment Tresanthera is treated as monotypic, with two 

rarely present in old capsules. Seeds compressed, infraspecific categories, and ranging from the coastal 

horizontal, scobiform, with reticulate exotesta. forests of Venezuela to the island of Tobago. 

Tresanthera is unique in the Rubiaceae in having 

poricidal anthers opening by a common lateral 

1. Tresanthera condamineoides H. Karsten, Fl. 

pore 

of triangular shape, just below the apex. The common 

Columb. 1: 37, pl. 19. 1869. 

lateral pore is the result of the two convex thecae that 

Figs. 3E, 4D, 7A-C, 15A,B, 42, 43, 44 

converge apically so that the anther becomes pointed. Shrubs to 

The connective tissue in the dorsal portion of the antrees, 

to 15-20 m tall, to 30 cm dbh, 

thers (and between the two thecae) is 

usually single-stemmed frondose trees; bark grayish. 

quite evident, 

black, and with irregular margins. The closely related 

Leafy branchlets glabrous, with soft pith in the cen- 

Rustia also has poricidal anthers, but 

ter, 

these 

easy to break, terete to subquadrangular, grassopen 

by 

two terminal pores (the thecae not converging). Both 

green; older branches terete, pale brown. Stipules 

Tresanthera and Rustia are exceptional in the Rubinarrowly 

triangular, acuminate, 3-7 cm long, ciliolate, 

aceae in having leaves with 

glabrous to sparsely puberulent outside, glabrous, with 

"pellucid glands." a basal area of colleters inside 

Tresanthera has protandrous (the stigma is 

(Fig. 3E), dark green, 

receptive 

only after the anthers are dried) flowers, with 

readily caducous, leaving a scar encircling the stem 

1.5-2.5 mm wide. Leaves 28-71 x 15-25 cm, L/W 

valvate-reduplicate aestivation, erect or nodding, and 

do not produce any particular fragrance. Because of 

1.5:1-3:1; elliptic, oblong, to oblanceolate; acute, 

obtuse to truncate at 

their particular morphology, the flowers are most 

base, acute to obtuse at apex, 

sometimes short deltoid-acuminate; blade dark 

probably buzz-pollinated by bees (as in some 

green 

species 

of Rustia), but their floral visitors/pollinators have 

above, pale green below, semi-coriaceous; drying olnever 

been observed. Tresanthera usually occurs as 

ive-green, stiff-chartaceous; glabrous above and be-


low, pellucid-punctate; primary and secondary veins 


each side; tertiary veins starting subparallel and openly 

reticulate in the center, faintly evident above, very 

evident below; petioles 3-7(-10) cm long, 3-7 mm 

thick, terete to adaxially concave, glabrous. Inflorescence 

single pauciflorous axes to thyrsoid-panicles, 

to 100 cm long; lateral branches (when present) 1-2 

pairs, opposite to subopposite, (5-)15-28 cm long; 

basal portion of axis not branched 14-33 cm long; 

rachis terete to obtuse-compressed, rachis and 

branches glabrous; flowers on rachis and lateral 

branches solitary or in several cymules subtended by 

a bracteole; bracts subtending basal branches leaf-like, 

semi-caducous, to 17 x 7 cm, lanceolate; distal bracts 

subtending the cymules and the flowers narrowly triangular, 

4-15 x 3-7 mm. Flowers long-pedicellate, 

pedicels 10-25 mm long, glabrous; hypanthium 

obconical, 5-10 x ca. 3 mm, glabrous to puberulent. 

Calyx reduced to a minutely undulate (crenulate) 

margin, 0.5-1.5 x 4-6 mm, glabrous. Corolla campanulate 

with reflexed lobes, 20-30 mm long, orangered 

outside, yellowish white to greenish white inside, 

semi-fleshy to subcoriaceous when fresh; tube with 

medial constriction, 1.1-1.8 x 5-9 mm, glabrous outside 

and inside, without internal ring of hairs; lobes 

5, '/3-/2 of corolla length, 7-12 x 6-8 mm, ovate to 

deltoid, acute at apex, glabrous outside, microscopically 

(40x) papillose inside. Stamens 5, partially exserted, 

equal, attached 5-7 mm from the base of the 

tube; filaments 4-6 mm long, enlarged and flattened 

at base, glabrous; anthers convex toward the center, 

lanceolate, acute at apex, (8-)10-13 x their general shape, but differ from the latter two in 

having anthers strongly convex toward the center. 

Also, the anthers in Rustia are apically poricidal, while 

in Condaminea they dehisce by lateral slits. 

The detailed original description by Karsten (1858) 

and its hand-colored illustration provide sufficient 

information to grasp the identity of T. condamineoides. 

In the Vienna Herbarium (W) two Karsten 

specimens of Tresanthera are preserved: one with a 

flowering inflorescence and a cymule with old capsules, 

bearing the label "Tresanthera condamineoides 

Krst. - Cumbre de Valencia - Pt Cabello - Dr 

Karsten" (unknown handwriting); the other with a 

terminal branch with two fully expanded leaves, and 

two cymules with old capsules, bearing the label 

"Tresanthera condamineoides Krst - Cumbre de 

Valencia - Pt Cabello" (unknown handwriting), with 

"H. Karsten. Dr." embossed on the label, and the annotation 

"! Sch." (in K. Schumann handwriting). I 

have selected the former specimen (having flowers in 

anthesis and mature capsules) as the lectotype of T. 

condamineoides, and the latter as isolectotype. Additional 

isolectotypes might also be found at LE (Tryon, 

1963), but I did not examine material from that institution. 

A dubious fragment has been found at Field 

Museum (F), with a few anthers and a small piece of 

leaf; on the envelope is written "Tresanthera Condaminensis 

[orth. var. Karst. - Karsten." No original labels 

are attached to the herbarium sheet, and the identity 

of the fragments preserved at F remain doubtful. 

The label of Eggers 5812 (holotype of T. pauciflora) 

preserved at Munich (M) bears the following 

1.3-2.8 mm, annotation in Schumann's handwriting: "Certe non! 

base rounded and bilobed, smooth throughout, dehisc- potius Rustia spec. nov., Rustia pauciflora m." 

ing by common lateral triangular pore, 2.5-4 mm Solereder (1890) simply cited the name Rustiapaucibelow 

the apex. Pollen exine foveolate (non-echinate). flora from Schumann's herbarium annotation, but 

Style exserted, terete, 20-28 mm long, glabrous; style without description, thus producing an invalid specific 

branches elliptic to narrowly ovate, 2.5-3 mm long, epithet. Schumann (1891), in Die Natirlichen Pflanzenstigmatic 

surface smooth; immature fruits elliptic, familien, simply listed "T. pauciflora (Solered.) K. 

reddish green, and semi-camose when fresh. Capsules Schum.," also without description, causing this epinarrowly 

elliptic to obovoid, 15-24 x 8-12 mm, dark thet to remain a nomen nudum. Finally, Solereder 

brown, sometimes with few lenticels, glabrous; disk (1893) published a detailed description of T. pauciobviously 

exceeding the calyx, black; disk septicidal flora, validating Schumann's specific epithet and statdehiscence 

present in some old capsules. Seeds 1-1.5 x ing that the single specimen to which he referred was 

0.33-0.53 mm. 

"Eggers 5812 (Herb. Monac. [M!])", where the holotype 

is still preserved. 

Tresanthera condamineoides is a rare species in- The main characters that Steyermark (1974) used 

creasingly threatened with extinction by human pres- to distinguish the three Tresanthera species recogsure. 

It is easily distinguishable within the family by nized by him were corolla color, inflorescence morits 

laterally poricidal anthers and pellucid-punctate phology, and leaf shape. Examining complete sets of 

leaves (with Rustia). The flowers of T. condamine- various collections, I noticed that the corolla color of 

oides vaguely resemble those of Rustia occidentalis Tresanthera ranges from yellowish orange to orangeand 

Condaminea corymbosa in being reddish outside red outside, and from yellowish white to greenish 

and greenish white inside, fleshy when fresh, and in white (to pearl-white) inside (as in Condaminea


I _ n .* 

s 

f 

4 

*? 

^p , a 

II ?11 

FIG. 41. Distribution of Tresanthera condamineoides var. condamineoides (circles) and T. condamineoides var. 

thyrsiflora (star). 

corymbosa), making this character taxonomically insignificant. 

The inflorescence morphology of Tresanthera 

varies from pauciflorous to sparsely cymose within 

the same set of collections (probably from single individuals), 

and is densely thyrsoid only in Bernardi 

5829. In Rustia, the closest genus to Tresanthera, I 

have observed subsecund inflorescences and erect 

inflorescences in the same individual. I have not observed 

individuals of Tresanthera in the field, but it 

is expected that the same is true for this genus, and 

therefore the secundiflorous or erect inflorescences are 

not considered a reliable taxonomic character. 

The leaves of Tresanthera are usually oblanceolate 

(but elliptic-oblong in Bernardi 5829), and the leaf 

bases vary from acute to obtuse (but are truncate in 

Bernardi 5829). The fruits of Tresanthera change in 

shape as they go through maturation, early on elliptic-ovoid 

and becoming narrowly obovoid with age. 

After proposing the three Tresanthera species, 

Steyermark (1974) stated that they might be reduced 

to one or two species as further collections accumulated. 

Steyermark did not study the types housed in 

the European herbaria, which are crucial for understanding 

the identity of these taxa. Additional recent 

collections (and their accurate label data) throughout 

its range have helped me to unravel the confused taxonomy 

of Tresanthera. I conclude that the characters 

used by Steyermark (1964, 1974) to separate the three 

taxa are of little diagnostic merit, and this genus is 

here treated as monotypic: T. thyrsiflora is reduced 

to varietal rank, and T. pauciflora is placed in synonymy 

with var. condamineoides. 

Key to the varieties of 

Tresanthera condamineoides 

1. Leaf blades oblanceolate to obovate, acute at 

base, L/W ca. 3:1; inflorescence a sparsely 

flowered thyrsoid panicle, with a single axis 

or with a pair of basal branches; corolla campanulate 

with reflexed lobes, ventricose at base, 

9-10 mm wide at base ......... la. var. condamineoides 

1. Leaf blades elliptic to oblong, truncate at base, 

L/W 1.5:1 to 2.5:1; inflorescence a densely 

flowered thyrsoid panicle, with a pair of long 

basal branches; corolla subcampanulate with 

patent lobes, conical at base, 4-5 mm wide at 

base ................................................. b. var. thyrsiflora 

la. Tresanthera condamineoides H. Karsten var. 

condamineoides. Tresanthera condamineoides H. 

Karsten, Fl. Columb. 1: 37, pl. 19. 1869. Type. 

Venezuela. Aragua: Puerto Cabello, Cumbre de 

Valencia (near the village of Campanero), hot wet 

forest on the slopes of coastal Cordillera, 100-200 

m, 1844-1847[?] Karsten s.n. (B*; lectotype, W, 

here selected; isolectotypes, W, LE?; dubious 

isolectotype, fragment at F). 

Figs. 3E, 4D, 7C, 15B, 41, 42A-F 

Tresanthera pauciflora K. Schumann ex Solereder, Bull. 

Herb. Boissier 1: 280. 1893. Tresanthera pauciflora 

K. Schumann, nom. nud., in Engler & Prantl, Nat. 

Pflanzenfam. 4(4): 19. 1891. Rustia pauciflora K. 

Schumann ex Solereder, nom. nud., Ber. Deutsch. 

Bot. Ges. 8: 99. 1890. Type. Tobago. Morue d'Or, 

forest of Cremorue River, 360 m, Nov 1889 (fl), 

Eggers 5812 (holotype, M; isotypes, GB, K, US).


FIG. 42. Tresanthera condamineoides var. condamineoides (A from Eggers 5812, US, isotype; B from Steyermark et 

121534, NY; C-E from Sandwith 1757, K; F from Fendler 2366, K). A, B. Inflorescence variation. A. Pauciflorous inflo- 

rescence with immature fruits, and mature leaf. B. Multiflorous inflorescence with mature fruits. C. Open flower at anthe- 

sis with dehished anthers, before style elongation. D. Stamen, adaxial view. E. Stamen, lateral view. F. Mature capsule. 

A


Leaf blades oblanceolate to obovate, acute at base, 

L/W ca. 3:1. Inflorescences single pauciflorous axes 

to sparsely flowered thyrsoid panicles with a pair of 

basal branches; bracts subtending basal branches nar- 

rowly triangular. Corollas campanulate with reflexed 

lobes, 25-30 mm long; tube ventricose at base, 13- 

18 x 9-10 mm (at base); lobes 13-/2 of corolla length, 

7-10 mm long. Stamens with filaments 5-6 mm long; 

anthers lanceolate, acute at apex, 11.5-12.5 x 2.2-2.5 

mm. Pollen exine regularly foveolate. Style 20-22 

mm long; immature fruits ovoid-elliptic. Capsules 

narrowly obovoid, 18-24 x 8-12 mm. Seeds 1.33- 

1.5 x 0.37-0.53 mm. 

Distribution (Fig. 41) and ecology. Lowland rain 

forests and cloud forests of medium elevations, in 

association with Gyranthera caribensis and Iriartea 

fusca, 100-1400 m, along the Coastal Cordillera of 

Venezuela and the forested slopes of Tobago. Flow- 

ering specimens were collected in August, Novem- 

ber, and December. Specimens with mature capsules 

have been collected in July and December. 

pauciflorous to multiflorous panicles, with lateral 

cymules ranging from single-flowered to few flowered; 

all the possible intergradations were found 

throughout the specimens examined. Unfortunately, 

in Bernardi 5829 (var. thyrsiflora) only young capsules 

are available, so to it was impossible to reliably 

compare the fruits of these two varieties. 

lb. Tresanthera condamineoides H. Karsten var. 

thyrsiflora (Steyermark) Delprete, var. et stat. 

nov. Tresanthera thyrsiflora Steyermark, Bol. Soc. 

Venez. Cienc. Nat. 25: 248. 1964. Type. Venezuela. 

Miranda: Merida, rain forest of Guatopo, 

400-600 m, small tree of the lower level of canopy, 

flowers red-brown, 23 Nov 1956 (fl), Bernardi 

5829 (holotype, NY[2]; isotypes, K[3], NY, 

VEN[2]). Figs. 7A,B, 15A, 41, 43A-E 

Leaf blades elliptic to oblong, truncate at base; L/ 

W 1.5:1 to 2.5:1. Inflorescences densely flowered 

thyrsoid panicles, with a pair of long basal branches; 

Specimens examined. VENEZUELA. ARAGUA: Near bracts subtending basal branches foliose, 10-15 x 4- 

Colonia Tovar, 1856-1857 (fr), Fendler 2366 (frag-F, GH, 8 cm. Corollas subcampanulate with patent lobes, 18- 

K). DISTRITO FEDERAL: N slope of Cordillera de la Costa, S 25 mm long; tube conical at base, 10-13 x 4-5 mm 

of Camuri grande, Topo Tacamahaco [Tacamanaco], wide; lobes ca. '/3 of corolla length, 9-10 x 6.5-7 mm. 

quebrada Rio Grande, 9 km E of Tanques de la Electricidad Stamens with filaments 4-5 mm long; anthers lande 

Caracas, 900-1000 m, 16 Jul 1973 (fr), Morillo et al. 3298 ceolate, blunt at apex, 8.5-9 x ca. 3 mm. Pollen ex- 

(MY, VEN[3]); between Fila Maestra and Rio Caruao, San ine irregularly foveolate. Style 22-28 mm long; im- 

Jorge de Caruao, 900-1150 m, 10?32'N, 66?22'W, 16-19 Apr mature fruits ovoid-elliptic. Capsules not seen. Seeds 

1993 (fr), Meyer & Llamosa 3627 (VEN); N slope, above 

1-1.2 x 0.33-0.5 mm. 

Naiguata, quebrada Rio Camuri Grande, 800-850 m, 

10034'N, 66?42'W, 11-12 Apr 1992 (fr), Meyer et al. 2039 

(VEN). SUCRE: Quebrada Nivardo, affluent of Rio Local name. Venezuela: cambur6n 

Oscuro, 

(Bernardi 5829). 

above Mundo Nuevo, W of Cerro Humo, Paria peninsula, 

700-750 m, 7 Aug 1966 (fl), Steyermark & Rabe 96163 (K, Distribution (Fig. 41) and ecology. This variety 

VEN[2]); Dto. Cagigal, Peninsula de Paria, Rio Cumana, is known only from a single collection (Bernardi 

below trail between El Paujil and El Brasil, 10?40'N, 5829) made in the rain forests of Guatopo (State of 

62?43'W, 850 m, 21 Feb 1980 (fr), Steyermark et al. 121534 Miranda, Venezuela). On its large inflorescence, col- 

(NY, VEN); Peninsula de Paria, Cumbre de La Estrella, W lected in November, are present floral buds, flowers 

of Manacal, N of El Paujil, 800-850 m, 10?40'N, 62041'W, in anthesis, and immature ovoid fruits. 

17 Oct 1984 (fl), Knapp & Mallet 6764 (BH). 

TOBAGO: Rd. Roxborough-Parlatuvier, 800-1300 m, 

The 

14 Oct 1937 (fl), Sandwith 1757 (K[2]); Tobago Forest Re- original description of Tresanthera thyrsiflora 

serve, along Roxborough-Parlatuvier, between 4 & 5 mi was based on the terminal portion of the inflorescence, 

posts, 1200-1400 m, 20 Aug 1959 (fl), Webster et al. 9805 not the complete inflorescence, and was described by 

(A, CAS, DAV); Easterfield, 20 Dec 1912 (fl-fr), Broadway Steyermark (1964a) as a closely flowered thyrse. 

3076 (F[2], G[6], GH, NY[2], P, US[2]). 

Having available the complete set of Bernardi 5829, 

I was able to match the lateral branches of T. thyrsi- 

Variety condamineoides differs from var. thyrsiflora, realizing that its inflorescence is instead a longflora 

in having oblanceolate, basally acute (vs. oblong, pedunculate, densely flowered, thyrsoid panicle, with 

basally truncate) leaf blades (Fig. 42A), larger and less a pair of long basal branches (Fig. 43A). In addition, 

constricted corollas with reflexed (vs. sub-erect) lobes, the flowers of Bernardi 5829 are clustered in short 

and larger anthers. The inflorescences of var. cymules as illustrated for T. condamineoides (in 

condamineoides are quite variable, ranging from Karsten, Fl. Columb. 1, pl. 19) and orange-red (as in


FIG. 43. Tresanthera condamineoides var. thyrsiflora (Bernardi 5829, K and NY). A. Habit of inflorescence and mature 

leaf. B. Open flower at anthesis with dehisced anthers, before style elongation. C. Stamen, adaxial view. D. Stamen, 

lateral view. E. Immature capsule.


T. condamineoides); the calyx crenulations of 

Bernardi 5829, to which Steyermark (1964a) referred 

in the protologue, are similarly illustrated for T. 

condamineoides (in Karsten, Fl. Columb. 1, pl. 19). 

CONDAMINEA 

3. Condaminea A. P. de Candolle, Prodr. 4(4): 402. 

Sep 1830; Ruiz & Pavon, Fl. Peruv. 2: 48-49, figs. 

188, 190, 198, 199. 1799; Schumann in Martius, 

Fl. Bras. 6(6): 257. 1889; Poiret in Lamarck, 

serted because of spreading lobes, subequal; anthers 

dorsifixed near the base or at medial zone, opening 

inward by longitudinal splits. Pollen tricolporate, 

exine reticulate. Style exserted; style branches membranous, 

ovate, reversed at maturity. Ovary 2-celled, 

obconical to obovate, glabrous, placentation axile; 

ovules many in each locule, horizontally inserted; 

immature fruits green and semi-fleshy when fresh. 

Capsules oblong, obovate to turbinate, woody; disk 

septicidal dehiscence absent even in old capsules. 

Seeds irregularly 3-4-angular, trapezoid, compressed, 

testa reticulate. 

Encycl. Suppl. 3: 570-571. 1814; Kunth in 

Humboldt, Bonpland & Kunth, Nov. Gen. & Spec. 

3: 311. 1818; A. Richard, Mem. Fam. Rubiacees 

198. Dec 1830; Endlicher, Gen. P1. 553 (N. 3262). 

1837; Meisner, PI. Vasc. Gen. PI. 158 (114). 1838; 

Hooker in Bentham & Hooker, Gen. PI. 2: 12, 44- 

45. 1873; Baillon, Hist. PI. 7:471. 1880; Standley, 

Publ. Field Columbian Mus., Bot. Ser. 7: 26, 203, 

278. 1930, 1931; Standley, Field Mus. Nat. Hist., 

Bot. Ser. 13: 55-57. 1936; Steyermark in Lasser 

& Steyermark, Fl. Venez. 9(1): 241-244. 1974; 

Dwyer, Fl. Panama, Ann. Missouri Bot. Gard. 67: 

115-117. 1980; Burger & C. M. Taylor in Burger, 

Fl. Costar., Fieldiana, Bot. n.s. 33: 107-108. 1993; 

Delprete in Harling & Andersson, Fl. Ecuador (in 

press). 1999. Type species. Condaminea corymbosa 

(Ruiz & Pav6n) A. P. de Candolle. 

Shrubs to small trees, much-branched shrubs, 

rarely single-stemmed treelets; bark pale brown to 

grayish. Stipules interpetiolar and intrapetiolar, 4 

(rarely 2) at each node, connate at base, overlapping 

in bud, foliose, lanceolate, acuminate, with one central 

vein with strongly ascending secondary veins and 

several parallel veinlets departing from base, caducous. 

Leaves sessile to short-petiolate, elliptic to lanceolate 

to oblanceolate, rounded, cordate, to auriculate 

at base, acute to obtuse, usually acuminate at apex, 

dark green and waxy above, flat to undulate, thickfoliaceous 

to very coriaceous; petioles (when present) 

short and stout; domatia absent. Inflorescences terminal, 

solitary, long pedunculate, open corymbose 

panicle with a pair of opposite branches terminating 

in cymules. Flowers protandrous, hypanthium 

obconical to obovate. Calyx cupular, truncate to 

lobed; lobes (when present) 4-5, rounded to ovate; 

Condaminea is the only Rubiaceae to have largefoliose, 

acuminate stipules made of four units, two 

interpetiolar and two intrapetiolar, usually connate at 

base, forming a short sheath above the petiole attachments 

(Figs. 3F,G, 44A, 45C,D). These four stipular 

units have been interpreted either as two interpetiolar 

bifid stipules per node or four (two inter- and two 

intrapetiolar) entire stipules per node, depending on 

the basal connections. 

Condaminea is also easily detectable by its sessile 

to short-petiolate leaves, which are commonly thickcoriaceous 

and have undulate margins. Because of its 

variable leaf shape (see Fig. 47A-H) and vestiture, 

various specific and varietal names have been assigned 

to this genus, which are all reduced to synonymy under 

C. corymbosa. 

The flowers of Condaminea are thick and fleshy, 

and with valvate aestivation. During anthesis the filaments 

bend toward the lower portion of the flowers, 

and the anthers open upward by longitudinal slits, 

forming a landing platform for visitor/pollinator insects 

(commonly bees). Flowers in Condaminea are 

usually flesh-red outside and pearl-white inside. 

Seeds in Condaminea are minute, horizontally inserted, 

non-winged, truncate at apex, and usually (3-) 

4-angled. The capsules of Condaminea could be confused 

with those of Rustia, the latter characterized 

by their persistent calyx (readily caducous in Condaminea). 

Condaminea is commonly found in almost pure 

stands of single-stemmed shrubs in disturbed soil, and 

secondary vegetation, but exceptionally could grow 

into small trees. Condaminea is here recognized as a 

genus of two species (C. elegans of dubious identity, 

see below), ranging from Costa Rica to Bolivia. 

calyx caducous, leaving a white scar on capsules. 

Corolla salverform to infundibuliform (tubular with 

spreading lobes), aestivation valvate narrowly reduplicate 

with contact zone, camose, red to purple outside, 

greenish white inside. Stamens 5, partially ex- 


Condaminea was founded by A. P. de Candolle 

(1830) and was dedicated to Charles Marie de La 

Condamine. La Condamine directed an expedition to


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Ecuador to-prove that the earth is not an exact sphere, 

but rather flattened at the poles. He later descended 

the Amazon River (La Condamine, 1745) on his way 

back to France. In addition, La Condamine was the 

first to describe Cinchona, as reported in the Spanish 

translation by Ruiz (1778). La Condamine's description 

and illustration were later used by Linnaeus 

(1753) for his description of Cinchona. 

Condaminea corymbosa was first described and 

illustrated by Ruiz and Pav6n (1799) as Macrocnemum 

corymbosum. Kunth (1818) included M. corymbosum 

into "Macrocnemum sect. 2) bracteis minutis, 

sessilibus (An genus distinctum?)" [with M. tinctorium 

(= Simira rubescens (Bentham) Bremek. ex 

Steyerm.)], which was the first suggestion to treat this 

taxon as a separate genus. Macrocnemum corymbosum 

was also accepted by A. Richard (1830). 

A number of species were transferred by de 

Candolle (1830) from Macrocnemum (probably influenced 

by Kunth, 1818) to the newly founded Condaminea 

as C. corymbosa, C. tinctoria (= Simira rubescens 

(Bentham) Bremek. ex Steyerm.), C. microcarpa 

(= Bathysa sp.?), C. venosa (not Condaminea!), and C. 

glabrata (not Condaminea!), reaching the highest 

number of species ever attributed to this genus. 

Schumann (1889) described C. corymbosa var.fi 

pubescens Spruce ex Schum., which is here treated 

as synonymous with C. corymbosa. Schumann (1891) 

maintained the species of Condaminea previously 

recognized by de Candolle (1830), but placing C. tinctoria 

as a Sickingia [= Simira]. 

Rusby (1896) described C. angustifolia from narrow-leaved 

specimens collected in Bolivia, and 

Standley (1929) described C. breviflora, which he 

separated from C. corymbosa because of smaller flowers 

and petiolate leaves. I consider both of these species 

to be synonymous with C. corymbosa. 

Standley (1936) tentatively recognized three species 

of Condaminea: C. microcarpa, C. corymbosa 

(with two varieties), and C. macrophylla; in addition 

he discussed the identity of C. glabrata and C. venosa 

(see Excluded Species, below). 

Steyermark (1974) stated (translated from Spanish): 

"Four or five species [of Condaminea] are known 

from Panama to Peru and Bolivia; only one is known 

in Venezuela [C. corymbosa]." 

Dwyer (1980) stated that "there are about three 

species in this genus." In Panama he recognized two 

species: C. corymbosa and his newly described C. 

petiolata (= Rondeletia panamensis DC.; see Excluded 

Species). 

Burger and Taylor (1993) also recognized Condaminea 

as a genus of"four or five species ranging from 

Costa Rica to Venezuela, Peru, and Bolivia." 

Key to the species of Condaminea 

1. Stipules (2-)3-7(-9) cm long, at least 4 mm 

wide at base; corollas 2-3 cm long; inflorescences 

lax corymbs, with distal cymules of 

>4 flowers; leaves elliptic, ovate to obovate 

(rarely lanceolate), L/W 1.5:1 to 3.5:1 (widespread 

from Costa Rica to Bolivia) .. 1. C. corymbosa 

1. Stipules 0.5-1.7 cm long, 2-4 mm wide at base; 

corolla 0.7-1.5 cm long; inflorescences very 

reduced corymbs, with distal cymules of 2-4 

flowers; leaves lanceolate to narrowly elliptic, 

L/W 3:1 to 4:1 (Peru: Madre de Dios, Manu 

National Park) ........................................ 2. C. elegans 

1. Condaminea corymbosa (Ruiz & Pav6n) A. P. de 

Candolle, Prodr. 4: 402. 1830. Macrocnemum 

corymbosum Ruiz & Pav6n, Fl. Peruv. 2: 48, fig. 

189.1799. Type. Peru. Chinchao, 1795 (fl), Tafalla 

& Manzanilla s.n. (lectotype, MA, selected by 

Delprete, 1999b; photo-MA at F, MO, TEX). 

Figs. 3F-H, 4F, 7D-F, 15C-F, 44A-D, 45A-C, 

46A-F, 47A-H, 48, 49 

Condaminea breviflora Standley, Publ. Field Columbian 

Mus., Bot. Ser. 6: 264. 1929. Type. Ecuador. Zamora- 

Chinchipe: E slopes of E Andes of Loja, 800-1200 

m, Oct-Jan 1906 (fl), Lehmann 5651 (holotype, F, 

N.578,485; photo-F at G, VEN; isotype, K). 

Condaminea angustifolia Rusby, Mem. Torrey Bot. Club 

6: 45. 1896. Type. Bolivia. Between Guanay and 

Tipuani, Apr-Jun 1892 (fl), Bang 1429 (holotype, 

NY; isotypes, F, G, K, MO, M, NY[2], US). 

Condaminea corymbosa (Ruiz & Pav6n) A. P. de 

Candolle var. pubescens Spruce ex K. Schumann in 

Martius, Fl. Bras. 6(6): 258. 1889. Type. Peru. San 

Martin: Vie. of Tarapoto, slopes close to Rio Mayo, 

Jul-Aug 1856 (fl), Spruce 4579 (lectotype, K, selected 

by Delprete, 1999b; isolectotypes, BR, F, G[2], 

GH, K[2], NY). 

Shrubs 2-5 m tall (exceptionally small trees to 

15 m tall, to 30 cm dbh), multicaulous shrubs, excep- 

tionally single-stemmed trees; bark fissured, grayish 

brown to yellowish brown. Leafy branchlets terete 

to subterete, glabrous, dark green; older branches rug- 

ose, pale brown; lenticels sparse, whitish, ca. 1 mm 

wide. Stipules 4 at each node, connate at base (basal 

2-8 mm), foliose, long lanceolate, acuminate, gla- 

brous to puberulent to hirsutulous outside, glabrous 

with basal area of colleters inside; (2-)3-7(-9) x 4- 

9(-13) mm, persistent, pale green (to reddish green), 

caducous, leaving a white-grayish linear scar. Leaves 

sessile, subpetiolate to short-petiolate, (15-)25-55 

(-65)x (10-)15-29cm,L/W 1.5:1 to 3.5:1; lanceolate 

to elliptic to oblanceolate to obovate, rounded to cor-

106 

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FIG. 45. Condaminea corymbosa. A. Habit of a mature shrub, 3-4 m tall. B. Inflorescence. C. Detail of a young stipule 

in apical bud, before opening into four units. D. Stipules in young vegetative bud (ca. 6 cm long). 

.


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m!. . 4 

G. C. 

angustfolia, Bang 1429 type. H. Lewis 88914. 

; 

C. P er 5251. D. Cro 69930. . Subsessile leaf, Brandbyge & Asanza 30431. F-H. Petiolate leaves. Schunke 9494. 

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T. -I 

FIG. 47. Leaf variation in Condaminea corymbosa. A-D. Sessile leaves. A. Steyermark 56659. B. Skutch 3981. 

C. Pittier 5251. D. Croat 69930. E. Subsessile leaf, Brandbyge & Asanza 30431. F-H. Petiolate leaves. F. Schunke 9494. 

G. C. angustifolia, Bang 1429 type. H. Lewis 88914.


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ascending at medio-distal portion of blade; tertiary 

veins subparallel; quaternary veinlets openly reticu- 

late, evident below; petioles (when present) to 25 mm 

long, 3-7 mm thick, terete to flattened adaxially. In- 

4a.SAr* 

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florescences open corymbose panicles with 1-2(-3) 

pairs of opposite branches terminating in variably 

reduced cymules; 18-50 cm long, basal branches 4- 

25 cm long; basal portion of axis not branched 2-30


cm long; rachis terete to decussately compressed, rachis 

and branches glabrous to puberulent to hirtellous; 

flowers on terminal cymules; distal bracts 2-7 x 2-4 

mm, deltoid to narrowly triangular, sometimes similar 

to stipules (foliose, many-nerved) to 2.5 x to 5 mm, 

lanceolate, acuminate; bracteoles subtending flowers 

1-3 x 1-2 mm, deltoid. Flowers pedicellate, pedicels 

1-10 mm long, glabrous to puberulent to hirtellous; 

hypanthium obconical to narrow-obovate, 6-12 x 3- 

6 mm, glabrous to hirtellous, sometimes verrucate; 

flower buds short-clavate, apiculate at tip. Calyx 

cupular, truncate to evidently lobed, 3-10 x 5-12 mm, 

glabrous to sparsely puberulent; lobes (when present) 

(4-)5, deltoid to rounded, 1-6 mm long; leaving a 

white scar on capsule. Corolla salverform (tubular 

with spreading lobes), 2-3 cm long, carose, creamwhite 

to purple outside, greenish white inside; tube 

cylindrical, 7-16 mm long, 3-6 mm wide at base and 

6-8 mm wide at orifice, glabrous outside, glabrous to 

sparsely pilose inside, with a ring of short-sericeous 

hairs 4-7 mm from the base, at the same point of filament 

attachment; lobes (4-)5, /3--2 of corolla length, 

(3-)5-9 x 3-7 mm, ovate to oblong, pointed at apex, 

glabrous outside and inside. Stamens 5, subequal, 

attached 4-9 mm from the base of the tube and bending 

toward lower portion at anthesis; filaments 4-8 

mm long, terete distally, flattened and widened at base, 

white, sparsely pubescent at base; anthers narrowly 

elliptic, 5-8 x 1-1.5 mm, dorsifixed at medial zone, 

base elliptic to sagittate, smooth throughout, dehiscing 

by longitudinal slit toward the inside, yellow. 

Pollen exine reticulate, columellate. Style exserted, 

15-22(-26) mm long, terete to moderately compressed, 

stout, glabrous, grass-green; style branches 

ovate, membranous, often unequal, reversed at maturity, 

3-5 mm long, stigmatic surface microscopically 

(40x) papillose; immature fruits green to reddish, 

semi-fleshy, glabrous to minutely puberulent. Capsules 

obovoid to oblong-turbinate, acute to obtuse at 

base, shallowly obtuse at apex, (10-)14-20 x the year except March and September. Fruiting specimens 

were collected throughout the year except July 

and November. 

Ecology. This species is rather frequent on disturbed 

slopes of the South American Andes, and occurs 

sometimes as the first successional woody shrub 

(often in pure stands of individuals 2-3 m tall) on landslides 

and in cleared or freshly disturbed soil of recently 

built mountain roads. Because of this, Condaminea 

corymbosa plays an important role in 

containing the frequent landslides in the rainy Andean 

slopes and protecting the young seedlings of endemic 

tree species. I would very much encourage artificial 

dissemination of this short-lived shrub for rapid establishment 

of woody species on disturbed slopes so 

as to protect newly built Andean roads from frequent 

landslides. 

Reproductive biology. The flowers of Condaminea 

corymbosa are carnose, purple to flesh-red outside 

and perlaceous inside, and are historically described 

as strictly actinomorphic. Personal observations 

(Fig. 46A-F) have revealed that the flowers of this 

species are obliquely presented (sometimes erect), 

never pointing downward, and that during anthesis the 

upper filaments bend toward the lower portion of the 

corolla throat (not caused by their weight alone), arranging 

the anthers parallel to each other and dehiscing 

upward by longitudinal furrows, forming an easy 

landing platform for possible pollinators (Fig. 46A- 

F). In Ecuador the most frequent visitors are bees (not 

identified), which are probably the pollinators of this 

species ("lap-pollination"). Occasional individuals of 

C. corymbosa were observed to be associated with ants 

(pers. obs.). I suspect these ants to be only casual visitors, 

and no obvious mutualism was observed. 

6-11 

mm, light to dark brown, without lenticels, glabrous 

to puberulent throughout, with a cream-white circular 

calyx scar, 0.5-1.1 mm wide, 5-8 mm diam. Seeds 

0.57-0.77 x 0.13-0.3 mm; irregularly 3-4-angular, 

trapezoid, compressed, testa reticulate. 

Representative specimens examined. COSTA RICA. 

CARTAGO: Field of Inst. Interamericano de Ciencias 

Agricolas, Turrialba, 600 m, Feb 1949 (fr), Le6n 1484 (US). 

HEREDIA: Near Rio Col6n, between San Isidro and Col6n, 6 

Aug 1971 (fr), Dwyer et al. 523 (NY). LIMON: Sukut, Alto 

Uren, margin of Rio Ur6n, 26 Oct 1985 (fl), G6mez et al. 

23844 (MO, NY); Reserva Indigena Talamanca, confluence 

of Rio Sukut with Rio Ur6n, 09?24'N, 82?58'W, 325 m, 3 

Jul 1989 (fl), Hammel et al. 17540 (MO). PUNTARENAS: Be- 

Distribution (Fig. 49) and ecology. Common 

shrubs (see Fig. 45A), exceptionally single-stemmed 

trees, in disturbed soil and roadside vegetation on 

mountain slopes, in dry, deciduous or moist primary 

and secondary forests, coastal and mainland rain forests 

to cloud forests on eastern and western Andean 

slopes, 120-2000 m, from Costa Rica to Bolivia. 

Flowering specimens were collected in all months of 

tween Villa Neilly and Agua Buena, Golfito, 500 m, 21 Oct 

1964 (fl), Jimenez 2451 (F[2], NY); Rio Sonador, El General 

Valley, 600 m, 31 Jan 1965 (fr), Williams et al. 28805 

(F); 24.7 km SE of Brujo, on rd. Coto Brus-San Vito, 300- 

350 m, 17 Mar 1978 (fl-fr), Almeda et al. 4264 (CAS); Callas 

Gordas, Coto Brus, 1160 m, 14 Nov 1985 (fl), Gdmez- 

Laurito 10694 (F); forest of Boruca, Feb 1891 (fr), Tonduz 

3947 (BR[2]). SAN Jos:: Vic. of El General, 640 m, Jan 1939 

(fl), Skutch 3981 (GH, K, NY, US[2]).


PANAMA. CHIRIQUi: 20Ykm N of David, 14 Dec 1966 

(fl), Lewis et al. 692 (GH, K, MO, NY, UC, US); Boqueli, 

1500 m, 26 Jun 1938 (fr), Davidson 792 (A); vic. of San 

Felix, E of Chiriqui, 120 m, Dec 1911 (fl), Pittier 5251 (F, 

GH, NY, US). DARItN: 4 km S of Jaque, 2 Feb 1981 (fr), 

D 'Arcy & Sytsma 14584 (MO, NY); Piifas, 2 Mar 1967 (fr), 

Duke 10621 (ECON, MO). 

COLOMBIA. ANTIOQUIA: Vic. of Medialuna, 1700 m, 

Chama, 19 May 1944 (fl-fr), Steyermark 56659 (F[2], US[2], 

VEN); Dto. Libertador, between San Rafael and Via El 

Morro, 18 Aug 1972 (fl), Benitez de Rojas 1513 (MY[2]); 

between Merida and San Rafael del Chama, 1600 m, 16 May 

Oct 1945 (fr), Molina s.n. (US); Quebrada Verruga, near 

Uramita, 1600 m, 8 Apr 1944 (fr), Core 566 (US). CALDAS: 

8 km up Rio Apii, 1015 m, 29 Nov 1944 (fl), St. John 20871 

(UC, P). CAQUETA: Florencia, 420 m, Dec 1930 (fl) Perez- 

Arbeldez 697 (COL, US). CAUCA: El Tambo, vic. of La Paz, 

1700 m, 29 Aug 1949 (fl), Idrobo 287 (US); Tierra Adentro, 

Rio Paez drainage, Segovia, Jun 1943 (fl), Hurtado 5 (US). 

CHOC6: Mecana, N of Bahia Solano, 06016'N, 77?21W, 3 

Jan 1984 (fl), Juncosa 1660 (MO); Bahia Solano (Puerto 

Mutis), 0-20 m, 4 Jan 1973 (fr), Gentry & Forero 7195 

(VEN); rd. Tutunend6-El Carmen, Alto Rio Atrato, 29 Apr 

1979, Forero et al. 6093 (COL). CUNDINAMARCA: Intersection 

of rds. from Pandi and from Arbelaez, 1300 m, 21 Feb 

1970, Uribe-Uribe 6385 (COL); S of Silvania, on rd. to 

Fusagasuga, 1400 m, 26 May 1972, Barclay 3463 (COL). 

HUILA: S of Acevedo, 16 Aug 1944 (fl), Little 8490 (MO, 

US). META: Valley of Rio Tigre, Colombia-Uribe trail, 

03?14'N, 74?36'W, 18 Dec 1942 (fl), Fosberg & Holdridge 

19409 (UC). NARISO: Mallama, trail Planada-Pialapi, 1700 

m, 27 May 1985 (fr), Benavides 5651 (MO). NORTE DE 

SANTANDER: Region Sarare, riverside of Rio Cubug6n, El 

Indio, 13 Nov 1941, Cuatrecasas 13074 (COL). PUTUMAYO: 

Near Ecuadorian border, Rio San Miguel, between the affluents 

Bermeja and Conejo, 300 m, 13 Dec 1940 (fl), 

Cuatrecasas 11059 (US). SANTANDER: La Corcova, Tona, 

1866 m, 12 Oct 1977 (fr), Renteria et al; 658 (COL, MO); 

Bucaramanga, May 1948 (fl), Sandemer 6055 (K). TOLIMA: 

Region of Herrera, Rio Saldafia, 03?22'N, 75?56'W, 1800 

m, Jan 1945 (fr), Hurtado 111 (US[2]); canyon of Rio Ata, 

SW of La Colonia, 5 Oct 1944 (fl), Little 8757 (COL, MO); 

Prov. Mariquita, Ibagu6, 2000 m, May 1854, Triana 1782 

(3262.1) (COL); Central Cordillera, Valley of Rio 

Bermellon, near Ibagu6, 1220 m, 29 Oct 1952 (st), Humbert 

26973 (NY). VALLE: Tulua, at limits of botanical gardens, 

1300 m, 30 Jun 1984 (fl), Devia 558 (COL, MO[2]); Mun. 

Argelia, rd. La Argelia-Toro, 1700 m, 2 Sep 1983 (fl), Devia 

316 (COL, MO). 

VENEZUELA. BARINAS: Dto. Pedraza, W of La 

Escampa, Parque Nacional Sierra Nevada, 08031'N, 

70046'W, 27 Jun 1988 (fl), Dorr et al. 5719 (NY). MERIDA: 

Near M6rida, 1700 m, 24 Feb 1964 (fl-fr), Breteler 3612 

(NY, US, VEN[2]); Rio Chama, 1500 m, 15 Nov 1953 (fr), 

Little 15872 (VEN); Dto. Libertador, 2 km from Belsante, 

quebrada Carvajal, Los Curos, 1800 m, 16 Nov 1962 (fr), 

Ruiz Terdn 1204 (VEN); Aricagua, 950 m, Jun 1955 (fl), 

Bernardi 2209 (VEN); Dto. Arzobispo Chac6n, rd. Canagua 

(1500 m) - Mucuchachi 1964 (fr), Ijjfsz-Madriz 422 (VEN). 

ECUADOR. CARCHI: Trail Chical-Quinyul, 78?13'W, 

01?02'S, 1200 m, 13 Jan 1983 (fr), Barford 41389 (AAU); 

E of Maldonado, 23 Aug 1943 (fr), Steere 8060 (F); 

Maldonado, 1500 m, 4 Oct 1981 (fl), Werling & Leth-Nissen 

226 (QCA[2]); Maldonado, rd. Maldonado-Chical, 

1500-1700 m, 14 Aug 1994 (fl), Delprete & Verduga 6412 

(NY[3], QCA, QCNE, TEX, UPS[2]). COTOPAXI: Rio Pilalo, 

km 52 on Quevedo-Latacunga, 750-900 m, 29 Oct 1981 (st), 

Dodson & Dodson 12001 (F). EL ORO: Ca. 10 km SE of 

Portovelo, 850 m, 8 Oct 1944 (fl), Wiggins 10947 (DS, F); 

El Placer, W of Morona, 340 m, 28 May 1979 (fl), Albert de 

Escobar 1373 (QCA). ESMERALDAS: Lita, on Ibarra-San 

Lorenzo, 550 m, 8 Jun 1978 (fr), Madison et al. 5051 (F, 

QCA, SEL). IMBABURA: Between Guayupe and Peias 

Negras, 78014'W, 00?44'S, 1050 m, 6 Dec 1980 (fl), Holm- 

Nielsen & Jaramillo 28968 (AAU); rd. Salinas-Lita, 4 km 

before Lita, 78?26'W, 00?50'N, 580 m, 25 Jul 1992 (fl), 

Freire-Fierro et al. 2333 (QCA). MORONA-SANTIAGO: Rd. 

Suc6a-Macas, 880-1080 m, 78?07'W, 02?20'S, 26 Sep 1979 

(fr), Holm-Nielsen et al. 20567 (AAU, QCA); Bomboiza, 

15-18 km SE of Gualaquiza, 700 m, 03?27'S, 78?34'W, 1- 

3 Nov 1986 (fl), Zaruma 802 (MO, NY, QCNE, QCA). 

NAPO: Canton Archidona, rd. Hollin-Loreto, 01?04'N, 

77?36'W, 1300 m, 3 Oct 1992 (fr), Delprete & Luteyn 6205 

(AAU, MO, NY, QCA, QCNE, TEX), 6207 (NY); Rio 

Quijos with Rio Salado, below Baeza, 2 Mar 1944 (fr), 

Ownbey 2686 (F[2], MO[2], US[2]). PASTAZA: Rio Bobonaza, 

between Cachitama and outlet of Rio Bufeo, 74?40'W, 

02?20'S, 19 Jul 1980 (fl), 0llgaard et al. 34667 (AAU, 

QCA); Mera, 2 km toward valley, 27 Jun 1992 (fl), Delprete 

et al. 6049 (AAU, MO, NY, QCA, QCNE, TEX); Mera, bank 

of Rio Pastaza, ca. 1100 m, 11 Nov 1955 (fl),Asplund 18347 

(B). PICHINCHA: Rd. La Uni6n del Toachi-San Francisco de 

las Pampas, km 3, 1100-1200 m, 19 Mar 1985 (fl), Harling 

& Andersson 23151 (QCA); Bosque Protector Mindo, ridge 

between Rio Mindo and Rio Bagasal, 00?02'S, 78?48'W, 

1400 m, 15 Jul 1992 (fl), Delprete & Webster 6104 (AAU, 

DAV, F, MO, NY[2], QCA, QCNE, TEX, UPS); Canton 

Santo Domingo de los Colorados, Uni6n del Toachi, 9 Oct 

1992 (fl), Delprete et al. 6218 (NY, QCA). SucUMBmOS: Section 

NW of San Rafael waterfall, 1300 m, 11 Oct 1990 (fl), 

Jaramillo et al. 13199 (QCA). TUNGURAHUA: Rio Negro, 

00?15'S, 78020'W, 1200 m, 18 Sep 1987 (fl), Zak & 

Jaramillo 3577 (AAU, GB); rd. Bafios-Puyos, km 11.3-18, 

78?15'W, 01?24'S, 1650 m, 30 Apr 1992 (fl), Freire-Fierro 

& Luteyn 2159 (QCA). ZAMORA-CHINCHIPE: Mtn. ridge N 

of Zamora, between Rio Janeiro and Rio Yacuambi, 29 Jun 

1944 (fr), Camp A-C25 (F); rd. Zumba-Valladolid, between 

Palanuma and Palanda, 04?40'S, 7908'W, 1200 m, 13 May 

(974 m), Andean Cordillera, Cerro 1985 (fl), Stein & D 'Alessandro 2776 (QCNE); Lim6n (Gende 

Las Flores, s.d. (fl), Ruiz Terdn & L6pez Figueras 658 eral Plaza Gutierrez), rd. San Bartolo-Uni6n, 24 Aug 1994 

(VEN[3]), 4 Jun 1953 (fr), Bernardi 563 (NY); between La 

Trampa and Lagunillas, 1188-2430 m, 29 Apr 1944 (fl), 

Steyermark 56208 (VEN); Mt. San Jacinto, N side of Rio 

(fl), Delprete & Verduga 6436 (NY[2], QCA, TEX). 

PERU. AMAZONAS: Quebrada Sasa, 600 m, 25 Aug 1975 

(fl), Kayap 2012 (F, MO, NY); Rio Cenepa, vic. of Huam-


* as- 1'''--"-'-'-40 ' 

\r~ - p 

V -, 

0~~~~ -~~. pi - 

I~~~~~~~~I 

6. ~ ~ ~a 1' : 

- . 2~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I 

t i~~~~~~~~~~~~~~~~~U 

* m ~~~~~~~~~~~~~~~~~~?? 

ma-wa ~ ~ ~ ~ . ? 

:Me~~~~~~~~~~~~ 

*~~~~~~~~~~~~~~~~ 

FIG. 49. Distribution of Condaminea 

ED~~~~~~~~~~~~~~~~ 

corymbosa (circles) and C. elegans (star). 

* 's M -usessa 

FI. 9 Dstiutono Cndmne crmbsa(irle)an . elgas str 

pami, 5 km E of Chavez Valdivia, 78?30'W, 04?30'S, 200 

m, 15 Aug 1978 (fr), Ancuash 1509 (MO); Prov. Bagna, 

Valley of Rio Maration, above Cascadas de Mayasi, near 

Campamento Montenegro (kms 276-280 of Marafion rd.), 

450 m, 3-4 Sep 1962 (fl), Wurdack 1831 (VEN). 

CAJAMARCA: Prov. Ja6n, 800 m N of Chirinos, 1700 m, 30 

Jul 1943 (fl), Hodge 6048 (ECON). Cuzco: San Miguel, 

Urubamba Valley, 1800 m, 26 May 1915 (fl), Cook & Gil- 

bert 913 (F, US); Prov. La Convenci6n, E of Quillabamba, 

1200 m, 20 Apr 1981 (fl), Young & Eisenberg 255 (MO, 

NY). HuANuco: Prov. Pachitea, Parque Nacional de Iparia, 

Rio Pachitea, 1 km above Tournavista, 22 Jun 1967 (fl),


Schunke 2072 (F, NY); Chinchao, Pampayacu, 1 km E of Condaminea corymbosa is easily recognizable for 

Cochero, 1050-1200 m, 16 Feb 1973 (fl),Schunke Vigo 5807 its four long-lanceolate foliose stipules (two 

(CAS, VEN); Prov. Leoncio Prado, Dto. Rupa Rupa, Calpar interpetiolar and two intrapetiolar) at each node (Figs. 

Bella, Rio Monz6n, 700-900 m, 2 Jul 1976 (fl), Schunke 

3F-H, 44A; Fig. 45C,D for stipules in apical buds), 

9494 (F, MO, NY); Rupa Rupa, environs of Tingo Maria, its 

09?18'S, 75059'W, 680 m, 24 Mar 1982 (fl), King & Ramirez- fleshy flowers, purple outside and perlaceous in- 

Rengifo 372 (B); Chinchao, 1 Aug 1964 (fl-fr), Dwyer 6178 side, and by its sessile to short-petiolate coriaceous 

(MO[4], UC). JUNIN: Huacapistana, 1800-2400 m, 5-8 Jun 

leaves. Condaminea corymbosa is most commonly a 

1929 (fl), Killip & Smith 24130 (F, NY, US); La Merced, short-lived, single-stemmed shrub 2-4 m tall, but I 

700 m, May-Jun 1929 (fl), Killip & Smith 23506 (F, NY, have encountered it as a tree 15 m tall with 30 cm dbh. 

US); Prov. Tarma, gorge of Rio Huasahuasi, 7 km E of The type of Macrocnemum corymbosum was de- 

Huasahuasi, 12 Aug 1957 (fr), Hutchison 1153 (F, NY, UC); scribed and beautifully illustrated in Ruiz and Pav6n's 

Prov. Chanchamayo, Rio Tulumayo valley, rd. from San famous Flora Peruviana et Chilensis (1799: pl. 189); 

Ram6n to Vitoc and Chilpes, 1000-1250 m, 11o15'S, no precise type was mentioned and the protologue 

75?15'W, 9 Mar 1985 (fl-fr), Stein & Todzia 2358 (MO, NY). vaguely states: "Habitat in collibus et runcationibus 

LORETO: Above Pongo de Manseriche, hills left of Rio 

callidis, versus Chinchao, Acomayo, Pillao et Mufia 

Marafion, 250 m, 26 Dec 1931 (fr), Mexia 6351 (CAS, F, K, vicos." Most of the material collected 

MO, NY, UC, US). PUNO: Prov. Sandia, near Ketlin, 2000 

by Ruiz and 

Pav6n in this area was lost on the 

m, 4 Jun 1942 (fl), Metcalf30676 (A, UC[2]). SAN MARTIN: 

ship San Pedro de 

Boqu6ron Pass, 92 km from Tingo Maria, 400 m, 16 Dec Alcantara returning to Spain, in a shipwreck along the 

1949 (fl), Allard 22129 (UC); Zepelacio, near coast of 

Moyobamba, Portugal. Ruiz and Pavon returned to Spain 

1100 m, Jun 1934 (fl), Klug 3672 (A, CAS, F, MO); Prov. in 1788, leaving the two botanists Juan Jose Tafalla 

Lamas, trail from Tabalosos to Lamas, E of Rio Mayo, 13- and Juan A. Manzanilla (between 1793 and 1797) to 

15 Sep 1937 (fr), Belshaw 3422 (F, GH, MO, NY, UC, US); re-collect in sites of the collections lost by their pre- 

Prov. San Martin, hills NE of Tarapoto, on rd. to decessors (Estrella, 1991). Tafalla and Manzanilla sent 

Yurimaguas, 1000 m, 06030'S, 76?20'W, 11 Feb 1985 (fl- their collections to Spain, where they were then filed 

fr), Stein & Todzia 2142 (MO, NY); Prov. Rioja, Disto. Pardo in the Herbarium Peruvianum by Ruiz and Pavon and 

Miguel, 1.5 km W of Aguas Verdes, 1320-1440 m, 24 Jun included in Flora Peruviana et Chilensis. In the Her- 

1997 (fr), Sdnchez Vega & Dillon 8906 (NY), 26 Jun 1997 

barium Peruvianum of Madrid (MA) are preserved 

(fl), 9052 (NY). UCAYALI: Prov. Portillo, Bosque Von 

four 

Humboldt, quebrada Tahuahillo, 200 m, 20 Jun 1981 (fl), 

specimens annotated as Macrocnemum 

Young 942 (MO, NY); rd. Fed. Basadre, between Pucallpa corymbosum by Pav6n (in his handwriting) with the 

and San Alejandro, 08?22'S, 74?31'W, 21 Jan-3 Feb 1987 following labels: "Pentand. Monog. - Macrocnemum 

(fl), Trigoso 1025 (K). 

corymbosum 

BRAZIL. MINAs GERAIS: Mun. Ituiutaba, San Vicente, 

29 May 1950 (fl), Macedo 2402 (MO[2], US[2], TEX). 

BOLIVIA. COCHABAMBA: Prov. Chapare, vic. of San 

Onofre, 1600 m, 12 Feb 1929 (fl), Steinbach 9292 (GH, K). 

LA PAZ: Yungas, 1890 (fl), Bang 610 (DS, F, K, NY[2], UC, 

US); Nord Yungas, Covipata, Hazienza El Choro, 1700 m, 

28 Jun 1930 (fl), Buchtien 8186 (NY, US); Sur Yungas, basin 

of Rio Bopi, San Bartolom6, near Calisaya, 750-900 m, 1- 

22 Jul 1939 (fr), Krukoff 10032 (F, NY); Inquisivi, bridge 

over Rio Khara, 2 km NW of Licoma Pampa, 16?48'S, 

67?13'W, 15 Jun 1988 (fr), Lewis 88825 (MO[3]); Apolo, 

27 Feb 1902 (fl), Williams 322 (NY[2], US); Coroico, 

Pajonales, 1500 m, 19 Mar 1928 (fl), Troll 1745 (B); Prov. 

Sur Yungas, W of Chulumani, 16?23'S, 67?34'W, 1900 m, 

17 Sep 1947, Fosberg 28672 (CAS, US[2]). 

Local names and uses. Venezuela: anime chiquita 

(Steyermark 56208), cacahuito (Steyermark 56306), 

guamo (Steyermark 56208). Colombia: lengua de vaca 

(Goudot s.n.). Peru: caspi blanco (Trigoso 1025), 

chullachasqui (Trigoso 1025), larangui (Hodge 6048), 

mun baikuanin (Berlin 1812). 

Leaves used for wrapping food, and wood used in 

sugar mills (Venezuela, Steyermark 56306). 

- Vulgo. Carato. - F. P. c. 1. NO 212. - 

Ex Chinchao. - Afio 95" (Fig. 48), "Macrocnemum 

corymbosum - Vulgo. Carato insilbatis - Fl. Per. Tab. 

189 - Cochero," "Macrocnemum corymbosum - Fl. 

Peruv. 2. Ta. 189," and "Macrocnemum corymbosum 

F. P." The first specimen, which was collected by 

Tafalla and Manzanilla in 1795 and has attached to it 

the complete description of this species (Pav6n's 

handwriting), was selected by Delprete (1999b) as the 

lectotype (Fig. 48); the other specimens are treated as 

paratypes. 

Condaminea corymbosa has leaves that vary from 

sessile through subsessile, to short-pedicellate (Fig. 

47A-H) and glabrous (mostly in Central America and 

northern South America) to short-petiolate and glabrous 

to pubescent (mostly in Peru and Bolivia); in 

shape they vary from broadly elliptic to narrowly lanceolate 

(Fig. 47A-H). The floral morphology is relatively 

constant throughout its distribution, except for 

the calyx, which tends to be slightly more expanded 

in southern South America. Because of its variation 

in shape and vestiture of the vegetative parts throughout 

its range, several specific and varietal epithets have 

been assigned to this species.


This species is known from only one Brazilian ing olive-green and chartaceous, glabrous above and 

collection made in the state of Minas Gerais (Mun. below; primary and secondary veins glabrous, fine, 

Ituiutaba, San Vicente, Macedo 2402), a locality quite prominent below; secondary veins 18-22 each side; 

distant from the southernmost Bolivian Andes. Nev- tertiary veins subparallel, quaternary veinlets openly 

ertheless, this disjunct collection does not differ in any reticulate, faintly evident below; petioles 9-16 mm 

morphological character from its Andean relatives. long, 1-2 mm thick, subterete. Inflorescences very 

I have attempted to maintain Condaminea reduced corymb, long pedunculate, with or without a 

corymbosa var. pubescens from the Andes of south- pair of opposite branches terminating in extremely 

ern Peru and Bolivia, where the leaves of this species 

reduced cymules of 2-4 flowers; 16-22 cm long, 

are almost consistently petiolate, with blades thick- branches (when present) 3.5-5 cm long; basal portion 

coriaceous, often lanceolate, undulate and of axis not branched 10.5-14.5 cm 

commonly 

long; rachis terpuberulent 

to pubescent beneath. However, as a re- ete, rachis and branches glabrous; distal bracts reduced 

sult of field observations of various to scales 1-2 mm 

populations 

long, deltoid; bracteoles subtendthroughout 

its range and critical study of herbarium ing flowers ca. 0.5 mm long, deltoid. Flowers pedimaterial 

of C. corymbosa, it is obvious that the above cellate, pedicels 6-10 mm long, glabrous; hypanthium 

set of vegetative characters intergrade clinally into the obconical, ca. 6 x ca. 3 mm, glabrous; flower buds 

not seen. 

more typical elements and that Calyx cupular, evidently lobed, 2-5 x 5-7 

infraspecific recognition 

for the populations concerned is unwarranted. mm, glabrous; lobes 5, ovate, 2-4 x 3-4 mm; calyx 

The specimens Boeke 1501 (NY-2 sheets) have 

caducous, leaving a scar on mature capsule. Corolla 

yellow-velutinous pubescence on vegetative and re- campanulate, ca. 12 mm long; tube subcylindrical, ca. 

6 x ca. 4 mm, 

productive parts, capsules abruptly truncated at 

glabrous outside and inside; lobes 5, 

apex, ca. /2 of corolla 

and label data that reports "Corolla 

length, ca. 6 x ca. 3 mm, 

yellow." As stated 

narrowly 

triangular, glabrous outside and inside. Stamens and 

above, the clinal gradation in external vestiture does 

style not seen (the single flower available is in depnot 

represent a reliable taxonomic character, and the 

auperate condition). Pollen exine reticulate, columelyellow 

flowers reported by Boeke are probably an late. Capsules oblong to obovate, acute at base, shalerroneous 

observation. 

2. Condaminea elegans Delprete, sp. nov. Type. 

Peru. Madre de Di6s: Prov. Manu, Parque Nacio- 

nal Manui, Cocha Cashu uplands, 400 m, 11?45'S, 

71000'W, 13 Sep 1986 (fl-fr), Nunfez 6104 (holo- 

type, F; isotypes, MO, NY). Figs. 7H, 49, 50A,B 

lowly obtuse at apex, 9-12 x 8-9 mm, black, glabrous 

throughout, without lenticels; disk flat, with a pale 

brown circular calyx scar at edge of apex, ca. 1 mm 

wide, 6-7 mm diam. Seeds 1.3-1.87 x 0.9-1.33 mm; 

irregularly 3-4-angular, trapezoid, compressed, yel- 

lowish beige, testa reticulate. 


from the hills (350-400 m) of the Amazonian forests 

in Parque Nacional Manu, Madre de Di6s Province, 

Peru. Specimens in end-flowering/early-fruiting stage 

were collected in September. Fruiting specimens were 

collected in October. 

Arbores parve, a C. corymbosa stipularum laminis 5-17 

mm longis basi 2-4 mm latis (nec ultra 20 mm longis basi 

ultra 4 mm latis), inflorescentiis 2-4-floris (nec multi-floris), 

necnon floribus quam 15 brevioribus distantes. 

Specimens examined. PERU. MADRE DE Di6s: Parque 

Nacional Manu, Rio Manu, Cocha Jarez, hills N of Cocha 

Small trees 4-7 m tall. Leafy branchlets terete, on side trail 3 km N of Mirador trail, 350 m, 12?05'S, 

thin and delicate, 3.5-4.5 mm thick, densely rugulose, 71?04'W, 5 Oct 1989 (fr), R. Foster & E. Vivar 13333 

pale brown, glabrous. Stipules 2 each node, com- (MO, US). 

pletely connate, induplicate in bud, splitting at maturity, 

lanceolate, glabrous outside and inside, with a This species is tentatively assigned to Condasmall 

group of colleters at base inside, 5-17 x 2-4 minea, because the flowering material is depauperate 

mm, reddish brown, persistent, leaving a grayish lin- and only partially known. Its stipules, although much 

ear scar. Leaves 19-31 x 6-1 1 cm, L/W ca. 3:1 to 4:1; reduced, resemble those of Condaminea, but the seeds 

lanceolate to oblanceolate, decurrent to acute (rarely (Fig. 7H) are thicker and larger than C. corymbosa 

obtuse) at base, acute and short-acuminate at apex, the and resemble those ofRustia. No pollen material was 

acumen to 5 mm long; olive-green and dull (not wax- available in any of the specimens studied. Unfortushiny) 

above, grass-green below, thin-foliaceous; dry- nately, there was only a single depauperate corolla


.I Y 

G 

5. 

? ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~? 

? ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ? 

....?. 

..:. 

..., ? 

~~~~~~~~~~.? 

.' ? . ? ~'\ 

r.; 

,_,..._" 

__ A- 

T~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG. 50. Condaminea elegans (NriSez 6104, F). A. Habit of innlorescence, typical stipules, and iong-petiolate leaves. 

B. Young capsule with deciduous calyx still attached, 

i ' iI


without anthers or style. Some old calices were present 

on some immature capsules of N~uez 6104; these are 

larger (5-6 mm long) and more lobed than any known 

member of Rustia, and closely resembled the calices 

of Condaminea. Fruit size and shape could place this 

species in either Rustia or Condaminea. The seeds of 

Nunez 6104 resembled those of Rustia in their size, 

shape, and external sculpturing, and were much larger 

than any known Condaminea species. Because of the 

thickness of the much-enlarged deciduous calyx (re- 

duced and persistent in Rustia), the bifid short-lan- 

ceolate stipules, the absence of pellucid glands in its 

leaves, and the much-reduced inflorescences, I pro- 

visionally place this species in Condaminea (but it 

might as well be a new genus!). The definitive place- 

ment of this species will be possible only when flow- 

ering collections (and pollen material) become avail- 

able. 

Condaminea macrophylla Poeppig & Endlicher, Nov. 

Gen. Sp. 3: 30. 1845. Type. Peru. Loreto: Prov. 

Maynas, vic. of Yurimaguas, May 1929-1932 (fl), 

Poeppig 2463 (lectotype, W, here selected; photo- 

W at A, F, MO, NY, VEN; isolectotype, frag-F) 

= Capirona decorticans Spruce, J. Linn. Soc. 4: 

200. 1860. 

Its large intrapetiolar stipules, large broadly ovate 

and long-petiolate leaves, and flower morphology 

are typical of Capirona decorticans, with which it is 

synonymous. 

Condaminea microcarpa (Ruiz & Pav6n) A. P. de 


microcarpon Ruiz & Pav6n, Fl. Peruv. 2: 49, pl. 

188a. 1799. Type. Peru. Chinchao, 1795 (fl-fr) 

Tafalla & Manzanilla s.n. (lectotype, MA, here 

selected; photo-MA at F, GH, MO, NY, TEX, 

VEN) = Not Condaminea! (Bathysa?, Elaeagia?). 

The specimen preserved in the Herbarium 

Peruvianum of Madrid (MA), was collected by Tafalla 

and Manzanilla (not by Ruiz and Pav6n; see discussion 

under Condaminea corymbosa, and Estrella, 

1991) in 1795 at Chinchao (Peru). The specimen label 

reads "Pentand. Monog. - Macrocnemum microparpon 

Rejected and Doubtful Species 

Condaminea glabrata Bartling ex A. P. de Candolle, 

Prodr. 4: 402. 1830. Type. Peru. Mountains of Huanuco, 

Haenkes. n. (type not found) = not Condaminea! 

De Candolle (1830) described this species as having 

bilobed stipules (which is characteristic of Condaminea) 

with rounded-obtuse lobes (which exclude 

it from Condaminea), and inflorescences with glomerulate 

flowers (which exclude it from Condaminea). 

Standley (1936) might have overlooked this 

species, since it was not cited in his treatment of Peruvian 

Rubiaceae. The type specimens were reportedly 

annotated by Bartling as "Macrocnemum (?) 

glabratum in Herb. Haenke" and published in Condaminea 

by de Candolle (1830). The identity of this 

species cannot be ascertained from the brief description 

presented by de Candolle (1830) (no types were 

found at G, and PR material was not available), and it 

can only be concluded that it is not a Condaminea. 

- F. P. c. 1. N. 245 - Ex Chinchao. Aflo 95," 

and it is this specimen that I select as lectotype. I have 

seen a set of detailed photographs of this specimen 

(sent to me by the staff of the Madrid Herbarium) and 

the original illustration, and I concluded that it is not 

Condaminea. Its terminal inflorescence, absence of 

calycophylls, small sessile flowers with exserted anthers, 

stamens attached at the medial portion of the 

corolla tube, small capsules dehiscing loculicidally, 

and many minute seeds readily suggest a member of 

the Rondeletieae, possibly Bathysa, Elaeagia, or 

Warszewiczia. 

Condaminea petiolata Dwyer, Ann. Missouri Bot. 

Gard. 67: 117. 1980. Type. Panama. Panama: 

Along Rio Maje, 2-3 km above waterfalls, near 

Baiano lake, 100 m, 5 May 1976 (fr), Croat 34706 

(holotype, MO; isotype, MO) = Rondeletia 

panamensis A. P. de Candolle. 

The general morphology, placentation, and capsule 

dehiscence of the depauperate type specimens suggested 

that this taxon belongs to Rondeletia. The absence 

of stipules and flowers makes it difficult to 

readily identify this species. Nevertheless, the type 

specimens have been identified by David Lorence 

(annotation label, and pers. comm.) as R. panamensis. 

Condaminea tinctoria (Kunth in Humboldt, Bonpland 

& Kunth) A. P. de Candolle, Prodr. 4: 402. 1830. 

Macrocnemum tinctorium Kunth in Humboldt, 

Bonpland & Kunth, Nov. Gen. Sp. 3: 399. 1820. 

Sickingia tinctoria (Kunth in Humboldt, Bonpland 

& Kunth) K. Schumann in Martius, Fl. Brasil. 6(6): 

228. 1889, non Simira tinctoria Aublet, Hist. PI. 

Gui. Fran9. 170. 1775. Type. Venezuela. Rio 

Orinoco, between Encamarada and Carichana, 

Humboldt & Bonpland (type not found). = Simira 

rubescens (Bentham) Bremekamp ex Steyermark 

(1972: 301).


This species was described by Kunth (1820) on the 

basis of flowering material. Recent fruiting collections 

showed that the seeds of this species are unique to 

Sickingia, a genus later synonymized with Simira by 

Bremekamp (1954a) and Steyermark (1972). This 

species is utilized for red dye used on fabrics and 

wooden artifacts. 

Condaminea utilis Goudot, Compt. Rend. 18: 260. 

1844. Type. Colombia. Cundinamarca: Vic. of 

Fusagasuga, s.d., Goudot s.n. (P?). = Elaeagia 

utilis (Goudot) Weddell (1849). 

This species is characteristic of Elaeagia in hav- 

ing a resin that exudes from its young vegetative buds, 

which is utilized in Colombia and Ecuador as a pro- 

tective coating ("varniz" or "varmiz del Pasto") for 

bowls and ornamental vases. 

Condaminea venosa (Ruiz & Pav6n) A. P. de 


venosum Ruiz & Pavon, Fl. Peruv. 2: 49, pl. 190b. 

1799. Type. Peru. (Acomayo?, Pati?), 1793-1796, 

Tafalla & Manzanilla s.n. (lectotype, MA, here 

selected; photo-MA at F, GH, MO, NY, TEX, 

VEN) = not Condaminea!. 

There are two specimens of this species preserved 

at MA (both specimens without date or locality) and 

annotated by Pav6n: one as "Macrocnemum venosum 

Sp. P1. Fl. Per." and the other as "Macrocnemum 

venosum Fl. P." I select the former specimen as lectotype. 

Both specimens were probably collected by 

Tafalla and Manzanilla in 1793-1796, after Ruiz and 

Pav6n returned to Spain (see discussion under Condaminea 

corymbosa, and Estrella, 1991). Standley 

(1936) apparently overlooked this species, since it was 

not cited in his treatment of Peruvian Rubiaceae. I 

cannot position this species with certainty, but from 

the detailed photos obtained from MA and de 

Candolle's (1830) brief description, it is not a Condaminea. 

PICARDAEA 

4. Picardaea Urban, Symb. Ant. 3: 376. 1903. 

Grisebach, Cat. P1. Cub. 124. 1866; Urban, Symb. 

Ant. 7: 391. 1912; Standley, N. Amer. Fl. 32: 6- 

7. 1918; Liogier, Fl. Cuba 5: 26. 1963; Liogier, 

Fl. Espafiola 7: 355, 357. 1995. Type species. 

Picardaea haitiensis Urban [= Picardaea cubensis 

(Grisebach) Britton ex Urban]. 

Shrubs to trees, much-branched shrubs, rarely 

single-stemmed trees; bark grayish brown, smooth. 

Stipules interpetiolar, adnate to the petioles, triangular, 

persistent. Leaves petiolate, lanceolate to oblanceolate, 

acute at base, obtuse and usually short-acuminate 

at apex, chartaceous; domatia often present. 

Inflorescences terminal, solitary, short pedunculate, 

corymbose trichotomous panicle terminating in lax 3- 

flowered cymules. Flowers protandrous, hypanthium 

obconical to obovate. Calyx cupular, truncate to 

lobed; lobes (when present) 4-5, rounded to ovate; 

calyx caducous, leaving a white scar on mature capsules. 

Corolla campanulate, aestivation valvate with 

contact zone, fleshy when fresh, red to purple outside, 

yellowish white inside; tube cylindrical, expanding 

distally; lobes 5, erect, triangular. Stamens 5, partially 

exserted, subequal; anthers dorsifixed near the base, 

opening by longitudinal splits. Pollen tricolporate, 

exine reticulate. Style exserted; style branches narrowly 

oblong, reversed at maturity. Ovary 2-celled, 

obconical, glabrous, placentation axile; ovules many 

in each locule, horizontally inserted; immature fruits 

green, semi-fleshy. Capsules oblong, woody; dehiscing 

loculicidally, the halves remaining attached at base 

and at apex, disk septicidal dehiscence absent even 

in old capsules. Seeds irregularly 3-4-angular, horizontally 

compressed, testa reticulate. 

The combination of fleshy valvate corollas, cap- 

sules that dehisce loculicidally with the halves remain- 

ing attached at apex and at base (like in 

Macrocnemum, Cinchoneae), and horizontal minute 

unwinged seeds make this genus unique in the Rubi- 

aceae. 

The floral morphology of Picardaea is similar to 

that of Condaminea, but its reduced inflorescences 

resemble those of Macbrideina (Rondeletieae). Its 

capsule dehiscence is similar to that of Macrocnemum 

(Cinchoneae), and its seed morphology is similar to 

that of Rustia, Pogonopus, and several other genera 

of the Rondeletieae. Because the seeds of Picardaea 

are minute, unwinged, and horizontally attached, this 

genus cannot be placed in the Cinchoneae as defined 

by Andersson and Persson (1991), which have seeds 

vertically attached and wings strongly bipolar. This 

genus is therefore included in Rondeletieae s.l. 


Urban (1903), dedicated the name of this genus to 

Luis Picarda (1848-1901), botanical collector in 

Cuba, who collected the specimen selected as lecto- 

type ofPicardaea haitiensis. He separated this genus 

from Rustia and Condaminea as follows: "Ex affini-

118 


tate Condamineae DC. et Rustiae Klotzsch, quarum 

prior calycis limbo lobato v. dentato circumcisse 

deciduo, corollae fauce villosa, ramulis compressis, 

stipulis magnis elongatis 2-partitis chartaceis, cymis 

trichotome corymbosis, ulterior antheris basifixis, 

poro v. rimula terminali dehiscentibus, stipulis amplis 

elongatis deciduis discrepat." 

Urban (1912) added a second species to Picardaea 

by transferring Macrocnemum cubense Griseb., following 

a suggestion he encountered in an unpublished 

manuscript by Britton. Until recently, Picardaea has 

been treated as genus of two species (Standley, 1918; 

Liogier, 1963, 1995), but it is here recognized as 

monotypic (see discussion below). 

1. Picardaea cubensis (Grisebach) Britton ex Urban, 

Symb. Ant. 7: 391. 1912. Macrocnemum cubense 

Grisebach, Cat. PI. Cub. 124. 1866. Type. Cuba. 

Guantanamo: Rio Yumuri, 17 Sep 1860-64 (fl), 

Wright 2661 (holotype, GOET; isotypes, G, GH, 

HAC, K, P). Figs. 3M, 4E, 8F, 15G,H, 51A-D, 52 

Picardaea haitiensis Urban, Symb. Ant. 3: 377. 1903. 

Type. Haiti. Petionville, 500-600 m, Aug 1893 (fl), 

Picarda 1129 (B*; lectotype, GH (fragment), here 

selected). 

Shrub 4-6 m tall, exceptionally a tree to 15 m tall, 

commonly much-branched shrubs, rarely singlestemmed 

trees. Leafy branchlets glabrous, terete, 

dark-green; older branches terete, grayish brown; lenticels 

sparse, small. Stipules shallowly triangular, 

often short-acuminate, glabrous outside, glabrous with 

basal colleters inside, 1.5-3 x 2.5-3.5 mm, dark green. 

Leaves 8.2-16.5 x (3-)4-12 cm, L/W 1.5:1 to 2.5:1; 

oblanceolate to obovate, cuneate to acute at base, 

obtuse to round at apex, sometimes tapering to a short 

acumen to 15 mm long; grass-green above and below, 

foliaceous; drying olive-green, stiff-chartaceous; glabrous 

above and below; primary and secondary veins 


each side; tertiary veins openly reticulate; petioles 7- 

15 mm long, 0.7-1 mm thick, adaxially concave to 

flattened; domatia (when present) a tuft of yellowish 

hairs in the axils of secondary veins. Inflorescences 

corymbose trichotomous paniculate, with opposite 

decussate lateral branches; (5-)7-15 x angular; bracteoles subtending flowers ca. 0.5 mm 

long, deltoid. Flowers pedicellate, pedicels (1-)2.5- 

4.5 mm long, glabrous to minutely puberulent; hypanthium 

obconical, 10-12 x 5-6 mm, glabrous to 

minutely puberulent; flower buds clavate. Calyx 

cupular, truncate to shallowly lobed (or 5 minute 

teeth), 2-3 x 7-10 mm, glabrous. Corolla campanulate, 

3.2-3.5 cm long, reddish purple (to pinkish) outside, 

yellowish white inside, camose when fresh; tube 

basally cylindrical, expanding distally, 10-12 mm 

long, 5-7 mm wide at base and 15-25 mm wide at 

the orifice, glabrous outside; pilose below filament 

attachments, glabrous above, inside; lobes 5, ca. 2/5 

of corolla length, 10-16 x 7-9 mm, triangular, glabrous 

throughout, minutely papillose at margins inside. 

Stamens 5, subequal, attached 10-12 mm from 

the base of the tube; filaments 9-12 mm long, glabrous, 

distally terete, basally flattened-adnate to tube; 

anthers narrowly elliptic, 5-7.5 x 1-1.5 mm, dorsifixed 

near the base, base rounded, smooth throughout. 

Pollen exine reticulate, columellate. Style exserted, 

25-30 mm long, glabrous; style branches narrowly 

oblong, 1.3-2 x 0.3-0.4 mm, stigmatic surface microscopically 

(40x) papillose. Capsules oblong, acute 

to obtuse at base, apex truncate, 15-19 x 6.5-8 mm, 

dark brown to black at maturity, with 3-5 parallel ribs, 

glabrous to minutely puberulent; disk glabrous. Seeds 

0.45-0.64 x 0.22-0.36 mm; irregularly 3-4-angular, 

horizontally compressed, beige, not winged, truncate 

at apex, testa reticulate. 

Distribution (Fig. 52) and ecology. In primary or 

secondary forests, and in remnants of thickets on 

cliffs, usually in proximity of rivers, in limestone soil, 

50-900 m, of E Cuba and Hispaniola. Flowering specimens 

were collected in January, February, July, August, 

October, and November. Fruiting specimens were 

collected in January, February, March, and August. 

Conservation status. This is a rare species much 

threatened by ongoing destruction of the natural vegetation 

of Hispaniola, and is encountered as a shrub 

in remnant forests on limestone cliffs (rarely as a tree). 

The reproductive biology of this species has not been 

studied. 

Specimens examined: CUBA. GUANTANAMO: Valley of 

7-10 cm, lat- Rio Yumuri, Baracoa, Feb 1941 (fl-fr), Le6n & Matos 19644 

eral branches 1-2 pairs, basal portion of axis not (GH, HAC, US); Altos del Yumuri, Aug 1939 (fl), Le6n & 

Victorin 17235 (GH, HAC, NY, TEX); 

branched (when present) up to 6 cm long; rachis 

Gorge of Rio Yamuri, 

7-9 Dec 1910 (fl), Shafer 7850 (NY, US); Valley of Rio 

decussately compressed, rachis and branches glabrous Yumuri, Baracoa, 16 Jan 1960 (fl), Liogier & Acuna 7673 

to minutely puberulent; distal cymules usually 3-flow- (HAC); Valley of Rio Yumuri, Baracoa, 25 Nov 1914 (fl), 

ered; distal bracts 0.5-1.5 x ca. 1 mm, narrowly tri- Ekman 3642 (S).


- .i. , . 

. . 

FIG. 51. Picardaea cubensis (A-C from Wright 2661, GH, isolectotype; D from Le6n 19644, GH). A. Habit of inflo- 

rescence with terminal leaves and foliage leaf. B. Open flower, with old anthers and exserted style. C. Open anther, dorsal 

view. D. Mature open capsule, showing typical dehiscence.


_ 70- 60 

- -A----- ----- --.-_ - 

--- -- 

I!0 ^"W 1 IO' 

*?r3'~~~. ~F. Di I ..i 

. 

HAITI. Massif du Nord, Port-de-Paix, More Fourrise, 

400 m, 14 Aug 1925 (fl-fr), Ekman 4641 (A, G, GH[2], NY, 

US); Massif des Matheux, Grand-Bois, rd. Cormillon- 

Thamazeau, 800 m, 19 Mar 1926 (st), Ekman 5772 (GH, S); 

Massifde La Hotte, Miragoane, Jul 1927 (fl), Eyerdam 180 

(US); Massif de la Hotte, Group More Rochelois, Mira- 

goane, path Le Brum-Quatre Chemins, 700 m, 30 Mar 1927 

(fr), Ekman 7919 (G, S); Massifde la Selle, Croix-des-Bou- 

quets, Badeau, Trou-a-1'Eau, 1000 m, 15 Mar 1927 (fr), 

Ekman 7859 (S); Massifde la Selle, Gd. Goave, Papelle, 29 

Nov 1927 (fl), Ekman 9384 (S); Massif de la Hotte, dept. 

Sud, Citadelle des Platons, 30 km NNW from Torbec, 

18?16'N, 73058'W, 750 m, 10 Mar 1983 (fr), Zanoni et al. 

25634 (NY). 

DOMINICAN REPUBLIC. Prov. Barahona, La Fili- 

pana, Loma Travesia, Barahona, 700 m, 5 Jan 1977 (fl), 

Liogier & Liogier 26174 (GH, NY); Prov. Barahona, Las 

Filipinas, 450 m, Nov (fl), Fuertes 649 (G[2], NY); Prov. 

Barahona, Sierra de Bahoruco, Los Naranjos, confluence of 

Rio Bahoruco with Rio Brazo Seco, 18?05'N, 71?08'W, 50 

m, 17 Jan 1986 (fl-fr), Zanoni & Pimentel 35991 (NY, US); 

Prov. Barahona, Sierra de Bahoruco, Arroyo La Travesia, 

18?07'N, 71?07'W, 850-900 m, 12 Jan 1988 (fr), Zanoni et 

al. 40662 (NY); Prov. Barahona, Sierra de Bahoruco, Ar- 

royo El Maniel, 2 km from Los Patos de Paraiso, 17?58'N, 

71013'W, 350-470 m, 17 Jan 1985 (fl), Zanoni etal. 33125 

80 

FIG. 52. Distribution of Picardaea cubensis. 

(NY); Polo, Los Arroyos, 800 m, 28 Feb 1969 (fr), Liogier 

14353 (GH, NY); Prov. Pedernales, 13 km N of Pedernales, 

along Rio Mulito, 18009'N, 71?46'W, 230 m, 21 Oct 1991 

(fl), Thompson et al. 9859 (CM). 

Urban (1912) transferred Macrocnemum cubense 

to Picardaea, separating it from P. haitiensis by its 

yellow-white corolla [as reported on Wright 2661], 

stipule shape, and leaf size and shape. Standley (1918) 

also maintained the two names. Recent collections [Leon 

& Victorin 17235, Ekan 4641,Liogier &Liogier26174] 

showed that the corollas of Picardaea are purple-red 

outside and yellowish white (rarely reddish white) 

inside (see type label data, above), the stipules are 

constantly interpetiolar and adnate to the petioles, deltoid 

and variably acuminate, and the leaf blades vary 

in size and shape throughout the range of the species. 

For the above reasons, the characters that were used 

to separate P. cubensis from P. haitiensis are no longer 

valid, and the two taxa are here treated as synonymous, 

the previous having taxonomical priority. 

Among the duplicates of Wright 2661 examined, 

only the one present at the Herbarium Grisebachianum 

(GOET) bears Grisebach's handwriting and is here 

20 

30


treated as the holotype of Macrocnemum cubense (= Shrubs, rarely treelets, much-branched; outer 

P. cubensis). The label [manu Wright] has the follow- branches often scandent, main trunk (when present) 

ing description: "A small tree or large bush 10-20 feet short. Stipules interpetiolar, free at base, sometimes 

on farallones. Corolla valvate, thick greenish tinge adnate to the petioles, small, deltoid to shallowly triwith 

red externally and dull red within." 

angular, persistent; stipules in apical buds with axil- 

The typification of Picardaea haitiensis is diffi- lary colleters that secrete a sticky yellow resin. Leaves 

cult. The specimens of Picarda 1129, examined by petiolate, elliptic to obovate, foliaceous; petioles short, 

Urban at Berlin, were destroyed; no photos of that thickened at base; domatia absent, or tuft domatia with 

specimen were found. Among the herbaria from which sparse hairs (rarely pocket domatia). Inflorescences 

loans were received, only one leaf-fragment was lo- terminal, frondose, pyramidal, laxly paniculate, each 

cated at the Harvard herbarium (GH). It is unlikely lateral branch subtended by leaf-like bracts, terminatthat 

other duplicates of Picarda 1129 are extant, and ing in a small cyme. Flowers protandrous; hybecause 

the leaves of the GH specimen are typical of panthium obconical to obovoid (rarely turbinate). 

this genus, I have selected the leaf-fragment at GH as Calyx extremely reduced, caducous; lobes 5; 1-5 

lectotype of P. haitiensis. 

calyx lobes expanded into colorful calycophylls in 

some flowers. Calycophylls shortly stalked and 

blades ovate to obcordate to reniform. Corolla tubular 

with short 

POGONOPUS 

lobes, membranaceous or fleshy when 

fresh; tube cylindrical, basally constricted, with a ring 

5. Pogonopus Klotzsch, Monatsber. Koenigl. Preuss. of hairs at base inside; lobes 5, erect to spreading, tri- 

Acad. Wiss. Berlin 1853: 500. 1853; de Candolle, angular to ovate; aestivation valvate-reduplicate with 

Prodr. 4: 403. Sep 1830; A. Richard, Mem. Rubiac. contact zones. Stamens 5, exserted well above the 

199. Dec 1830; Weddell, Ann. Sci. Nat. 4: 66-71. 

corolla, attached near the base of the corolla tube (just 

1854; Linden & Planchon, Bot. Zeit. 12:365. 1854; above the constriction); filaments thin, with a tuft of 

Orsted, Centr.-Amer., fig. 13. 1863; Hooker in hairs at base; anthers elliptic-oblong, dorsifixed at 

Bentham & Hooker, Gen. PI. 2: 47. 1873; medial zone, dehiscing by longitudinal slit. Pollen tri- 

Schumann in Martius, Fl. Bras. 6(6): 264-266, fig. colporate, exine reticulate. Style exserted well above 

126. 1889; Schumann in Engler & Prantl, Nat. the corolla; style branches linear to oblong. Ovary 2- 

Pflanzenfam. 4(4): 21, fig. 6P. 1891; Werham, celled, turbinate, glabrous, placentation axile; ovules 

J. Bot. 50: 241-242, tab. 520. 1912; Standley, Publ. 

many in each locule, horizontally inserted; immature 

Field Columbian Mus., Bot. Ser. 7: 25, 44, 277, fruits green and semi-carose when fresh. Capsules 

370.1930, 1931; Sandwith, Kew Bull. 1949:258- obovoid, oblong to globose, truncate at apex, often 

259. 1949; Steyermark, Acta Bot. Venez. 6: 108- with white lenticels; dehiscing loculicidally, disk sep- 

110. 1971; Steyermark in Lasser & Steyermark, ticidal dehiscence present in old capsules. Seeds 

Fl. Venez. 9:235-241, fig. 36. 1974; Dwyer, Ann. 

many, horizontal, irregularly shaped, 3-5-angled, 

Missouri Bot. Gard. 67: 329-333, fig. 69. 1980; compressed. 

Burger & Taylor in Burger, Fl. Costaricensis, 

Fieldiana, But. n.s. 33: 218, fig. 16. 1993. Type Pogonopus is commonly encountered as beautiful 

species. Pogonopus ottonis Klotzsch [= P. speci- multi-stemmed shrubs (rarely single-stemmed trees), 

osus (Jacquin) K. Schumann var. speciosus]. with highly ornamental red-purple calycophylls. It is 

Carmenocania Wemham, J. Bot. 50:241. 1912. Type spe- easily recognizable because of its long-tubular pinkcies. 

Carmenocaniaporphyrantha Wemham [= Pogon- red flowers, small deltoid stipules with a tuft of hairs 

opus speciosus (Jacquin) K. Schumann var. speciosus]. between them, and small capsules with horizontally 

Chrysoxylon Weddell, nom. poster. homonym., Monogr. inserted seeds. Its seeds are minute, non-winged, trun- 

Cinch. 100. 1849, species typica Chrysoxylonfebri- cate at the apex, and posses a reticulate and minutely 

fugum Weddell, nom. inval. [= Pogonopus tubulosus tuberculate exotesta (within the primary reticulation). 

(A. Richard in A. P. de Candolle) K. Schumann]; Its bark is often reported to be a good remedy for 

non Chrysoxylon Casaretto, Nov. Stirp. Dec., decas 

malaria and intermittent fevers, and more 

7: recently for 

59. 1843 (= Plathymenia Bentham, Mimosaceae). 

its antitumor 

Howardia Weddell, Ann. Sci. Nat. 4(1): 66, fig. 10. 1854; 

properties (for additional information see 

non Howardia Klotzsch, nom. under 

poster. homonym., Uses). 

Monatsber. Deutsch. Akad. Wiss. Berlin 1859: 584, Pogonopus (see Figs. 53, 55, 56, 57) is closely re- 

607. 1860 [= Aristolochia Linnaeus, Aristolochi- lated to Pinckneya, from southeastern North America. 

aceae]. Type species. Howardiafebrifuga (Weddell) The latter differs from the former in having corolla 

Weddell [= P. tubulosus (A. Richard) K. Schumann]. cream-white, its throat without a ring of hairs (present


in Pogonopus), corolla lobes long-narrow pubescent var. speciosus); and H. richardi (based on Macroinside, 

and coiling outward at anthesis (vs. spreading), cnemum tubulosum A. Rich.) and H. febrifuga (based 

seeds arranged in two vertical ranks, with orbicular on Chrysoxylonfebrifugum), both using Bolivian colwing 

(vs. many minute unwinged seeds), and exotesta lections (and both = P. tubulosus). Weddell (1854) 

secondarily reticulate (within the main reticulation). also commented that Calycophyllum tubulosum (A. 

The distribution of Pogonopus is an interesting Rich. in DC.) DC. shared many affinities with his Howcase 

of geographical vicariance (see Fig. 54). Re- ardia and, instead of making a new combination, he 

stricted to tropical seasonal forests, this genus occurs reduced this species under synonymy of his H. richardi. 

in two areas of distribution that mirror to each other Hooker (1873) established the tribe Condamiwith 

respect to the equator. In the northern hemisphere neeae, and in his third subtribe Pinckneyinae he placed 

Pogonopus ranges from Chiapas (southern Mexico) Pogonopus and Pinckneya. 

to northern Colombia, from 6?N to 16?N; while in the Baillon (1880) included Pogonopus under Pincksouthern 

hemisphere it occurs from central Peru to neya, simply stating (translated from French): "In 

northern Argentina, from 9?S to 22?S. Pogonopus also those named Pogonopus, the corolla, instead of beoccurs 

along elevational gradients and is usually en- ing tomentose internally, is glabrous. The fruit is 

countered at medium elevations of(50-)500-1400 m. ovoid, whilst in the true Pinckneya [as Eupinckneya] 

I attribute the occurrence of Pogonopus in the low- it is more globular and subdidymous. They are shrubs 

lands to its introduction into cultivation for its orna- of the two Americas, with rather large and showy 

mental and medicinal properties. Pogonopus is here flowers in terminal or axillary clusters of cymes." 

treated as a genus of three species and two varieties. Wernham (1912) described Carmenocania, a genus 

that he included in the Mussaendeae. The type 


collection, Schlim 755, is from the vicinity of the city 

of Carmen, in the province of Ocafia (hence the ge- 

Pogonopus was established by Klotzsch in 1853; neric name), Venezuela [and not Colombia as reported 

its name was derived from the Greek words iwoycov 

by Wernham (1912), Standley (1930a), and Sandwith 

[pogon = beard] and oOG [pous = feet], because of 

(1949)]. Wernham apparently overlooked that this was 

its basally bearded filaments. The first species dethe 

same collection previously used by Weddell 

scribed under this genus was P. ottonis [= P. speciosus 

(1854) to establish Howardia grandiflora. 

(Jacq.) K. Schum. var. speciosus], from material col- 

Standley (1918) reduced Chrysoxylon and 

lected in Venezuela, with the specific name dedicated 

Howardia to synonymy under Pogonopus, but subto 

its collector [Otto 901]. Klotzsch (1853) also noted 

sequently he (Standley, 1930a) listed Pogonopus (in 

that another species (from Costa Rica) should be inthe 

Condamineeae) and Carmenocania (in the 

cluded in Pogonopus, namely Macrocnemum exsert- 

Mussaendeae) without noting their similarities. 

um Orsted (Orsted, 1852), but he did not make the of- 

Sandwith (1949) recognized the mistakes made both 

ficial transfer. When Klotzsch (1853) described P. 

by Wernham (1912) and Standley (1930a) and synottonis, 

he was perhaps unaware that this species was 

onymized Carmenocania under Pogonopus. 

already described by Jacquin (1797) as Macrocnemum 

Steyermark (1971) divided Pogonopus speciosus 

speciosus. The latter was transferred to Pogonopus into three subspecies: exsertus, speciosus, and sandmuch 

later by Schumann (1889). 

withianus. He later reduced subsp. sandwithianus to 

Augustin Pyramus de Candolle (1830) transferred 

a variety under P. speciosus subsp. speciosus 

Macrocnemum tubulosum A. Rich. in DC. [= Pogon- 

(Steyermark, 1974). In the present treatment P. exopus 

tubulosus (A. Rich. in DC.) K. Schumann] to 

sertus is returned to specific rank (Orsted, 1863) and 

Calycophyllum, which in turn was placed next to 

the subspecific divisions are no longer maintained. As 

Pinckneya [as "Pinkneya"], within his Cinchoneae. 

here treated, Pogonopus consists of three species, one 

Weddell (1849) described Chrysoxylon, an invalid 

with two varieties, collectively ranging from southname 

because this generic epithet was previously used 

ern Mexico to northern Argentina. 

by Casaretto (1843) for a mimosoid genus. Realizing 

the invalidity of his Chrysoxylon, Weddell renamed 

this genus Howardia, ignoring (purposely or otherwise) 

the fact that Klotzsch (1853) had meanwhile 

described it as Pogonopus (the valid name for this 

genus). Under Howardia, Weddell (1854) described 

four species: H. caracasensis and H. grandiflora, both 

using Venezuelan collections (and both = P. speciosus 

Key to the species of Pogonopus 

1. Corolla lobes erect to narrowly spreading; 

corolla tube narrowly cylindrical, not expanded 

or ampliate at the orifice, mem branaceous 

when fresh, glabrous to minutely 

puberulent outside.


2. Corollas 25-30 mm long, with erect lobes; two halves. Leaves 

tube cylindrical for the whole 

9-19(-25) x (3-)5-7(-10) cm, 

length; 

lobes '/s of corolla length; blades of lanceolate, 

calycoelliptic 

to elliptic-obovate, acute at base, 

phylls


IP 

F I 

FIG. 53. Pogonopus exsertus (A-D from Nee 9125, MO; E from Croat 8959, MO). A. Habit of inflorescence. B. Open 

flower with exserted anthers. C. Close up of calyx . D. Detail of external pubescence of corolla lobe. E. Detail of intemal 

glandular pubescence of corolla lobe. F. Mature capsule.


....... _ --[-.-..... .....--.---.. ... 

dp 

~ or- 1lil 1- - #- A- ---I-- __ 1 

FIG. 54. Distribution of Pogonopus exsertus (circles), P. speciosus var. speciosus (diamonds), P. speciosus var. 

sandwithianus (squares) and P. tubulosus (triangles). 

callous zone, at pilose ring); filaments 20-28 mm 

long, with a basal tuft of hairs; anthers elliptic-oblong, 

1.9-2.3 x ca. 0.8 mm, dorsifixed at medial zone, with 

a narrow space between the thecae, base and apex 

rounded. Pollen exine reticulate. Style exserted, 27- 

35 mm long, microscopically (40x) papillose, green; 

style branches linear to oblong, 2-2.5 x 0.5 mm, re- 

versed, stigmatic surface microscopically (40x) glan- 

dular. Capsules obovoid, base acute, apex shallowly 

hemispherical, 5-8 x 4-6 mm, with sparse lenticels, 

dark brown, minutely puberulent below the disk; disk 

glabrous, black, with lenticels, with a white tubercule 

at the point of attachment of the style. Seeds 0.38- 

0.58 x 0.23-0.35 mm. 

Distribution (Fig. 54) and ecology. Seasonal ev- 

ergreen and deciduous forests, secondary tropical 

moist forest, and relative remnants, in steep wooded


slopes and thickets, usually near small creeks or in 

river banks, 50-1000 m, from S Mexico to NW Colombia. 

Flowering specimens were collected in January, 

February, and July through December, but especially 

from September through January. Fruiting 

specimens were collected in January, March, September, 

November, and December. 

Representative specimens examined. MEXICO. 

Shattuck 688 (MO); N of Paraiso, near Gaillard, 22 Nov 1970 

(fl), Croat 12652 (MO, NY); Rio Pedro Miguel, near E 

Paraiso, 7 Jan 1924 (fl), Standley 29972 (A, GH, US); between 

Juan Mina and El Vigia, 11 Jan 1911 (fr), Pittier 2396 

(NY); tributary of Rio Chagres, 3 km SW of Cerro Brewster, 

14 Dec 1967 (fl), Lewis et al. 3392 (COL, UC, VEN); 

Serrania de Maje, trail along Rio Ipeti Grande, 300 m, 

08?55'N, 78?31'W, 28 Jan 1984 (fl), Churchill & Nevers 

4446 (MO[2]); Forest Preserve, jet. Boyd-Roosevelt Hwy. 

with Madden Rd. 1 km from Madden Dam, 1967, Mori 421 

CHIAPAS: Rio Testecapa, 10 km SE of Mapastepec, 180 m, 

24 Dec 1972 (fr), Breedlove & Thorne 30715 (NY); Mal 

(MO). UNCERTAIN LOCALITY: Maume and Gorgone, Jan 1858 

(fr), Wagner 3495 (M), 3496 (M). 

Paso, Carrillo Puerto, 250 m, 6 Oct 1984 (fl), Ventura & COLOMBIA. ANTIOQUIA: Uraba, Dabeiba, Riosucio, 

L6pez 424 (MO); Escuintla, 15 Nov 1947 (fl), Matuda 16811 

(NY); Huixtla, Rio Huixtla, 4 Dec 1985 (fl), Ventura & 

350 m, 11 Jan 1947 (fl), Uribe-Uribe 1441 (COL, F, NY); 

rd. Antioquia-Anza, Rio Cauca, 06?25'N, 75?52'W, 2 Nov 

L6pez 2856 (NY); Azulejo, Margarita, 250 m, 7 Jan 1950 

(fr), Matuda 18748 (MO). 

GUATEMALA. ALTA VERAPAZ: Finca Mocca, 750 m, 

25 Nov 1919 (fl), Johnson 57 (NY, US); Tucuru, 18 Aug 

1979 (fl), Boeke & Utzischneider 2925 (MO); Pancajche, 18 

1988 (fl), Zarucchi et al. 7044 (F, NY). CHOC6: Mun. 

Riosucio, Parque Natural Los Katios, Camino Peye-Tilupo, 

60 m, 22 Nov 1976 (fl), Le6n 455 (COL, MO); Parque 

Nacional Los Katios, Quebrada de La Tigra, 18 Jan 1983 

(fr), Zuluaga 1184 (COL). 

Nov 1920 (fl-fr), Johnson 1022 (US). SAN MARCOS: Finca 

Armenia, San Rafael, at the base of Cuesta to Carrizal (cul- 

CULTIVATED. UNITED STATES: FLORIDA: Miami, 

Fairchild Tropical Garden, 24 Jan 1969 (fl), Gillis 7535 (A), 

tivated), 1300-1600 m, 9 Aug 1980 (fr), Dwyer 15265 (MO); 

near Malacatan, 400 m, Nov-Dec 1940 (fl), Grant 630 (A, 

24 Nov 1971 (fl), Gillis 11168 (A). HAWAII: Oahu, U.S. 

Nursery, 4 Jan 1937 (fl), E.L.C. s.n. (A). GUATEMALA. 

GH). SANTA ROSA: Plains N of Los Cerritos, rd. 

Chiquimulilla-El Ahumado, 75 m, 7 Dec 1940 (fl), Standley 

79549 (US). SUCHITEPtQUEZ: Cocales (cultivated), 215 m, 

ALTA VERAPAZ: Near Tucuri, 500 m, Oct 1917 (fl), Popenoe 

802 (A, GH). HONDURAS. Lancetilla Experimental Station, 

11 Jul 1934 (fl), Yuncker 4529 (NY). COSTA RICA. 

5 Jan 1939 (fl), Standley 62055 (A). 

PUNTARENAS: Golfito nursery, 23 Aug 1950 (fl), Allen 5621 

EL SALVADOR. AHUACHAPAN: Near Salto de Atehue- (DS). CUBA. SANTA CLARA: Cienfuegos, Soledad, Harvard 

cia, 600 m, 22 Jan 1947 (fr), Standley & Padilla 2885 (MO). Tropical Garden, 1 Feb 1932 (fr),Jack 8430 (A[2], NY, US), 

SAN MIGUEL: La Libertad, vic. of La Laguna, 830 m, Cruz 11 Mar 1931 (fr), Jack 8173 (A). PUERTO RICO. Rio 

WB-00482 (MO). SAN SALVADOR: La Cebadilla, 1922 (fl), Piedras, Experimental Station, 1 Dec 1964 (fl), Wagner 711 

Calder6n 1225 (GH, MO, NY, US); on Rt. 382, W of Sanso- (A), 30 Aug 1965 (fr), Wagner 897 (A); Humacao Playa, 22 

natejct., 6 Sep 1964 (fl), Elias et al. 583 (MO). SAN VICENTE: Jan 1966 (fl), Howard 16044 (A); Pennock's Nursery, Hato 

Vic. of San Vicente, 2-11 Mar 1922 (fr), Standley 21672 Rey, Howard & Nevling 16910 (A). 

(GH, US). SANTA ANA: Trail to Cerro Verde, 1000 m, 30 Sep 

1959 (fr), Montalvo 3849 (MO); E side of Lake Coatepeque, Local names and uses. El Salvador: chorcha de 

790 m, 2 Dec 1970 (fl), Harmon & Fuentes 4965 (MO, NY). 

COSTA RICA. GUANACASTE: Colonia Carmina, 100 m, 

Feb 1912 (fl), Jimenez 541 (US). PUNTARENAS: Cant6n de 

Osa, Rio T6rralba, E of Palmar del Norte, 100 m, 08?56'N, 

83?20'W, 3 Jan 1990 (fl), Hammel et al. 17726 (MO). SAN 

gallo (Calderon 1922). Guatemala: quina (Standley 

79549). Panama: chibigui (Bristan 1425), madrofio. 

Reportedly a local remedy for malaria in Guatemala 

(Standley 79549). 

JosE: Vic. of El General, cliff above Rio Pacuan, 610 m, Jan 

1939 (fl), Skutch 3950 (A, MO, NY, US[2]); Puriscal, San 

Martin, 09?44'N, 84024'W, 800 m, 29 Nov 1989 (fl), Jimenez 

et al. 751 (MO); trail San Miguel de Turricanes-Piedras 

Negras de Mora, Rio Virilla, 5 Dec 1985 (fl), Gdmez et al. 

Pogonopus exsertus is a very elegant, glabrescent 

shrub with slender branches, easily distinguishable 

from the other two species in having thin, delicate, 

lax inflorescences. It has been found in cultivation 

24504 (MO). 

PANAMA. COLON: Around Porto Bello, 18 Jan 1911 (flfr), 

Pittier 2476 (GH[2], NY, US); ca. 2 km E of Transisthmian 

Hwy to Salamanca, 100 m, 19 Dec 1972 (fr), Gentry 

6720 (MO, NY); near Salamanca, hydrografic station, Rio 

Pequeni, 70-80 m, 15 Dec 1934 (fl), Dodge etal. 16977(BR, 

G, GH, NY, US). DARIEN: Banks of Rio Piedras, 26 Jun 1959 

(fr), Stern et al. 724 (GH, MO, US); Rio Aruza, 24 Oct 1967 

(fl), Bristan 1346 (MO[2]); Parque Nacional del Darien, 

ridge between Rio Topalisa and Rio Pucuro, 08?03'N, 

77?17'W, 750-850 m, 18 Oct 1987 (fl), Cuadros etal. 3884 

(MO). PANAMA: Barro Colorado Island, 6 Jan 1932 (fr), 

throughout its natural distribution and elsewhere (see 

above). Pogonopus exsertus is well deserving of specific 

rank because it differs in many floral and vegetative 

morphological characters from its South American 

counterparts (P. speciosus and P. tubulosus). 

Pogonopus exsertus closely resembles P. tubulosus, 

from Peru and southward (rather than P. speciosus, 

from the Caribbean coast of Colombia and Venezuela), 

in having glabrescent, narrowly cylindrical, 

membranous corollas; the former differing from the 

latter in having corolla lobes erect (vs. narrowly


spreading), shorter corollas (to 3 cm long), and smaller bescent hairs; flowers on distal branches in lax 

calycophylls that are basally and apically acute. cymules of 9-50 flowers; distal bracts usually leaf- 

Pogonopus exsertus is easily distinguished from P. like, petiolate, elliptic-obovate, to 22 x 8 cm, decreasspeciosus 

in the latter having thick and fleshy corol- ing in size distally, often reduced to deltoid scales 20las 

that are gradually expanded toward the orifice, and 30 x 5-6 mm; bracteoles subtending flowers to 3 x 

having reproductive and vegetative parts densely yel- ca. 0.5 mm, linear, densely golden-puberulent. Flowlow 

pubescent. 

ers pedicellate, pedicels 2-10 mm long, puberulent 

At least two specimens of this species [Wagners.n. and pubescent intermixed, to hirsutulous to pilose; 

(M), Maume and Gorgone, Jan 1858 (fr); and Hayes hypanthium obconical to obovoid, 2.5-4 x 2-2.8 mm, 

164 (BR), Isthmus of Panama, Oct 1861 (fl)] were yellow appressed puberulent, with lenticels; young 

annotated by Schumann as "Pogonopus speciosus flower buds minute glandular-pilose (var. speciosus) 

(tubulosus) var. panamensis K. Schumann," but this or golden-hirsutulous (var. sandwithianus). Calyx 

name was never published. 

extremely reduced, 0.3-0.7 x 2.5-4 mm, densely yellow 

pubescent; lobes 5, barely distinguishable, narrowly 

triangular to linear, 1-3 x 0.5-1 mm; in some 

flowers 1-5 calyx lobes expanded into calycophylls. 

2. Pogonopus speciosus (Jacquin) K. Schumann in Calycophylls (3-)4.5-6.5 

Martius, Fl. Bras. 6(6): 265. 1889. 

Figs. 3K,L, 4G, 8C,D, 16C, 54, 55A-E, 56A-F 

Shrubs 4-8 m tall, rarely small trees to 10 m tall, 

to 30 cm dbh, much-branched, with spreading crown; 

main trunk (when present) short, terete; central 

branches erect, outer branches often scandent; bark 

brown. Leafy branchlets sparsely to densely antrorse 

short-pubescent to hirsutulous to hirsute; older 

branches sparsely short pubescent or sparsely hirsutulous; 

lenticels cream-white, punctiform to linear, to 5 

mm long. Stipules deltoid to shallowly triangular, 

acuminate, sparsely long pilose (var. speciosus) or 

hirsutulous to hirsute (var. sandwithianus), with elongated 

colleters (to 0.5 mm long) inside; 3.5-6.5 x 3.5- 

5 mm, persistent. Leaves 11-25 x 5-10.5 cm, elliptic 

to obovate, cuneate to decurrent at base, acute to 

obtuse and acuminate at apex, acumen to 3 cm long; 

dark green above, pale green below, stiff-foliaceous; 

drying olive-green, stiff-chartaceous; glabrous above, 

glabrous to minutely puberulent to hirtellous below; 

primary and secondary veins sparsely short appressedpubescent; 

secondary veins 9-14 each side, tertiary 

veins starting subparallel and subreticulate in the 

middle; petioles 0.8-2.2 mm long, 1-1.5 mm thick, 

adaxially concave to flattened, minutely puberulent; 

domatia absent (var. sandwithianus) or a tuft of sparse 

erect to curled hairs, rarely pocket domatia. Inflorescences 

laxly paniculate, pyramidal, with opposite 

decussate branches, each lateral branch subtended by 

leaf-like bracts, terminating in a small cyme; 11-30 cm 

long, basal branches 4-20 cm long, lateral branches 

2-5 pairs, basal portion of axis not branched 5-13 cm 

long; rachis terete, with punctiform to linear lenticels, 

rachis and branches sparsely short puberulent to 

densely golden-pubescent (to hirtellous), branchlets 

of terminal cymes densely puberulent, with few pu- 

x (1-)2.5-4.5 cm, blades 

ovate to widely ovate, base obtuse to rounded (rarely 

cordate), apex acute (with the very tip rounded), palmately 

veined (3-5 veins each side), glabrous to glabrescent; 

primary and secondary veins sparsely puberulent; 

pink, lavender to crimson red; stalks 1.5-2 

cm long. Corolla tubular gradually expanded (var. 

speciosus, Fig. 55F) or abruptly ampliate (var. 

sandwithianus, Fig. 55B) at orifice, 25-34 mm long, 

aestivation valvate-reduplicate with contact zone, 

thick and fleshy when fresh, rose-pink to pink-red, 

sometimes with white-pink lobes; tube subcylindrical 

with distal expansion, 20-28 mm long, 2.5-4 mm 

wide at base and 3-7 mm wide at orifice, sparsely (at 

base) to densely (distally) antrorse puberulent-pubescent 

(var. speciosus, Fig. 55G), or golden-hirsute (var. 

sandwithianus, Fig. 55C) outside; with a white callous 

basal area 5-7.5 mm long, topped by a dense ring 

of hispid or golden ascending hairs (ca. 1 mm long), 

the medial and superior zones of throat glabrous to 

pubescent; lobes 4-5, spreading, 2.9-6.5 x 2.2-5 mm, 

narrowly to widely ovate, antrorse yellow pubescent 

outside (var. speciosus), or golden hirtellous (var. 

sandwithianus), with sparse glandular hairs inside. 

Stamens 5, exserted, slightly unequal, attached 6-7 mm 

from the base of the tube (just above callous zone, at 

pilose ring); filaments 27-37 mm long, white, with a 

basal tuft of hairs; anthers elliptic, 2-3 x 0.8-1.1 mm, 

dorsifixed at medial zone, base rounded or elliptic, 

apex acute or round, with no space between the thecae, 

purple to blue when fresh, drying black. Pollen 

exine reticulate. Style exserted, 32-40 mm long, glabrous, 

green; style branches oblong to oblate, 1.25- 

1.5 x 0.5-0.75 mm, reversed, stigmatic surface microscopically 

(40x) papillose or with glandular hairs. 

Capsules ellipsoid-ovoid (var. speciosus, Fig. 551) or 

oblong to subglobose (var. sandwithianus, Fig. 55E), 

base rounded, apex round to shallowly hemispheri-


FIG. 55. A-E. Pogonopus speciosus var. sandwithianus (from fresh material and Delprete & Apreza 6359, TEX). 

A. Habit of inflorescence. B. Flower (note corolla tube abruptly ampliate). C. Detail of external vestiture of corolla. 

D. Detail of internal glandular pubescence of corolla lobe. E. Portion mature capsule (note hirsute pubescence). F-I. Pogonopus 

speciosus var. speciosus (Williams 12408, F). F. Flower (note corolla tube gradually ampliate). G. Detail of external vestiture of 

corolla. H. Detail of internal glandular pubescence of corolla lobe. I. Portion mature capsule (note appressed pubescence). 

re 

tnn

Systematic Treatment of the Genera Studied 

... 

129 

?~~~~ ? ::, ~' ..? :. .O j.,:.~ ..'.~..~, ? ., '131 1 :~...,:..., __ 

. . 9' 

_@).. .o '-~ _:: ...'


glabrous to minute puberulent below, veins 

sparsely short appressed-pubescent, secondary veins 

sparsely appressed-pubescent below; capsules 9-14 each 

ellipsoid-ovoid, 7-8 mm side; 

long, golden 

petioles 0.8-2.2 mm long, 1-1.5 mm 

puberulent; 

disk shallowly hemispherical (Venezuela: thick, minutely puberulent; tufts domatia, with sparse 

Caribbean coast) ............................. 2a. var. speciosus 

erect to curled hairs (rarely pocket domatia). Inflo- 

1. Corollas golden-hirsutulous outside; corolla tube rescences pyramidal, laxly paniculate; 11-30 cm long, 

abruptly ampliate at orifice, 3-4 mm at base, basal branches 6-20 cm long, lateral branches 2-5 

5-7 mm at orifice; lobes 4-6.5 mm long, ca. '/7 pairs, basal portion of axis not branched 5-12 cm long; 

of corolla length; leaf blades sparsely to densely rachis and branches sparsely short puberulent, branchgolden-hirtellous 

below, veins sparsely to densely lets of terminal cymes densely puberulent, with sparse 

hirsutulous to hirsute below; capsules oblong to 

subglobose, 6-7 mm long, short pubescent; disk 

pubescent hairs intermixed; distal branches lax 

with outer elevated ring (do-nut shaped) and cymules of 20-50 flowers; distal leaf-like bracts to 

concave (Colombia: Sierra Nevada de Santa Marta 22 x 8 cm, decreasing in size distally, often reduced 

and Serrania de Perija) ........... 2b. var. sandwithianus to deltoid scales 20-30 x 5-6 mm; bracteoles subtending 

flowers to 3 x ca. 0.5 mm, linear, densely goldenpuberulent. 

Flower pedicels 3-10 mm long, puberulent 

and pubescent intermixed; young flower buds 

2a. Pogonopus speciosus (Jacquin) K. Schumann var. minute glandular-pilose throughout (hairs to 0.5 mm 

speciosus. Pogonopus speciosus subsp. speciosus long). Calyx 0.3-0.7 x 2.5-4 mm, densely yellow 

var. speciosus Steyermark in Lasser & Steyermark, pubescent; lobes barely distinguishable, narrowly tri- 

Flora de Venezuela 9(1): 239. 1974. Chrysoxylon angular, 1-2 x 0.5-1 mm; in some flowers 1-3 calyx 

speciosum (Jacquin) O. Kuntze, Revis. Gen. PI. lobes expanded into calycophylls. Calycophylls (3- 

278. 1891. Mussaenda speciosa (Jacquin) Poiret )4-6.5 x (1-)2.5-4.5 cm, blades ovate to widely ovate, 

in Lam. Encyc. Suppl. 4: 37. 1816. Macrocnemum base obtuse to rounded (rarely cordate), apex acute 

speciosus Jacquin, PI. hort. schoenbr. 1: 19, tab. (with the very tip rounded), palmately veined (3-5), 

43. 1797. Type. Cultivated in Sch6nbrunn Botani- glabrous to glabrescent; primary and secondary veins 

cal Gardens, Vienna, brought from Venezuela, sparsely puberulent; pink, lavender to crimson red; 

Caracas, 1787-1788, Bredemeyer s.n. (lectotype, stalks 1.5-2 cm long. Corolla tubular, gradually ex- 

W, here selected; isolectotypes, W[3]). Figs. 8C, panded at orifice, 25-32 mm long, rose-pink to pink- 

54, 55F-I 

red, sometimes with whitish pink lobes; tube cylin- 

Pogonopus caracasensis (Weddell) Nichols, Diet. Gard. drical with gradual expansion, 20-28 mm long, 

3:175. 1886. Howardia caracasensis Weddell, Ann. 2.5-3.5 mm wide at base and 4-6 mm wide at orifice, 

Sci. Nat. 4(1): 70, pl. 10 (figs. 4-7). 1854. Type. sparsely (at base) to densely (distally) antrorse puberu- 

Venezuela. Distrito Federal: La Guayra, Oct 1843 lent-pubescent outside; with a white callous basal area 

(fl), Funck 463 (holotype, P). 

6-7.5 mm long, with a dense tuft of golden ascend- 

Pogonopus ottonis Klotzsch, Monastber. Koenigl. ing hairs (ca. 1 mm long), the medial and superior 

Preuss. Acad. Wiss. Berlin 1853: 500. 1853. Type. zones of throat glabrous; lobes 2.9-4.2 x 2.2-3.4 mm, 

Venezuela. Caracas, 1838, E. Otto 901 (B*; isotypes, 

narrowly ovate, antrorse yellow pubescent outside, 

BR?, K?, W?). 

with 

Howardia grandiflora Weddell, Ann. Sci. Nat. 4(1): 70. sparse short glandular hairs (0.2-0.4 mm long) 

1854. Type. Venezuela. Ocafia: Carmen, ca. 3000 inside. Stamens attached 6-7 mm from the base of 

m, Aug 1846-1852 (fl), Schlim 755 (holotype, P; the tube; filaments 31-37 mm long, subequal, with a 

isotypes, BR[2], G[2], K, frag-F). 

basal tuft of hairs; anthers elliptic, 2-2.6 x 0.8-1.1 

Carmenocania porphyrantha Werham, J. Bot. 50: 241, mm, base rounded, apex acute. Style 32-38 mm long, 

fig. 520. 1912. Type. Venezuela. Ocafia: Carmen, microscopically (40x) multicostate; style branches 

ca. 3000 m, Aug 1846-1852 (fl), Schlim 755 (holo- oblong, 1.25-1.5 x 0.5 mm, stigmatic surface microtype, 

K; isotypes, BR[2], G[2], P, frag-F). scopically (40x) papillose. Capsules ellipsoid-ovoid, 

apex round to shallowly hemispherical, 7-8 x 5-6 

Leafy branchlets sparsely to densely antrorse mm, sparsely golden-puberulent below the disk; disk 

short-pubescent; older branches sparsely short pubes- shallowly hemispherical, with a white tubercule at the 

cent. Stipules deltoid, acuminate, sparsely long-pi- point of attachment of the style, with some lenticels. 

lose; 4-5 mm x 4-5 mm, persistent. Leaves 11.3-24.2 Seeds 0.6-0.67 x 0.31-0.49 mm wide. 

x 5-10.5 cm, elliptic to obovate, cuneate to decurrent 

at base, acute to obtuse and acuminate at apex, acu- Distribution (Fig. 54) and ecology. Steep, dry 

men to 3 cm long; glabrous above, glabrous to mi- forested slopes, semi-deciduous and transitional fornutely 

puberulent below; primary and secondary veins ests, disturbed forest margins, and shady ravines, 300-


1200 m, in the coastal cordillera of Venezuela. Flow- Guarenas, 700 m, 22 Dec 1939 (fl-fr), Williams 12408 (A, 

ering specimens were collected during rainy season ECON, F, UC); Parque Nacional El Avila, southern slope, 

in January, August, September, October, and Decem- NNE of Caucagiiita, Qda. Tacamahaca, 10?30'N, 44?45'W, 

ber. Fruiting specimens were collected in 

850-1000 m, 3 

January, 

Aug 1977 (fl), Steyermark et al. 114008 

March, and November. 

(VEN). YARACUY: San Felipe, Rio Cocorotico, Camino del 

Isleflo, 18 Oct 1950 (fl-fr), Trujillo 1413 (MY). 

CULTIVATED. BRAZIL. Rio de Janeiro: Rio de 

Representative specimens examined. VENEZUELA. 

Janeiro, 3 Feb 1891 (fl), Glaziou 18305 (G[2], BR), 18309 

ARAGUA: Slopes of La Fila Maestra of Interior Coastal Cor- 

(P), and 18905 (P). 

dillera, NE of San Casimiro, 650 m, 26 Oct 1963 (fl), 

Steyermark 91807 (NY, P); Parque Nacional, below Rancho 

Grande, toward Maracay, 9 Dec 1943 (fl), Steyermark 54921 

Local names. Venezuela: coralito (Machado 11- 

(A, VEN); near Aragua-Miranda border, between Hda. 1984), naranjillo montafiero (Saer 642, Steyermark et 

Negro and Hda. Negrito, S of La Fila Maestra of Interior al. 114008). 

Coastal Cordillera, NE of San Casimiro, 650 m, 26 Oct 1963 

(fl), Steyermark 65741 (VEN); Parque Nacional Aragua, 600 This variety differs from Pogonopus speciosus var. 

m, 25 Aug 1938 (fl), Williams 10299 (F[2], G, MO, UC, US); sandwithianus in having corollas gradually expanded 

Carmen, 300 m, s.n. (fl), Williams 10415 (A, F[2], US); Mun. 

(Fig. 55F) at the orifice (vs. abruptly expanded, see 

Ocumare de la Costa, Parque Nacional Pittier, 800 m, 

Fig. 55B) with ascending pubescence, leaves puberu- 

Castillo 1966 (MO); Rancho Grande, 19 Sep 1975 (fl), lent 

Rodriguez 266 (MY); Maracay, Parque Ecol6gico, Fac. 

(vs. hirsute) below, and capsules ellipsoid-ovoid 

Agron6mica, 2 Jun 1957 (fl), Trujillo 3416 (MY). DISTRITO (vs. oblong-subglobose). 

FEDERAL: Between Caracas and La Guaira, Oct 1916 The 

(fl), complex nomenclatural history of this species 

Rose & Rose 21691 (GH, US); Valley of Tacagua, rd. is explained in the section dedicated to the taxonomic 

Caracas-La Guaira, near El Pauji, 1 Feb 1925 (fr), Pittier history of Pogonopus (see above). Schlim 755-P is the 

11672 (A, VEN); vic. of Caracas, 2 Nov 1916 (fr), Rose & holotype of Howardia grandiflora; Schlim 755-K is 

Rose s.n. (US[2]); vic. of Caracas, 1842, Lansberge 214 (A, the holotype of Carmenocania porphyrantha; and 

MO, NY, US); Caracas, La Guaira, 950 m, Nov 1938 (fl), Schlim 755-BR was annotated (unknown handwriting) 

Williams 10646 (UC); Quebrada Las Adjuntas, 10?32'N, as Pinckneya ionantha. Schlim 755 is the isotype of 

66?52'W, 11 Nov 1984 (fl-fr), Machado 11-1984 (VEN); rd. 

both Howardia grandiflora (Weddell, 1854), and 

to electricity plant behind Naiquetia, 12 Mar 1974 (fr), Gentry 

& Morillo 10339 (MO, NY); Parque Nacional El Carmenocaniaporphyranta (Wernham, 1912), which 

Avila, 

are both 

1200 m, 19 Jul 1978 (fl), Maroto & Salas 123 (MO); vic. of synonymous with Pogonopus speciosus var. 

Colonia Tovar, 1854-1855, Fendler 584 (MO, NY); near speciosus. 

Zigzag, between Caracas and Puerto Cabello, 18 Oct 1921 In the herbarium of the Museum of Natural His- 

(fl), Pittier 58 (F, G[2], GH, NY, US); Maracay (St. Ottilien), tory of Vienna (W), are preserved four specimens, 

450 m, Aug 1925 (fl), Zehnter 38 (M[2]); El Valle, Hda. once part of the Jacquin herbarium, all labeled as 

Sosa, 11 Sep 1949 (fl), Trujillo & Ferncndez 293 (MY); "Macrocnemum/Calycophyllum speciosum Jacq. 

Colonia Tovar, 1852 (fl), Moritz 840 (B*, F, G, P). GUARICO: 

Hwy. to Caucagua, 10 km NWN of Altagracia de Orituco, 

440 m, 18 Nov 1973 (fl-fr), Davidse 4174 (MO, VEN); Dto. 

Libertador, Rio Chichiriviche, 1-2 km S of Chichiriviche, 

20-50 m, 10?32'N, 67?14'W, 9 Oct 1976 (fl), Steyermark & 

Carrerio-Espinoza 112706 (F, MO, NY, VEN). LARA: Dto. 

Jim6nez, Paso de Angostura, represa de Yacambu, confluence 

of Qda. Honda y Rio Yacambu, 09?4 1'N, 69?3 'W, 35 

km E of Sanare, 500 m, 27-28 Dec 1973 (fr), Steyermark & 

Carreno-Espinoza 108790 (VEN); El Altar, near La Miel, 4 

Jan 1967 (fr), Smith VI-280 (VEN); Serrania de Terepaima, 

S of Barquisimento, 800-1000 m, Aug 1930 (fl), Scer 

d'Heguert 642 (M, VEN). MIRANDA: Rio Guarita, Morro de 

La Gaurita, 10?27'N, 66?48'W, 900-950 m, 17 Aug 1975, 

Steyermark & Berry 112141 (VEN); Morro de La Guarita, 

880 m, 11 Jan 1976 (fr), Berry 1847 (VEN); near El Encantado, 

7 Oct 1917 (fr), Pittier 7468(VEN); Qda. Las Comadres, 

near Las Mostazas, 1100 m, Nov 1924 (fl),Allart 236 (VEN); 

Parque Nacional Guatopo, 16 Jan 1967 (fr), L6pez-Palacios 

1939 (VEN), 17 Jan 1967 (fr), 1940 (NY); vic. of El Petare, 

11 Sep 1921 (fl),Pittier9799(F, GH[2], K, NY, US, VEN); 

- 

Caracas - Bredemeyer." Bredemeyer collected in 

Venezuela in 1787-1788 (Lanjouw & Stafleu, 1954), 

and part of his living collections were transported to 

the greenhouses of the Schonbrunn gardens. In turn, 

Jacquin described Macrocnemum speciosus from living 

material cultivated in the Schonbrunn greenhouses. 

Of these four specimens, one bears the annotation 

(in Schumann's handwriting): "Pogonopus 

speciosus - det. Schumann in Fl. Bras.," and I select 

this specimen as lectotype ofM. speciosus. The three 

other specimens are treated as isolectotypes. 

Specimens of Moritz 840 (B*, F, P; photo-B at G, 

GH, MO, NY, VEN; photo-P at F, MO, VEN), collected 

in Colonia Tovar (Venezuela) were annotated 

by Klotzsch as "Pogonopus moritzii Klotzsch" and by 

Schumann as "Pogonopus speciosus var. ottonis K. 

Schumann." Because both names remained unpublished, 

Moritz 840 should not be considered type 

material.


2b. Pogonopus speciosus (Jacquin) K. Schumann var. 

sandwithianus Steyermark. Pogonopus speciosus 

(Jacquin) K. Schumann subsp. speciosus var. sandwithianus 

(Steyermark) Steyermark in Lasser & 

Steyermark, Flora de Venezuela 9(1): 240. 1974. 

Pogonopus speciosus (Jacquin) K. Schumann 

subsp. sandwithianus Steyermark, Acta Bot. Venez. 

6: 110. 1971. Type. Colombia. Magdalena: Santa 

Marta, near Bonda, 50 m, 20 Aug 1898 (fl), H. H. 

Smith 100 (holotype, NY; isotypes, A, BR, CM, 

K, LL, MO, GH, COL, F, US, VEN). 

Figs. 3K,L, 4G, 8D, 16D, 54, 55A-E, 56A-F 

Pinckneya ionantha Planchon & Linden, Bot. Zeit. 12: 

365. 1854. Type: Colombia. Magdalena: Valley of 

Rio Magdalena, Momp6s, 1847, Schlim s.n. (lectotype, 

BR, here selected). 

Leafy branchlets hirsutulous to hirsute, with 

golden-yellow hairs; older branches sparsely hirsutulous. 

Stipules shallowly triangular, strongly acuminate; 

hirsutulous to hirtellous; 3.5-6.5 x 3.5-5 mm. 

Leaves 10-29 x 5.5-10 cm, elliptic to obovate, cuneate 

to decurrent at base, acute to obtuse and acuminate 

at apex, acumen to 3 cm long, sparsely puberulent 

(at least on midrib) above, sparsely to densely 

golden-hirtellous below; primary and secondary veins 

sparsely to densely hirsutulous to hirsute, secondary 

veins 9-16 each side; petioles 4-36 mm long, 1-2.5 

mm thick, densely golden-hirsutulous, giving a velvety 

appearance; domatia absent. Inflorescences 

subpyramidal, laxly paniculate; 14-19(-22) cm long, 

basal branches 4-8 cm long, lateral branches 2-3 

pairs, basal portion of axis not branched 6-8(-13) cm 

long; rachis and branches densely golden-pubescent 

(to hirtellous), becoming sparsely pubescent in older 

parts, branchlets of terminal cymes densely goldenhirtellous; 

distal branches congested cymules of 9- 

20 flowers; distal bracts leaf-like, to 12.5 x 5.5 cm, 

decreasing in size distally, down to 3 x 1.5 cm; 

bracteoles subtending flowers 1.5-1.5 x ca. 0.5 mm, 

linear, densely golden-pubescent. Flower pedicels 2- 

8 mm long, densely hirsutulous to pilose; hypanthium 

densely hirsutulous to hirtellous; young flower buds 

golden-hirsutulous throughout (hairs to 0.7-1.1 mm 

long). Calyx 0.3-0.7 x 3.5-4 mm, densely yellow 

pubescent; lobes usually linear, 1.5-3 x 0.5-1 mm; 

in some flowers 1-5 calyx lobes expanded into 

calycophylls. Calycophylls 4.5-5(-9) x 2.5-4.5(-5.5) 

cm, blades ovate to widely ovate, base obtuse to 

rounded (rarely subcordate), apex acute (with the very 

tip rounded), palmately 3-5-veined, glabrous to glabrescent; 

primary and secondary veins sparsely puberulent, 

pink, dark pink, to pink-red; stalks 1-2 cm 

long. Corolla tubular, abruptly ampliate at orifice, 26- 

34 mm long, rose-pink, lilac to pink-red; tube subcylindrical 

with abrupt expansion, 23-25 mm long, 3-4 

mm wide at base and 5-7 mm wide at orifice, densely 

erect golden-hirsutulous outside; with a white callous 

basal area 5-7 mm long, with a dense tuft of whitehispid 

hairs (ca. 1 mm long), medial zone of throat 

glabrous, superior zone sparsely long-pubescent; lobes 

4-6.5 x 3.8-5 mm, widely ovate, golden hirtellous- 

pubescent outside, with sparse elongate glandular 

hairs (0.5-0.7 mm long) inside. Stamens attached ca. 

7 mm from the base of the tube; filaments 27-34 mm 

long; anthers narrowly elliptic, 2.6-3 x 0.8-1 mm, 

base narrowly elliptic, apex round, short-ligulate. 

Style 32-40 mm long, microscopically (40x) papil- 

lose; style branches oblate, ca. 1.5 x ca. 0.75 mm, 

stigmatic surface with microscopic (40x) glandular 

hairs. Capsules oblong to subglobose, apex sub- 

truncate to shallowly hemispherical, 6-7 x 5-6 mm, 

short-pubescent below the disk; disk an elevated ring 

(do-nut shaped), concave, with a brown tubercule at 

the point of attachment of the style. Seeds 0.44-0.64 x 

0.35-0.42 mm. 

Local names and uses. Colombia: quina (Kernan 

167), quina morada (Delprete & Apreza 6359). 

Distribution (Fig. 54) and ecology. Seasonal de- 

ciduous forested slopes, often in limestone soil, 50- 

1200 m, endemic to Sierra Nevada de Santa Marta 

(Magdalena Department, Colombia) and Serrania de 

Perija (Cesar Department, Colombia). Flowering 

specimens were collected in July, August, September, 

November, and December. Fruiting specimens were 

collected in February, August, and December. 

Reproductive biology. The flowers of this vari- 

ety are visited by both butterflies and bees of uniden- 

tified species. Some flower buds are initially robbed 

by wasps, which make lateral incisions at the height 

of the nectaries, and are secondarily robbed by ants 

that enter through these incisions (Fig. 56E). 

Specimens examined. COLOMBIA. CESAR: Mun. 

Valledupar, Pueblo Bello, 1200 m, 13 Jul 1983 (fl), Cuadros 

1677 (COL, US); Serrania de Perija, 8 km NE of [Augustin] 

Codazzi, 275 m, 10 Feb 1945 (fr), M. L. Grant 10891 (NY); 

near Codazzi, 150 m, 30 Sep 1943 (fl), Haught 3705 (F, NY, 

US); Cerro Chimichagua, 35 km NE of El Banco, 11 Aug 

1938 (fl), Haught 2236 (COL, F, NY, US). MAGDALENA: 

Parque Nacional Tayrona, 50-150 m, I 11 8'N, 74005'W, 50- 

150 m, 2 Nov 1972 (fl), Kirkbride 2637 (COL, MO, NY, 

US, VEN); Parque Nacional Tayrona, quebrada Cinto, 50 

m, Sep 1979 (fl), Moreno 193 (COL); Parque Nacional


Tayrona, 74?02'W, 11?20'N, 23 Aug 1986 (fr), Gentry & 

Cuadros 55525 (MO); Sierra Nevada de Santa Marta, rd. 

from San Pablo to San Pedro de la Sierra, 2 km before El 

Mico, 10?53'N, 74?09'W, 300 m, 29 Dec 1992 (fl), Delprete 

& Apreza 6359 (AAU, COL, F, GB, MO, NY, TEX[3]); 

Bonda, near Santa Marta, 26 Aug 1944 (fr), Romero- 

Castaneda 132 (COL, F[2]); Mun. Fundaci6n, N of Santa 

Rosa, 400-800 m, 6 Aug 1971 (fl), Romero-Castaneda 

11198 (COL, MO, NY); Sierra Nevada de Santa Marta, Rio 

Frio, Jul 1925 (fl), Walker 1277 (F, GH, MO, US); trail 

Cenizo-Santa Rosa, Rio Tucurinca, 31 Jul 1944 (fl), 300 m, 

Kernan 167 (US[2]); Magdalena Valley, La Jagua, 8 Sep 

1924 (fl), Allen 622 (F, MO); Linden s.n. (BR). 

This variety is readily distinguishable from Pogonopus 

speciosus var. speciosus in having corollas 

abruptly ampliate (Fig. 55B) at the orifice (vs. gradually 

ampliate, Fig. 55F) and golden hirsutulous (Fig. 

55C) outside (vs. appressed-pubescent, Fig. 55G), leaf 

blades sparsely to densely golden-hirtellous below, 

secondary veins and capsules golden-hirsute, and capsular 

disk with an outer elevated ring [doughnut 

shaped] (vs. hemispherical). Also, this variety has 

ard in A. P. de Candolle) A. P. de Candolle, Prodr. 

4: 367. Sep 1830. Macrocnemum tubulosum A. 

Richard in A. P. de Candolle, Prodr. 4: 403. Sep 

1830. Macrocnemum tubulosum A. Richard, nom. 

superfl., Mem. Fam. Rubiacees. Dec 1830 (reimp. 

Mem. Soc. Hist. Nat. Paris. ser. 3, 5: 279. 1834). 

Howardia richardi Weddell, nom. superfl., Ann. 

Sci. Nat. 4(1): 69, pl. 10 (figs. 8-10). 1854. Type. 

Brazil. Locality, date, and collector unknown (holotype, 

P-Richard Herbarium; photo-P at F). 

Figs. 8E, 16D, 54, 57A-E 

Pogonopus febrifugus (Weddell) J. D. Hooker in 

Bentham, G. & J. D. Hooker, Gen. P1. 2: 47. 1873. 

Howardiafebrifuga Weddell, Ann. Sci. Nat. 4(1): 

67, pl. 10 (figs. 1-3). 1854. Chrysoxylonfebrifugum 

Weddell, nom. inval., Monogr. Cinch. 100. 1849. 

Type. Bolivia. Enquisivi, Dec 1846 (fl), Weddell 

4190 (lectotype, P, here selected; isolectotype, US). 

Pogonopus febrifugus (Wedd.) J. D. Hooker var. 

macrosema Hutchinson, Kew Bull. 1910: 200. 1910. 

Type. Bolivia. "Forest of eastern Bolivia," 6 Mar 

1910 (fl), Gosling s.n. (lectotype, K, here selected). 

been found only in the Sierra Nevada de Santa Marta 

and the Serrania de Perija (Colombia, Figs. 54, 56A), 

whereas var. speciosus is endemic to the Venezuelan 

Caribbean coast (Fig. 54). 

Sandwith (1949) helped to clarify the complex 

taxonomic history of Pogonopus speciosus (see 

above) and pointed out the differences between the 

specimens collected in Venezuela (Schlim 755, Funk 

463) and those collected in Sierra Nevada de Santa 

Marta (Smith 100, Dawe 698). Steyermark, following 

Sandwith's (1949) observations, described this 

taxon first as a subspecies (Steyermark, 1971) of P. 

speciosus and later as a variety (Steyermark, 1974) 

of P. speciosus subsp. speciosus, dedicating its epithet 

to its first careful observer, Sandwith himself. 

The type of Pinckneya ionantha was cited in the 

original description (Linden & Planchon, 1854) as: 

"von Schlim [s.n.] in Neu-Granada [Colombia], an 

den brenden Ufern des Magalenenflusse [valley of 

Magdalena River] in der Prov. Mompox [Mompos] 

gesammelt." In the herbarium of BR there is a specimen 

simply labeled "Pinckneia ionantha [orth. var., 

handwriting unknown], Neu-Granada, Schlim." This 

specimen is here selected as the lectotype of this taxon. 

Shrubs 2.5-8 m tall, exceptionally singlestemmed 

small trees to 10 m tall, to 30 cm dbh, muchbranched 

shrubs to treelets, with central branches or 

central stem erect, outer branches usually scandent; 

bark smooth, brownish gray. Leafy branchlets 

antrorse minute-puberulent; older branches sparsely 

puberulent to glabrate; lenticels cream-white, punctiform 

to linear, to 5-6 mm long. Stipules very shallowly 

triangular, not acuminate to short acuminate, 

glabrescent to puberulent outside, with elongated 

colleters (to 0.5 mm long) inside, 3-4 x 4.5-6 mm, 

persistent; older stipules often breaking into two 

halves. Leaves (8-)13.5-25 

3. Pogonopus tubulosus (A. Richard in A. P. de 

Candolle) K. Schumann in Martius, Fl. Bras. 6(6): 

265. 1889. Chrysoxylon tubulosum (A. Richard in 

A. P. de Candolle) O. Kuntze, Revis. Gen. P1. 1: 

278. 1891. Calycophyllum tubulosum (A. Rich- 

x 5-12.5 cm, narrowly 

to widely elliptic to elliptic-obovate, cuneate to longdecurrent 

at base, acute to obtuse and short-acuminate 

at apex, acumen to 3 cm long; dark green above, pale 

green below, stiff-foliaceous; drying olive-green to 

yellow-green, stiff-chartaceous; glabrous above, glabrous 

to puberulent to white-pilose below; primary 

and secondary veins glabrous to puberulent to sparsely 

pilose, secondary veins (5-)7-12 each side, tertiary 

veins subparallel, quaternary veinlets reticulate; petioles 

4-24 mm long, 1-2.5 mm thick, adaxially concave 

to flattened, glabrous to minute puberulent; 

domatia absent or tuft-domatia of sparse pubescent 

hairs. Inflorescences short-pyramidal, paniculate with 

opposite decussate branches, each lateral branch subtended 

by leaf-like bracts, terminating in a lax cymule; 

(7.5-)14-29 cm long, basal branches (4-)6-15 cm 

long, lateral branches to 4 pairs, basal portion of axis 

not branched 3.5-8.5 cm long; rachis terete, with puncti-


..~~~~~~~~~~~~. 

OOF ~ ~ ~ 

-Ac 

I~~~~~~J 

FIG. 5'7. Pogonopus tubulosus (A-E from Alfaro 35 9, M O; F firom Lourteig 714, F). A. Habit of inflorescence. B. Open 

glandular pubescence of corolla lobe. F. Mature capsule.


form to linear lenticels (to 3 mm long), rachis and 

branches glabrous to puberulent; flowers on distal 

branches alternate or in lax cymules of 4-8 flowers; 

distal bracts leaf-like, petiolate, elliptic-obovate, to 16 x 

7 mm, decreasing in size distally down to 3-9 x 1-3 mm; 

bracteoles subtending flowers 0.5-1.2 x ca. 0.5 mm, 

narrowly triangular. Flowers pedicellate, pedicels 6- 

20 mm long, minute puberulent; hypanthium 

obconical to narrowly oblong to obovoid, 3-4.5 x 2- 

3.5 mm, glabrous to minute puberulent; young flower 

buds minutely puberulent at apex. Calyx tube ex- 

tremely reduced, 0.3-0.6(-1) x 2.8-5.2 mm, glabrous 

to minute puberulent; lobes 5, deltoid to acuminate, 

0.5-1.5 x 0.3-0.5 mm; in some flowers 1-5 lobes 

expanding into foliose expansions of different sizes, 

from pale green to pink elongate lobes, to dark-pink 

calycophylls. Calycophylls (3.2-)4.5-15.5 x (2-)4- 

13 cm, blades elliptic-acuminate to ovate to bilobate 

with or without a small mucron in the middle, base 

obtuse to cordate (rarely acute), apex round to obtuse 

(rarely acute), palmately veined, blade glabrous above 

and below, primary veins and/or blade margins sparsely 

puberulent, bright pink, pink-red, wine-red to crimson; 

stalks 0.7-2.5 cm long. Corolla narrowly tubular, 

with a moderate constriction near the base, 30-50 mm 

Distribution (Fig. 54) and ecology. Fairly com- 

mon in primary and secondary forests and their rem- 

nants, in seasonal-dry forests (500-1400 m), and up 

to cloud forests (1600-2800 m), sometimes in clay- 

soils, of Peru, Bolivia, western Brazil, and northern 

Argentina. Flowering specimens were collected in 

January, February, March (most collections), April, 

May, June, July, and December. Fruiting specimens 

were collected in February, March, May, June, July, 

and November. 

Representative specimens examined. PERU. Cuzco: 

Prov. Calca, Chanchamayo, Lares Valley, 12 Mar 1929 (flfr), 

Weberbauer 7940 (F, GH); Prov. La Convenci6n, Sambray 

valley, W affluent of Vilcanota, 1000-1500 m, 10 Mar 

1936 (fr), Mexia 8035 (F, G, GH, K, MO, US). HUANCAVE- 

LICA: Vuelo-Pata, above Virgen-Pampa, SE of Tintay, 1300 

m, 13 Apr 1964 (fl), Tovar 4629 (US[2]). JUNIN: E of La 

Merced, 4 Nov 1962 (fr), Schunke 6214 (F); rd. La Merced- 

Villa Rica, 75?17'W, 10?54'S, 6 Jan 1984 (fl), Smith et al. 

5605 (F, MO, NY); La Merced, May-Jun 1929 (fl), Killip 

& Smith 23811 (F, NY); Prov. Tarma, San Ram6n, 800 m, 

16 Mar 1976 (fl), Plowman & Kennedy 5649 (F, GH, K). 

BRAZIL. ACRE: Rio Acre, Str. Seringal Sao Francisco, 

Mar 1911 (fl), Ule 9848 (B, G, US); Cobija, Jan 1912, Ule 

9848b (K, MG, UC); Cruzeiro do Sul, sub-base do Projecto 

RADAM, 15 Mar 1976 (fi), Ramos & Mota 368 (INPA, 

MG). MATO GROSSO: Jauru, km 16 W on rd. MT-248, 7 May 

1995 (fl), Hatschbach et al. 62451 (US). MATO GROSSO DO 

long, membranaceous (not fleshy) when fresh, pale SUL: Mun. Aquidauana, Piraputanga, 18 Feb 1970 (fl), 

purple, lilac to dark red outside, cream-white inside; Hatschbach 23784 (MO, NY, US); Mato Grosso, Dec 1827 

tube subcylindrical to narrowly obconical, abruptly (fl), Riedel 1229 (BR); Campos dos Urupas, Feb 1919 (fl), 

ampliate at orifice, 28-46 mm long, 2.5-6 mm wide Kuhlmann 2353 (R, RB). ROND6NIA: Vilhena, 12?58'S, 

at base and 4-7 mm wide at orifice, glabrous to minute 61?01'W, 8 Apr 1977 (fr), Barroso et al. 66 (RB); Mun. Ouro 

puberulent outside; with a white callous basal area 10- Preto do Oeste, Reserva Ecol6gica do CNPq-INPA, 10? 1 'S, 

15 mm long, topped by a ring of white sericeous hairs, 62?63'W, 29 Jun 1984 (fr), Cid et al. 4896 (F, GH, INPA, K, 

the medial and superior zones of throat glabrous; lobes MG, NY, US); Ouro Preto do Oeste, rd. Cuiaba-Porto Velho, 

km 

5, narrowly spreading, 3-4.3 x 382, 10?11'S, 62?63'W, 1 Jul 1984 

2.3-4 mm, narrowly to 

(fr), Cid et al. 4939 

(INPA, MG, NY, US); Mun. Jam, rd. Cuiaba-Porto Velho, 

widely ovate, glabrous to sparsely puberulent outside, km 428, 3 Jul 1984 (fl-fr), Cid et al. 5022 (K, MG, NY, US); 

microscopically (40x) glandular (0.02-0.15 mm long) Mun. Jiparana, Gleba km 20, 9 Apr 1983 (fl), Paulino Filho 

inside. Stamens exserted, slightly unequal, attached & Silva 83-96 (INPA), Silva 6125 (INPA, MG). 

10-15 mm from the base of the tube (ust above cal- BOLIVIA. LA PAZ: Tipuani-Guanay, Dec 1892 (fl), 

lous zone, at pilose ring); filaments 38-56 mm long, Bang 1716 (A, F[2], G[2], GH[3], M, MO[2], NY[2], US[2]), 

white, with a basal tuft of sericeous hairs; anthers nar- Apr-Jun 1892, 1344 (A, F, G, GH, K, M, MO, NY[3], 

rowly elliptic, 2.4-3 x 1-1.1 mm, dorsifixed at me- US[21); Prov. Nor-Yungas, 10 km N of Yolosa, 16?09'S, 

dial zone, base rounded, apex acute, purple-blue when 67?43'W, 1200 m, 25 Mar 1982 (fl-fr), Solomon 7321 (MO, 

fresh, drying black. Pollen exine reticulate. Style ex- NY[2]); 41.5 km N of Loyosa, 16?00'S, 67036'W, 1000 m, 

26 Mar 1982, Solomon 7359 

serted, 35-54 mm long, glabrous, white; style branches 

(INPA); Prov. Nor-Yungas, 

Milluguaya, 900 m, Dec 1917 (fl), Buchtien 761 (F, GH, MO, 

membranaceous, linear-oblong, 1.5-2.5 x 0.5-0.7 mm, 

NY, US); Prov. Sur-Yungas, basin of Rio Bopi, San 

reversed, stigmatic surface microscopically (40x) papil- Bartolom6, 750-900 m, 1-22 Jul 1939 (fr), Krukoff 10054, 

lose. Capsules subglobose to ellipsoid, base rounded, 10167 (A, F, G[2], K, MO, NY, UC); Coroico, 1000 m, 16 

apex subtruncate, 7-10 x 5-9 mm, with circular white Mar 1928 (fl), Troll 1514 (B). SANTA CRUZ: Lagunillas, 900 

lenticels, dark brown, glabrous to puberulent below m, May 1949 (fl), Cdrdenas 4261 (GH, US); Las Tejitas, 

the disk; disk sometimes exceeding the calyx, gla- 420 m, 2 Mar 1945 (fl), Peredo 525 (A, NY); Prov. Florida, 

brous, dark brown. Seeds 0.51-0.76 x 0.29-0.47 mm. gorge of Rio Pirai, 1 km NE of Rio Bermejo, 800 m, 18?1 1'S, 

63?33'W, 16 Feb 1988 (fl-fr), Nee & Saldias 36320 (NY); 

Prov. Cordillera, Alto Parapeti, 800 m, 25 Jan 1980 (fl), de 

Michel 41 (UB); Prov. Nuflo de Chavez, San Ram6n, 21 Feb


1991 (fr), Quevedo 316 (NY); Prov. Nuflo de Chavez, study Weddell (1849) also named this species Chrysoarea 

of BOLFOR project, Las Trancas-95, 1631 'S, 61 ?50'W, xylonfebrifugum nom. inval., using material he col- 

450 m, 20 Mar 1995 (fl), Mamani 436 (NY); Prov. Chiquitos, lected in Bolivia, and later (Weddell, 1854) validated 

Serrania de Sunsas, Tucavaca, 4 Jul 1995 (fr), Jardim & 

it by transferring it to Howardia. In the Paris her- 

Cadden 2153 (NY). TARIJA: Prov. O'Connor, San Sim6n, 

barium there is a 

20 km E of Entre Rios, 1300 m, 7 Feb 1937 (fl), West 8261 

specimen (Weddell 4190), collected 

(GH, MO, UC); rd. Tarija-Villa Montes, San Sim6n, 22 

in Bolivia 

May 

(Prov. Enquisivi), that bears on the attached 

1971 (fr), Krapovickas et al. 19143 (F, GH); Prov. Gran label (in Weddell's handwriting) the names "Chryso- 

Chaco, Sierra de Aguarague, 1200 m, 8 Jun 1977 (fl), xylonfebrifugum 

Krapovickas & Schinini 31024 (F[2], MO); Prov. Gran 

Chaco, Chaco, rd. Villa Montes-Jomiba, 500 m, Feb 1980 

(st), Rojas 1546 (UB); Villamontes, 1923, Pflanz 2053 (B). 

ARGENTINA. JUJUY: Depto. Ledesma, El Sauzal, 5 Jul 

1911 (fr), Lillo 10785 (F, GH, NY[2], UC, US); Depto. 

Ledesma, El Quemado, 4 Jun 1973 (fr), Turpe et al. 4690 (K, 

NY[2], US); Calilegua, 2 Jun 1943 (fr), Bartlett 20361 (GH, 

NY[2]); Depto. Ledesma, Parque Nacional Calilegua, 28 Jun 

1989 (fr), Kiesling 7109 (NY). SALTA: Urundel, 400 m, 18 

Jun 1941 (fr), Zabala 165 (F, GH, US); Depto. Orin, Campamento 

Rio Pescado, 16 Jan 1951 (fl), Scolnik 1678 (F[2]); 

Depto. Oran, Vespucio, 13 Jul 1937 (fr), Cabrera 4159 (F, 

NY[2]); Depto. Oran, Rio Pescado, 400 m, 1 Jun 1951 (fr), 

Sleumers.n. (B). TUCUMAN: Depto. Capital, Instituto Miguel 

Lillo, 450 m, 1 Apr 1974 (fl-fr), Borsini s.n. (CAS). 

Local names and uses. Peru: bouganvillea (Cuzco, 

Mexia 8035), flor de hoja (Cuzco, Mexia 8035). Bolivia: 

quina (Cardenas 2593; Peredo 525; Fosberg 28653; Nee 

& Coimbra 35199), sacha quina (Krapovickas et. al. 

19143). Argentina: cascarilla (Schultz 978), palo marfil 

(Lillo 10785), sacha quina (Borsini s.n.; Meyer 4811). 

Pogonopus tubulosus has been used by native 

people and European settlers as a secondary source 

of quinine to cure malaria and intermittent fevers, and 

reported by many as: "Bark used for quinine" (Bolivia, 

Gosling s.n., 3 Jun 1910), "Corteza febrifuga" 

(Argentina, Schultz 978), and "used as Cinchona bark" 

(Bolivia, Cdrdenas 2593). This species is also widely 

cultivated as an ornamental shrub, because of its bright 

red calycophylls: "Muy decorative por las grandes 

bracteas rosado-vivo" (Schultz 978). 

- Howardiafebrifuga Wedd," which 

is here selected as the lectotype of Howardiafebrifuga. 

Another specimen of Weddell 4190 is present 

at US, which is here treated as isolectotype. Another 

specimen (ex Paris Herbarium) collected by Weddell 

in Prov. Enquisivi is preserved at US, and this is a 

probably an isolectotype (its label lacks Weddell's 

collection number). Two other Weddell collections (ex 

Paris Herbarium) from "Prov. de la Cordillera, Bolivia" 

are at BR and K, and one (ex Paris Herbarium) 

from "Prov. de Santa-Cruz de la Sierra, Bolivia" is at 

F. These last three specimens were cited among the 

syntypes by Weddell (1849, 1854) and therefore are 

paratypes of H. febrifuga. 

Hooker (1873) transferred Howardiafebrifuga to 

Pogonopus. Hutchinson (1910) published P.febrifugus 

var. macrosema using specimens collected by Cecil 

Gosling in eastern Bolivia. One of the latter (Gosling 

s.n., 6 Mar 1910) was located at K, on which has been 

penciled (unknown handwriting): "P. febrifugus var. 

macrosema." This specimen is here selected as the lectotype 

of this variety. The same specimen also bears 

a label that has the annotation (in Hutchinson's handwriting): 

"Pogonopus amarus, Hutchinson sp. nov.," 

but this name was never published. 

Some herbarium specimens (Rusby 2090, G, P) 

were annotated by Britton as "Pogonopus bolivianus 

Britton, sp. nov.," but this specific name has never 

been published. Another specimen (Bridges s.n., BR- 

Martius Herbarium) was annotated by Schumann as 

Pogonopus tubulosus var. boliviensis, a varietal name 

that also remained unpublished. 

In the Richard Herbarium (P) a specimen was 

found with a label annotated (in A. Richard's handwriting): 

"Macrocnemum tubulosum A. Rich. - Bresil," 

which I consider to be the holotype of this species. 

Weddell (1854) renamed this species as Howardia 

richardi, a superfluous name based on the type of M. 

tubulosum. The holotype of M tubulosum annotated 

by A. Richard has two additional labels: one with the 

annotation (in Weddell's handwriting) "Howardia 

Richardi Wedd.," the other with morphological pencil-drawings 

and lightly penciled the name "Calycophyllum 

tubulosum." Because of the additional labels, 

this specimen preserved in the Richard herbarium 

is here selected also as the holotype for both names. 

Rejected Species 

Pogonopus erythroxylon (Standley) J. H. Kirkbride, 

Acta Amaz. 15: 54. 1985. Capirona erythroxylon 

Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 

22: 48. 1940, non Sickingia erythroxylon 

Willdenow, Gesellsch. Naturf. Freunde Neue Schr. 

3: 446. 1801. Non Simira erythroxylon 

(Willdenow) Bremekamp, Acta Bot. Neerl. 3:153. 

1954. Type. Ecuador. Surroundings of Guayaquil, 

"Vulgo Palo Colorado, F.H.D. 139.L495, Afio de 

1800," Tafalla s.n. (holotype, [F.H.D. 139.L495] 

MA; isotype, MA; photo-MA at F, TEX) = Simira 

ecuadorensis (Standley) Steyermark.


This species was originally described 

& 

by Standley 

Hooker, Gen. PI. 2: 45. 1873; Baillon, Hist. PI. 

(1940) as Capirona erythroxylon, citing two 

7: 397, 493. 

speci- 

1880; Poiret in Lamark, Encycl. Suppl. 

mens collected by Ruiz and Pavon in 1800 and 9: 

pre- 

230; Schumann in Martius, Fl. bras. 6(6): 259served 

in the Herbarium Peruvianum (MA). Both 260. 1889; Schumann in Engler & Prantl, Nat. 

"putative collectors" returned from South America to Pflanzenfam. 4(4): 18, fig. 6A-E. 1891; Urban, 

Madrid in 1788, leaving their collecting efforts under Symb. Ant. 410-411. 1900; Standley, N. Amer. 

the responsibility of Juan Jose Tafalla. Between 1799 Fl. 32: 5-6. 1918; Rizzini, Rev. Bras. Biol. 7(2): 

and 1803, Tafalla lead a collecting expedition in the 275-277. 1947; Bremekamp, Rec. Trav. Bot. 

forests Neerl. 31: 260-261. 

(now mostly destroyed) surrounding 

1934; Standley, Publ. Field 

Guayaquil, southern Ecuador (Estrella, 1991). It is Columbian Mus., Bot. Ser. 7: 203,370-371. 1931; 

most likely that the type specimens of C. erythroxylon Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 192. 

were collected in this area. Tafalla periodically sent 1936; Steyermark, Ceiba 3: 18-19. 1952; 

his own collections to Pavon, who filed them in the Steyermark, Mem. New York Bot. Gard. 12(3): 

Herbarium Peruvianum (MA). The type specimens of 178-186. 1965; Steyermark, Acta Bot. Venez. 8: 

C. erythroxylon should, therefore, be cited as collected 248-249. 1973; Steyermark in Lasser & 

by Tafalla in the vicinity of Guayaquil (Ecuador), and Steyermark, Fl. Venez. 9(3): 244-253. 1974; 

not collected by Ruiz and Pav6n in Peru (as reported 

Howard, Fl. Lesser Antilles 397. 1989; Dwyer, 

Ann. Missouri Bot. Gard. 67: 

by Kirkbride, 

82-84, 

1985a). 

fig. 18. 1980; 

The type specimens were annotated by Krause as Burger & Taylor in Burger, Fl. Costaric., 

Capirona and later described as C. erythroxylon by 

Fieldiana, Bot. n.s. 33: 94-95, fig. 37. 1993; 

Standley (1940). Kirkbride (1985a) transferred this Delprete, Brittonia 48: 35-44, fig 1. 1996; Delprete 

species to Pogonopus because of its "valvate aestiva- in Harling & Andersson, Fl. Ecuador (in press). 

tion and many small, ovules in each locule ... with 1999. Type species. Chimarrhis cymosa Jacquin. 

one calyx lobe sometimes expanded into a large col- Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio Janeiro 

ored foliar organ." I studied a detailed set of photos 4:177, pl. 23. 1925. Type species. Pseudochimarrhis 

of the two Madrid types taken by Kirkbride and con- turbinata (A. P. de Candolle) Ducke [= Chimarrhis 

cluded that C. erythroxylon matches the types of subgen. Pseudochimarrhis (Ducke) Delprete 

Sickingia ecuadorensis [Mille 884 (F)] (= Simira turbinata A. P. de Candolle]. 

ecuadorensis), which was collected in the forests of 

Guayaquil, and with which it is here reduced to syn- Trees, often tall canopy trees, bole irregular, with 

onymy. The flowers of the types have imbricate aes- or without buttresses; buttresses sometimes very large, 

tivation, but one photo of the set taken by Kirkbride especially in the Amazonian species; bark smooth to 

shows a few loose flowers much bigger than those on deeply fissured; wood yellow, very hard. Stipules 

the specimens, and with valvate aestivation. These interpetiolar, above leaf attachment, free or connate 

dubious flowers, certainly fragments of a mixed col- at base, adnate to the petioles, persistent or caducous, 

lection, were the ones that probably directed Kirkbride leaving a scar encircling the stem above the node. 

(1985a) to transfer this species to Pogonopus. The Leaves elliptic to ovate to obovate, usually acuminate 

common name "palo colorado" (= red wood), anno- at apex; not pellucid punctate; petioles often thickened 

tated on the Tafalla's specimen, also supports the at base but not pulvinate; domatia a tuft of hairs (tufttransfer 

of this species into Simira, which typically pit domatia in C. jamaicensis) or absent. Infloreshas 

red wood. 

cences subterminal (axillary on the terminal nodes), 

corymbose, with opposite to subopposite lateral 

branches terminating in cymules. Flowers 

protogynous, sessile to pedicellate, very fragrant; 

CHIMARRHIS 

hypanthium obconical to turbinate. Calyx extremely 

6. Chimarrhis Jacquin, Select. stirp. amer. hist. 61. reduced, truncate or with barely distinguishable lobes, 

1763; Chimarhis Rafinesque, orth. var., Ann. Gen. persistent; in few species (e.g., C. brevipes and C. 

Sci. Phys. 6: 84. 1820. Jacquin, Select. stirp. amer. gentryana) in some of the flowers one calyx lobe exhist. 

77-78. 1788; Jussieu, Mem. Mus. 6: 381. panded into a white to greenish white calycophyll. 

1820; de Candolle, Prodr. 4: 403. Sep 1830; A. Corolla funnelform, deeply lobed, with reflexed 

Richard, Mem. Rubiac. 278-280. Dec 1830; lobes, white, greenish white or cream-white; tube fun- 

Endlicher, Gen. P1. 553 (N. 3260). 1838; Meisner, nelform to ciatiform; with a ring of pubescent hairs 

PI. vase. gen. 158 (N. 114). 1838; Grisebach, Fl. inside on the tube or on the base of the lobes; lobes 

Br. W.-Ind. 323. 1861; J. D. Hooker in Bentham (4-)5(-6); aestivation narrowly imbricate, superfi-


cially resembling valvate aestivation. Stamens 5, al- rolla, followed by the opening of the stigmas (resemternate 

to the petals, totally or partially exserted; fila- bling two lips exserted beyond the corolla, Fig. 4J), 

ments attached on the upper part of the tube, slender, which then become receptive. Shortly after the corolla 

basally flattened, basally pilose to villous; anthers opens, the style is abscissed, the filaments elongate, 

elliptic, dorsifixed, dehiscing by lateral slits. Pollen and the anthers are exserted well above the corolla 

tricolporate, exine densely thick-reticulate. Style ex- (Fig. 4K), becoming functional and opening by latserted; 

style branches protruding above corolla before eral slits. The flowers of this genus are minute (a few 

anthesis, very short, rounded to ovate, reflexed at millimeters long), cream-white, and sweet-fragrant. 

maturity. Ovary 2-celled, turbinate to obovate, pla- The corollas of Chimarrhis have been described 

centation peltate on the septum; ovules many in each as having valvate aestivation, and for this reason this 

locule, vertically inserted; immature fruits green and genus was placed in the Condamineeae sensu Hooker. 

semi-carnose when fresh. Capsules woody, globose, Close observation of the corollas of Chimarrhis reoblong 

or obovoid; dehiscing septicidally, disk-locu- vealed that the aestivation is instead narrowly imbrilicidal 

dehiscence present in old capsules of some cate, superficially resembling valvate aestivation. 

species. Seeds suborbicular, with central hilum and More specifically, the tips of the lobes do not overlap 

narrow concentric wing, margin undulate. (valvate), but the lateral margins of the petals obviously 

do overlap (imbricate) in bud and in early stages 

Chimarrhis is unique in the Rubiaceae in its axil- of anthesis. 

lary-subterminal inflorescences, capsules with peltate Chimarrhis is often confused with Calycophyllum 

placentation, and minute seeds umbilically attached (Calycophylleae), but the latter has terminal infloresand 

with minute concentric wings. The seeds of at cences (vs. lateral subterminal), seeds with strongly 

least one species (C. barbata) have a tendency to de- bipolar wings, smooth thin bark exfoliating in long 

velop bipolar wings, resembling those of the tribes vertical strips, and white wood (vs. yellow in Chimar- 

Cinchoneae and Calicophylleae. All the species of rhis). Chimarrhis is also often confused with Bathysa 

Chimarrhis are trees, varying from mid-canopy to (Rondeletieae) and Simira (Rondeletieae), but the latupper-canopy 

levels. Mature individuals of the genus ter two differ from the previous in having terminal 

usually develop buttresses, especially those growing inflorescences and different seed morphology. Simira 

in the Amazon basin (see Figs. 68A, 71A). The bark differs from Chimarrhis in having larger seeds, horiof 

these trees is smooth and grayish in young indi- zontally inserted, laterally attached, with well-develviduals; 

and fibrous, deeply fissured, reddish, and oped lateral-orbicular wings, and reddish wood. 

falling off in small irregular pieces in mature individu- Bathysa is similar to Chimarrhis in having small 

als (see Figs. 67C,D, 69C,D, 76B). 

cream-white fragrant flowers, small fruits, minute 

The tree architecture of this genus has been stud- seeds, and yellowish wood; the previous differing 

ied in only a few species. Chimarrhis microcarpa from the latter in having floral aestivation strongly 

(Halle et al., 1978), C. glabriflora (pers. obs.), and imbricate, roundish petals with irregular margins (vs. 

C. hookeri (pers. obs.; Fig. 68A) display the narrowly triangular with entire margins in Chimar- 

"Aubreville Model (Terminalia branding)," whereas rhis), and horizontal seeds, laterally attached, and 

C. cymosa (Halle et al., 1978) develops the "Petit's minute lateral wing (vs. vertical seeds, ventrally attached, 

Model," according to the architectural classification minute concentric wing peltate to a central hilum). 

of Halle et al. (1978). 

After careful observation, I concluded that the 

Leafdomatia were found to be a reliable taxonomic species of chimarrhis can be divided into two easily 

character, variable within certain species or consis- recognizable and probably monophyletic groups, ustent 

within others (Fig. 58A-L; see Domatia, above). ing the persistence and general morphology of their 

Chimarrhisjamaicensis and C. cubensis can be rec- stipules, and their geographic distribution: subgen. 

ognized solely by their characteristic domatia (Fig. Chimarrhis, with readily caducous stipules; and 

58B,C), while in the species with glabrous blades (C. subgen. Pseudochimarrhis (Ducke) Delprete, with 

latifolia, C. brevipes, C. gentryana, and C. barbata) persistent stipules. The second subgenus is a new stadomatia 

were not found. 

tus for the genus Pseudochimarrhis Ducke (1925), 

Several species have been reported to be either which he separated from Chimarrhis and transferred 

heterostylous or dioecious (Grisebach, 1862; Urban, to the Cinchoneae. To the two species already included 

1899), but all Chimarrhis species have strongly in subgen. Pseudochimarrhis (C. turbinata and C. 

protogynous flowers, which are neither dioecious nor barbata) I add C. duckeana (described below), C. 

heterostylous. Flower anthesis in this genus starts with brevipes, and C. gentryana, all with persistent deltoid 

the protrusion of the style above the still-closed co- stipules. Subgenus Pseudochimarrhis occurs mostly

II ~~~~~AIPJ.~~~~~~~~~~~~~~~~~~ ..... 

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= 

C hooker (topright Nel & Zaruma 7021 NY bottomleft Spruce 4930 K isotype) H. C latfolia J... (Hallen 6302 MO) C brev 

~~~~~~~~~~~~~~'-. 

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'"""" ... ~~~'"-..'..;."': " : 

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.'. ': : .' : '~:. .~. . " - : I;:.~.--:..:'".-,'..- 

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FIG. 58. Leaf domatia in Chimarrhis, abaxial views and blade cross sections. A. C. parviflora (Harber & Bello 1 735, F). B. C. cu 

(Proctor 22591, DAV). D. C cymosa (Webster et a! 9205, DAV). E. C. microcarpa (Berti 241, K). F. C glabriflora (top-right Du 

(Vtisquez & Arevalo 8969, TEX). K. C barbata (Kuhlmann RB1 7384, US, isotype). L. C. turbinata (top-right Boom et a! 8756, N


in the lowlands of the Amazon basin, while the duced Sprucea (= Simira) and Sickingia (= Simira) 

subgen. Chimarrhis is distributed in the Antilles, in to synonymy under Chimarrhis. 

Central America, the Andean slopes, and western Schumann (1889, 1891) reported Chimarrhis to be 

Amazonian rain forests. 

a genus of three species, recognizing C. cymosa, C. tur- 

Chimarrhis is here recognized as a genus of 14 binata DC., and describing C. hookeri (Schumann, 

species (9 in subgen. Chimarrhis, and 5 in subgen. 1889) from Spruce 4930, previously pointed out by 

Pseudochimarrhis). 

Hooker (1873). Schumann (1889, 1891) placed 

Chimarrhis in the Condamineeae. 


Urban (1899) subdivided C. cymosa into three 

subspecies: subsp. genuina from the Lesser Antilles; 

Chimarrhis was founded by Jacquin (1763) with subsp.jamaicensis from material collected in Jamaica; 

the description of C. cymosa from Martinique. Jacquin 

reported it to be locally called "Bois de riviere," and 

and subsp. microcarpa from material collected in 

Cuba (Wright 1262). 

derived its Latin name from the Greek Xtjqrappy 

(chimarro = torrent), because it was originally encoun- 

Schumann and Krause (1908a) (four years after 

Schumann's death) described Chimarrhis dioica as a 

tered in the proximity of torrents. He also reported species exceptional to Chimarrhis in having dioecious 

Chimarrhis as having one seed per locule, an error that flowers. This species was later chosen by Steyermark 

was reproduced by Poiret (1811). 

(1963) as the type species of his new genus Dio- 

Achille Richard (1830) placed Chimarrhis cymosa icodendron. 

(as "Chimarrhis corymbosa [hort. var.] Jacquin") in Standley (1918) did not recognized Urban's (1899) 

synonymy under Macrocnemum, but admitted that the subspecies and reduced them to synonymy (along with 

genera were poorly known to him. He correctly noted Macrocnemum longifolium) under C. cymosa. In adthat 

the capsules of Chimarrhis are polyspermous and dition, he transferred Rustia ferruginea Standl. (= 

septicidal, as in M. candidissimum (= Calycophyllum Bathysa sp.) to Chimarrhis. Later, Standley (1926) 

candidissimum), M. jamaicense, and M. tetrandrum. described C. microcarpa based on specimens collected 

Because he erroneously reported C. cymosa as C. cor- in Trinidad, and C. pittieri (Standley, 1930b) based 

ymbosa, when including this species into Macrocnemum, on specimens collected in Venezuela. Standley 

the combination M. corymbosum (= Condaminea (1931c, 1936) merely listed C. dioica (= Dioicocorymbosa) 

already existed, and he had to establish dendron dioicum) and C. pittieri (= Bathysa pittieri), 

the new name M. longifolium (based on Jacquin's maintaining Chimarrhis in the Condamineeae (sensu 

collection from Martinique). 

Hooker). 

De Candolle (1830) recognized Chimarrhis as a Adolpho Ducke (1876-1958) was very influential 

good genus, placing Macrocnemum longifolium (and 

Chimarrhis corymbosa) under C. cymosa, and dein 

understanding the phenology and clarifying the 

taxonomy of the Amazonian species of Chimarrhis, 

scribing C. turbinata on the basis of material collected 

by Patris in Cayenne (French Guiana). He placed 

primarily due to his 50 years of accurate fieldwork in 

the Brazilian Amazon. He often collected one tree in 

Chimarrhis in the tribe Hedyotideae, subtribe flowering stage and returned to the same individual 

Rondeletiinae (as "Rondeletieae"). 

Joseph Dalton Hooker (1873) attributed four speseveral 

months later to collect its fruiting material, 

mounting both collections on the same herbarium 

cies to Chimarrhis: two from the West Indies (but only 

one species was described at that time!), the third 

species C. turbinata, and the fourth from a Spruce's 

collection (Spruce 4930) from Tarapoto, Peru, but 

without giving it a name. He placed Chimarrhis in his 

newly founded tribe Condamineeae, subtribe Condamineinae 

(as "Eucondamineae"). 

Baillon (1880) placed Chimarrhis in his "Cinchona 

Series" (or Cinchoneae). He reported it "exceptional 

sheet. Ducke (1922) separated Pseudochimarrhis 

(with C. turbinata as type species) from Chimarrhis, 

stating that the latter genus differed from the former 

in having narrower, longer flowers, and vertically 

arranged seeds, characters that would place it in the 

Cinchoneae. Three years later, Ducke (1925) added a 

second species to Pseudochimarrhis (P. barbata). 

Benoist (1920) described Bathysa difformis (= C. 

turbinata), which he later transferred to Pseudoin 

this group as it would be in any other in which it 

might be placed," later adding that its "stamens are 

of two kinds according as we examine this or that 

flower with filaments short or sometimes long" (probably 

suggesting dimorphic flowers, which do not exist 

in any Chimarrhis species), and erroneously rechimarrhis 

(Benoist, 1933). 

Bremekamp (1934) stated that the separation of 

Pseudochimarrhis from Chimarrhis, proposed by 

Ducke (1922, 1925) was unacceptable, reporting: 

"The seeds of C. cymosa Jacq., the type species of the 

genus, however are arranged in exactly the same way


as those of C. turbinata, and the longer and narrower duced Pseudochimarrhis to Chimarrhis and synonyflower 

is a character of so little importance that it is mized the other species accordingly. He maintained 

in itself not sufficient to justify the creation of a new Chimarrhis barbata and C. turbinata under Chimargenus. 

Pseudochimarrhis turbinata (DC.) Ducke is rhis; described C. brevipes and C. bathysoides [= Bathysa 

therefore referred back to Chimarrhis and P. barbata bathysoides ((Steyerm.) Delprete]; raised C. cymosa 

Ducke became Chimarrhis barbata (Ducke) Brem. subsp. jamaicensis Urb. and subsp. microcarpa to 

comb. n." In the same work Bremekamp described C. specific rank; maintained C. williamsii, C. hookeri, 

longistipulata, which is here treated as synonymous C. glabriflora, and C. parviflora as previously dewith 

C. microcarpa (see discussion under this spe- scribed; and divided C. microcarpa into var. microcies). 

Pseudochimarrhis difformis has never been carpa and var. speciosa (which are here raised to spepositioned 

in Chimarrhis, but this species is synony- cific rank). Finally, in the list of excluded species, 

mous with C. turbinata. 

Steyermark (1965) reduced C. dioica and C. venezuel- 

Rizzini (1947) treated Chimarrhis turbinata as a ensis to synonymy under Dioicodendron dioicum; 

dubious taxon, proposing C. duckei Rizz. (nom. treated C. decurrens as synonymous with Allenanthus 

superfl.) as a substitute name for Pseudochimarrhis erythrocarpus Standl.; transferred C. goudotii, C. pisoniturbinata 

(DC.) Ducke. 

aeformis, and C. pittieri to Sickingia [= Simira]; and 

Steyermark (1965) published the only monograph returned C. ferruginea to Rustia. 

of Chimarrhis, where he recognized 12 species, and Dwyer (1980) and Burger and Taylor (1993) recwith 

only the new species fully described. Following ognized Chimarrhis as a genus of 14 species ranging 

Bremekamp's (1934) observations, Steyermark re- from Costa Rica to South America and the West Indies. 

Key to the subgenera and species of Chimarrhis 

1. Stipules readily caducous before adjacent leaves expansion (Central America, Antilles, and South 

American Andes) ....................................................................................... subgen. Chimarrhis 

2. Capsular disks glabrous. 

3. Anthers 1-2 mm long (Antilles). 

4. Domatia absent or few hairs present; young leafy branchlets 3-4 mm thick; leaf blades 

7-11 x 3-5 cm, with 7-8 secondary veins each side; inflorescences 7-10 cm long; 

capsules 2-2.5 x 2 mm (Cuba) ............................................................ .................... 3. C. cubensis 

4. Domatia well-developed as tufts in most vein axils; young leafy branchlets 4-15 mm 

thick; leaf blades 10-47 x 5-20 cm, with 8-15 secondary veins each side; inflorescences 

>10 cm long; capsules 3-5 x 2.5-3.5 mm. 

5. Tuft-domatia without pit; leafy branchlets 10-15 mm thick, semi-succulent, costate; 

leaf blades 21-47 x 10-20 cm, with 11-15 secondary veins each side; inflorescences 

15-24 cm long; capsules 3-4 x 2.5-3 mm (Lesser Antilles) ............................................ 5. C. cymosa 

5. Tuft-domatia with pit; leafy branchlets 4-7 mm thick, woody, not costate; leaf blades 

(8-)10-16 x 5.5-8 cm, with 8-9 secondary veins each side; inflorescences (10-)12-22 

cm long; capsules 4-5 x 3-3.5 mm (Jamaica and Haiti) ............................................ 4. C. jamaicensis 

3. Anthers 0.5-0.7 mm long (Central and South America). 

6. Leafy branchlets 9-12 mm thick, succulent to semi-succulent, smooth (Colombia, Ecuador, 

Peru, and B razil) ................................................................................................................... 6. C. glabriflora 

6. Leafy branchlets 1.5-8 mm thick, woody, exfoliating in small longitudinal strips. 

7. Domatia absent; stipules 6-25(-30) mm long, 2-4 mm wide at base; inflorescences 

4-12 cm long; leaf blades 5-15(-20) x 4-7.5 cm, with length:width ratio 2:1 to 2.5:1; 

capsules obovoid to turbinate, acute at base (Nicaragua, Costa Rica, Panama) ......... 1. C. parviflora 

7. Tuft-domatia present at least in some axils; stipules 25-35 mm long, 5-7 mm wide at 

base; inflorescences 10-22 cm long; leaf blades (9-)14-24 x 6.5-12 cm, with length:width 

ratio 1.5:1 to 2:1; capsules subglobose, rounded at base (Venezuela, Trinidad and Tobago, 

the Guianas, Brazil). 

8. Corolla funnelform, 4-5 mm long; inflorescences very compactly branched (L/A = 

3:1); leafy branchlets 2.5-5(-6) mm thick (Venezuela, Trinidad and Tobago, the 

Guianas and Brazil) ........................................ ....................... 8. C. microcarpa 

8. Corolla campanulate, 1.6-2.4 mm long; inflorescences laxly branched (L/A = 2:1); 

leafy branchlets (4-)5-8 mm thick (Venezuela) ....................................................... 9. C. speciosa 

2. Capsular disks puberulent to pubescent.


9. Capsules 2-2.5 mm long; corollas 2.5-3.5 mm long (Ecuadorand Peru) ................................... 7. C. hookeri 

9. Capsules 4-5 mm long; corollas 4-5 mm long (Costa Rica) ........................................................ 2. C. latifolia 

1. Stipules persistent after adjacent leaves abscission (mostly Amazon basin) .......................... subgen. Pseudochimarrhis 

10. Calycophylls present at least in some of the flowers; leaf blades pandurate. 

11. Leaf blades with 5-6 secondary veins each side, length:width ratio 1:1 to 1.5:1; calycophylls 

obovate; inflorescence L/A = 2.5-3:1; basal portion of axis not branched ["peduncle"] (3.5-) 

6-9.5; capsules obovoid (Venezuela) ............ ..................................................................... 13. C. brevipes 

11. Leaf blades with 7-10 secondary veins each side, length:width ratio 1.5:1 to 1.7:1; calycophylls 

ovate-elliptic; inflorescence L/A = 1.5-2:1; basal portion of axis notbranched 4.5-7.5; capsules 

narrowly obconical to turbinate (Amazonian Ecuador and Peru, and W Brazil). ................... 14. C. gentryana 

10. Calycophylls absent (rarely present in 1-2 flowers per inflorescence); leaf blades elliptic, ovate, or 

obovate (not pandurate). 

12. Disk minutely puberulent, obviously exceeding the calyx (Brazil: Amazonas). 

13. Leaf blades 5.5-10(-14) x 4-7 cm; leafy branchlets thin (2-4 mm wide), woody; capsules 

w ith oblong-ovoid disks .......................................................................................................... 11. C. barbata 

13. Leaf blades >12.5 x 7 cm; leafy branchlet thick (4-6 mm wide), succulent; capsules with 

obtuse to rounded disks . ....................................... .................... 12. C. duckeana 

12. Disk pubescent to pilose, not exceeding the calyx (Surinam, Brazil: Amapa, Para, Amazonas). 

........................................................................................................................................................ 10 . C . tu rb in a ta 

CHIMARRHIS JACQUIN SUBGEN. 

CHIMARRHIS 

1. Chimarrhis parviflora Standley, Trop. Woods 11: 

26.1927. Type. Panama. Bocas del Toro: Changui- 

nola Valley, 1927 (fl), Cooper & Slater 120 (ho- 

lotype, US, N.1315890; isotypes, A, F, G, GH, 

NY). Figs. 9A, 17A,B, 58A, 59A-G, 61 

Trees to 30 m tall, 40-50(-60) cm dbh, with clear 

bole to 10 m, with small buttresses or flutes; bark soft, 

pale brown; wood yellow to golden-yellow, very brittle. 

Leafy branchlets 1.5-4 mm thick, terete to compressed, 

ochraceous, glabrous to appressed puberulent; older 

branches glabrate, terete, grayish to pale brown. 

Stipules free at base, contorted in bud, narrowly triangular, 

acuminate, appressed puberulent to gray-sericeous 

outside, glabrous with basal colleters inside, 6- 

branchlet, 4.5-12 x 3-7 cm; lateral branches 1-2 pairs, 

basal portion of axis not branched 2-7 cm long; rachis 

decussately compressed to subterete, reddish 

brown; L/A = 1.6-2.1; rachis and branches densely 

curled to appressed gray-puberulent; distal bracts 

(when present) deltoid, ca. 0.5 mm long; bracteoles 

subtending flowers absent. Flowers pedicellate; 

pedicels 0.5-2 mm long, glabrous to minute puberulent; 

hypanthium turbinate, 1-1.5 mm long, glabrous. 

Calyx extremely reduced with wavy margin or barely 

distinguishable lobes, to 0.5 mm long; lobes to 0.4 mm 

long, ovate to deltoid, ciliolate at apex. Corolla funnelform, 

deeply lobed, 2-4 mm long, white to creamwhite; 

tube short cylindrical, 1-1.5 x 1-1.5 mm, glabrous 

outside and inside; lobes (4-)5, 1.5-2 x 1-1.5 

mm, narrowly ovate, truncate at apex, glabrous outside, 

densely villous at medio-basal zone. Stamens 

(4-)5, exserted well above the corolla, equal, attached 

1-1.5 mm from the base of the tube; filaments 3-4 

mm long, terete, basally flattened, connate to throat, 

densely villous at base; anthers elliptic, 0.6-0.7 x 0.2- 

0.4 mm, dorsifixed at medial zone, reversed at matu- 

25(-30) x 2-4 mm, reddish with a grayish tinge, readily 

caducous, leaving a white scar encircling the stem, 0.5- 

1 mm wide. Leaves 6-17(-22) x 4-7.5 cm, L/W = 2:1 

to 2.5:1, elliptic to oblong-obovate, acute-decurrent 

(ca. 25?) at base, obtuse and mucronate at apex, the 

deltoid acumen 0.7-1 cm long; dark green above, pale rity, base rounded. Pollen exine irregularly foveolategreen 

below, foliaceous; drying olive-green to brown, reticulate. Style exserted, 2-4 mm long, as long as the 

stiff-chartaceous; glabrous above, glabrous to sparsely corolla, glabrous, minutely costate; style branches 

appressed puberulent below, hairs whitish to yellow- extruding as two lips above corolla before anthesis, 

ish; primary, secondary, and tertiary veins appressed widely ovate to widely oblong, 0.3-0.4 mm long, stigpuberulent 

below; secondary veins (5-)8-14 each side; matic surface microscopically (40x) papillose; ovatertiary 

veins starting subparallel and reticulate in the ries often infested by insects, developing into globose 

center; petioles 10-23 mm long, 1-2 mm thick, ter- galls containing minute yellow larvae. Capsules oboete, 

glabrescent to appressed minute-puberulent; void to turbinate, acute at base, apex rounded, 1.5domatia 

absent. Inflorescences usually 2 (rarely 3-4) per 2.5 x ca. 1.5 mm, dark brown without lenticels, glanode, 

corymbose with opposite to subopposite decus- brous throughout; disk loculicidal dehiscence present 

sate branches, densely cymose at distal end of each in old capsules. Seeds 0.5-0.88 x 0.38-0.5 mm.


C 

iH^HBRl IIf I 

I 

FIG. 59. Chimarrhis parviflora (A-F from Cooper & Slater 120, US, holotype; G from Jimenez 3418, F). A. Habit of 

inflorescences. B. Flower with exserted anthers. C. Variation in calyx lobes. D. Detail of corolla, inside view. E. Stamen. 

F. Apical portion of style. G. Mature capsules.


Distribution (see Fig. 61) and ecology. Primary 

and secondary rain forests, and respective remnants, 

from low to medium elevations (Bosque pluvial 

premontano), 100-900 m, of the Caribbean slopes 

(rarely of the Pacific slopes), usually in proximity of 

creeks and small streams, in alluvial soil or volcanic 

loam of Nicaragua, Costa Rica, and Panama. Flower- 

ing specimens were collected in March, April, May, 

June, and July. Fruiting specimens were collected in 

May, July, August, and September. 

Colorado Island, trail 30 m S of Creek 7, 13 Jul 1970 (fr), 

Croat 11244 (MO[2], VEN). VERAGUAS: Valley of Rio Dos 

Bocas, between Escuela Agricola Alto Piedra and Calovebora, 

NW of Santa Fe, 08?30'N, 81 ?10'W, 400-450 m, 31 

Aug 1974 (fr), Croat 27755 (MO). 

Local names and uses. Costa Rica: higueron 

(Hartshorn 1211), pejiballito (Dayton & Barbour 

3007). Panama: yema de huevo (Cooper 401), llema 

de huevo (Cooper & Slater 3/10708), man wood (Dunlap 

33). As in most species of this genus, C. parviflora is 

reported to be good timber wood. 

Representative specimens examined. NICARAGUA. 

ZELAYA: Cerro El Escobin, 11?33'N, 84?21,W, 30 Jul 1982 

(fr), Sandino 3412 (MO); Colonia Serrano, on trail to Colonia This species is easily distinguished from the other 

Yolaina, 100 m, 1 1?35'N, 84?22'W, 28 Jul 1982 (fl), Sandino Central American taxa by its minute capsules (Fig. 

3294 (MO); Kurimwuasito, 50 km SE of Rio Blanco, 15 Mar 59G) with glabrous disk, and its slender-woody leafy 

1984 (fl), Sandino 4842 (MO); Kurinwacito, 13?09'N, branchlets. It is commonly encountered as a tree with 

84?55'W, 100 m, 23 Mar 1984 (fl), Moreno 23868 (MO). soft flaky white bark, with wood quickly turning rust- 

Rio SAN JUAN: Buenos Aires, 1 km N of Sabalo, above Rio colored when cut (probably due to its alkaloid con- 

Sabalo, 1 102'N, 84?28'W, 70 m, 10 Sep 1985 (fr), Moreno 

tent); leaves thickly membranous, waxy dark green 

26317 (MO). 

and faintly shiny above; and 

COSTA RICA. ALAJUELA: 2 km N of Bijagua, trail 

vegetative buds lightto 

San Miguel, 400-500 m, 8 Jul 1976 (fr), Utley & Utley 5337 green to whitish, with readily caducous stipules. The 

(CAS, F); Reserva Biol6gica Monteverde, Rio Peias young ovaries are often infected by insects which 

Blancas, 10?20'N, 84?43'W, 850 m, 28 Mar 1987 (fl), Haber cause them to enlarge abnormally into globose galls 

& Bello 6837 (F, GH, MO, TEX); Upala, Colonia Libertad, (sometimes mistaken for mature fruits). 

Valley of Rio Cucaracho, 450 m, 6 May 1988 (fl), Herrera Chimarrhis parviflora is most similar to C. 

1976 (F, MO); rd. Canas-Upala, 13.8 km N ofBijagua, 100- microcarpa (from Venezuela and the Guianas), the 

150 m, 26 Jun 1976 (fl), Croat 36431 (MO); Cton. San former differing from the latter in having the termi- 

Carlos, Villa Guisada, 725 m, 14 Apr 1939 (fl), Smith 1890 nal nodes of the branchlets appressed-puberulent; leaf 

(A, F, MO, NY). CARTAGO: Turrialba, Instituto, 600 m, 1 veins ascending-pubescent (vs. glabrous), domatia 

Sep 1950 (fr), L&on 2143 (F); Turrialba, E of Pavones, 700 

absent (vs. 

m, 8 May 1972 

present; 

(fr), Holdridge 6723 (F). HEREDIA: Finca La 

Figs. 58A, 58E); corollas 2-4 mm 

Selva, Sabalo-Esquina trail, 10?26'N, 84001'W, 30 long (vs. 4-5 mm long; Figs. 59B-D, 71C,D); corolla 

May 

1974 (fl), Hartshorn 1500 (F, MO); Finca La Selva, OTS 

lobes acute at apex (vs. obtuse to truncate); infloresfield 

station, South Boundary trail, 100 m, 21 May 1985 cences with L/A = 

(fl), 

3:1 (vs. L/A = 2:1; Figs. 59A, 71B), 

Wilbur 37205 (CAS, F, K, MO, US) vic. of Puerto Viejo, and basal portion of axis not branched 2-7 cm long 

canal entering Rio Sarapiqui, 100-150 m, 11 Aug 1965 (fr), (vs. 7 cm or more). 

Jimenez 3418 (F, MO, NY, US); 1 km E of Cariblanco, 850 

m, 21 May 1973 (fl), Hartshorn 1211 (F). LIM6N: Cerro 

Coronel, E of Laguna Danto, 10?4 'N, 83?38'W, 60-100 m, 

15 Mar 1987 (st), Stevens et al. 24859 (MO); La 2. Chimarrhis latifolia 

Florida, 

Standley, Publ. Field 

250 m, 7 May 1943 (fl), Dayton & Barbour 3007 (MO, US). 

Columbian Mus., Bot. Ser. 4: 265. 1929. Type. 

PUNTARENAS: Osa Peninsula, Reserva Forestal Golfo Dulce, Panama. Chiriqui: Progreso, Jul-Aug 1927 (fr), 

Rancho quemado, 08?43'N, 83?35'W, 200-400 m, 28 May Cooper & Slater 260 (holotype, F, N.573,160; 

1988 (fl), Hammel et al. 16858 (F, MO). SAN JOSE: Parque photo-F at F, G, TEX, VEN; isotype, F). 

Nacional Braulio Carrillo, Estacion Carrillo, 500 m, 25 Jul 

Figs. 9B, 17C,D, 58H, 60A-F, 61 

1984 (fr), G6mez et al. 22944 (F, MO); Parque Nacional 

Braulio Carrillo, path beyond Rio Sucio, 400-500 m, May Trees to 30 m tall, 30 cm or more dbh, canopy trees 

1984 (fl), G6mez et al. 22770 (MO, NY, VEN). 

with long buttresses; wood yellow, heavy, very brittle. 

PANAMA. BocAS DEL TORO: Region of Almirante, Daytona 

Farm, Jan-Mar 1928 (fr), Cooper & Slater 401 

Leafy branchlets 4-9 mm thick, subterete to com- 

(A, F, 

K, US); Changuinola Valley, 1927, Cooper & Slater 3 pressed, ochraceous, glabrous; older branches terete, 

(F, 

US); Changuinola Valley, 1 Aug 1923 (fr), Dunlap 33 (F, 

grayish to pale brown. Stipules free at base, contorted 

G, US). COL6N: Along Rio Guanche, S of Portobelo in 

Hwy., bud, narrowly triangular, acuminate, glabrous with 

09030'N, 79?40'W, 50 m, I 1 Apr 1982 (fl), Knapp et al. 4586 ciliate margins outside, glabrous with basal colleters 

(LL). PANAMA: Barro Colorado Island, slope 300 m S of Bat inside, 20-35 x 5-13 mm, reddish brown, readily 

Cove, 16 Jul 1978 (fr), Foster 2877 (K, MO, US); Barro caducous, leaving a dark-brown scar encircling the


tr q 

,aW~~~~~~~~~~~~~~~~~~~~~, 

?- .I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

FIG. 60. Chimarrhis latifolia (A-E from Allen 6302, MO; F from Allen 5762, F). A. Habit of inflorescences. B. Flower 

with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style. F. Mature capsule.


FIG. 61. Distribution of Chimarrhis parvifolia (triangles) and C. latifolia (circles). 

stem, 0.5-1 mm wide. Leaves (13-)16-28 x (5-)7- 

11 cm, L/W = 1.5:1 to 2:1; broadly elliptic to ellipticobovate, 

acute-decurrent at base, obtuse and mucronate 

at apex, the deltoid acumen 0.5-1 cm long; dark 

green above, pale green below, foliaceous; drying 

brown above, olive-green below, chartaceous; glabrous 


glabrous, secondary veins 7-12 each side; tertiary 

veins starting subparallel and reticulate in the center; 

petioles 20-45 mm long, 1.3-1.8 mm thick, terete to 

adaxially concave, glabrous; domatia absent or rarely 

a few hairs at the axil. Inflorescences 2 per node, 

corymbose with opposite to subopposite decussate 

branches, densely cymose at distal end of each 

branchlet; 8-15 x 8-13 cm, lateral branches 1(-2) 

pairs, basal portion of axis not branched 2.5-7 cm 

long; rachis decussately compressed to subterete, reddish 

brown, rachis and branches glabrous to sparsely 

minute puberulent; distal bracts usually absent or deltoid, 

1-2 mm long, ciliolate; bracteoles subtending 

flowers absent. Flowers sessile to short-pedicellate; 

pedicels (when present) to 3 mm long, minute puberulent; 

hypanthium turbinate, 1.5-3 mm long, glabrous, 

reddish brown. Calyx extremely reduced with wavy 

margin or barely distinguishable lobes, 0.5-1 x 2-2.5 

mm; lobes (when present) to 2 mm long, shallowovate 

to shallow-triangular. Corolla funnelform, 

deeply lobed, 4-5 mm long, white to greenish white; 

tube short cylindrical, 1-2 x ca. 1.5 mm, glabrous 

outside and inside; lobes 5, 2.5-3.5 x 1-1.5 mm, oblong 

and rounded to truncate at apex, glabrous outside, 

densely villous at medio-basal zone. Stamens 

5, exserted well above the corolla, equal, attached 2.5- 

4 mm from the base of the tube; filaments 4-5 mm 

long, terete, basally flattened, connate to throat, 

densely white-villous at base; anthers elliptic, 0.7-1 

x 0.3-0.4 mm; dorsifixed at medial zone, reversed at 

maturity, base rounded. Pollen exine reticulate-foveolate. 

Style exserted, 3.5-5 mm long, as long as the 

corolla, glabrous, minutely costate; style branches 

extruding as two lips above corolla before anthesis, 

rounded, ca. 1 mm long, stigmatic surface microscopically 

(40x) papillose; ovary-galls not found. Capsules 

obovoid to oblong, acute at base, apex rounded to 

subtruncate, 4-5 x 3-3.5 mm, dark brown without 

lenticels, glabrous below the disk; disk black, densely 

minute-puberulent; disk loculicidal dehiscence present 

in old capsules. Seeds 0.49-0.78 x 0.33-0.58 mm.



secondary forests, forming stands of canopy trees, 

sometimes in secondary forests in association with 

Ochroma and Schizolobium, or in remnant patches of 

mature forests at low elevations (0-300 m) of the 

Pacific coast of Costa Rica and Panama. Flowering 

specimens were collected in October, and one speci- 

men (Jimenez & Zuniga 760) in late blooming-early 

fruiting was collected in December. Mature fruiting 

specimens were collected only in January. 

Field Mus. Nat. Hist., Bot. Ser. 11: 192. 1936. 

Chimarrhis cymosa Jacquin subsp. microcarpa 

Urban, Symb. Ant. 1: 411. 1899. Type. Cuba. 

Granma: Monte Verde, Jan-Jul 1859 (fl), Wright 

1262 (lectotype, G, here selected; isolectotypes, 

BR, G, HAC[5], K, P). 

Figs. 9C, 17E, 58B, 62A-F, 63 

Trees to 10-20(-30) m tall, without or with very 

small buttresses; bark light brown, soft; wood yellow, 

very hard. Leafy branchlets 3-4 mm thick, terete, 

Reproductive biology. The white, fragrant flowglabrous; 

older branches grayish to pale brown; leners 

are reported to be visited by bees and butterflies 

ticels not found. Stipules free at base, contorted in 

(Jimenez & Zuniga 760). 

bud, narrowly triangular, acuminate, glabrous outside, 

Specimens examined. COSTA RICA. PUNTARENAS: glabrous, with two basal rows of colleters inside, 15- 

Vic. of Jalaca Station, 30 m, 8 Oct 1951 (fl), Allen 6302 (DS, 25 x 5-8 mm, reddish brown, readily caducous, leav- 

F[2], GH, MO, US); Esquinas forest preserve, 0 m, 10 Jan ing a whitish scar encircling the stem, 0.5-1 mm wide. 

1951 (fr), Allen 5762 (DS, F, GH, US); Reserva Biol6gica Leaves (5-)8-12.5 

Carara, Lomas Pizote, Bijagual trail, 09?47'N, 84?35'W, 300 

m, 8 Dec 1989 (fl-fr), Jimenez & Zuniga 760 (CR, F, K[2], 

MO); vic. of Palmar Norte de Osa, Rio Terralba, 20 Aug 

1950 (st), Allen 5613 (F); Cant6n de Osa, Fila Retinto, Quebrada 

Benjamin, 08?59'N, 83?28'W, 50-200 m, 3 Jan 1990 

(fr), Hammel et al. 17719 (CR, F, MO); Cant6n de Osa, 

Peninsula de Osa, trail to Rancho Quemado, 08?40'N, 

83?31'W, 100m, 19Nov 1992 (fr),Zamoraetal. 1901 (NY); 

Corcovado National Park, Sirena Field Station, 08?30'N, 

83?35'W, 14 Aug 1984 (st), Gentry & OTS class 48572 

(MO). SAN Jost: Cerros de Turrubales, Lagunas, 09?50'N, 

84?31'W, 200 m, 15 Feb 1993 (fr), Morales et al. 1124 (NY). 

Local names and uses: Costa Rica: yema de huevo 

(Allen 6302). Panama: jagua amarilla (Cooper & Slater 

260). As in most other species, the wood of this species 

is used in construction. One label (Cooper & Slater 

260, holotype) bears the following comments: "Tree 

25 m, with high buttresses and yellow wood. Wood 

heavier and harder to split than 'Jagua blanca', grain 

very crooked and crossed, making it hard to split." 

This species is easily distinguishable from the other 

species present in Central America (C. parviflora) by 

its glabrous condition, larger leaves, larger capsules, 

and absence ofdomatia. It has been suggested (Burger 

& Taylor, 1993) that this species may be synonymous 

with C. cymosa (of the Lesser Antilles), but the latter 

differs in having succulent (with large spongy pith) 

costate (Fig. 65A), leafy branchlets (1-1.5 cm thick!), 

characteristic tuft-domatia (Fig. 58D), and much 

larger leaves. 

3. Chimarrhis cubensis Steyermark, Mem. New 

York Bot. Gard. 12(3): 184. 1965. Chimarrhis 

cymosa Jacquin var. microcarpa (Urban) Standley, 

x 3-5 cm, L/W = (2:1) 2.5:1; elliptic 

to elliptic-obovate, acute-decurrent at base, acute 

at apex, with a deltoid acumen ca. 7 mm long; dark 

green above, pale green below, semi-coriaceous; drying 

brown above, olive-green below, semi-coriaceous; 

glabrous above, sparsely yellow-pubescent below; 

primary and secondary veins erect pubescent to hirsutulous, 

secondary veins 7-8 each side; tertiary veins 

mainly reticulate (rarely subparallel); petioles 6-10 mm 

long, 1-1.5 mm thick, adaxially flattened, minutely 

winged, glabrous; domatia absent or rarely a few hairs 

at the axil. Inflorescences 2(-3) per node, corymbose 

with opposite decussate branches, densely cymose at 

distal end of each branchlet; 7-10 x 2.5-5 cm, one 

pair of lateral branches, basal portion of axis not 

branched 5.5-7.5 cm long; rachis decussately compressed 

to subterete, reddish brown, rachis and 

branches yellow short-puberulent; distal bracts absent 

or reduced to deltoid scales 0.5-1 mm long; bracteoles 

subtending flowers absent. Flowers sessile to shortpedicellate; 

pedicels (when present) to 1.5 mm long, 

minute puberulent; hypanthium turbinate, 1-1.5 mm 

long, glabrous, reddish brown. Calyx extremely reduced 

with wavy margin (rarely with 5 minute teeth), 

0.5 x 1.5 mm. Corolla funnelform, deeply lobed, 4- 

5 mm long, cream-white; tube short cylindrical, 1.5- 

2 x ca. 1.5 mm, glabrous outside and inside; lobes 5, 

2-3 x ca. 1 mm, oblong-ovate, rounded to truncate at 

apex, glabrous outside, white-villous at medial zone, 

the remaining basal and distal zones glabrous inside. 

Stamens 5, exserted well above the corolla, equal, 

attached ca. 2 mm from the base of the tube; filaments 

3-4 mm long, terete, basally flattened, connate to 

throat, densely white-villous at base; anthers narrowly 

elliptic, rectilinear, 1.5-2 x ca. 0.3 mm, dorsifixed 

near the base, reversed at maturity, base rounded, truncate 

apiculate at apex. Pollen exine foveolate-reticu-


FIG. 62. Chimarrhis cubensis (A-E from Wright 1262, BR, isotype; F from Wright 1622, BR, paratype). A. Habit of 

inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style. 

F. Mature capsule.


s0 70 so 

FIG. 63. Distribution of Chimarrhis cubensis (circles), 

* 0^0 

. . -cymosa (triangles). 

/ Tc.r.c, V? ^i 

FIG. 63. Distribution of Chimarrhis cubensis (circles), C. jamaicensis (squares) and C. cymosa (triangles). 

late. Style exserted, 3.5-5 mm long, as long as the 

corolla, glabrous, minutely costate; style branches 

extruding as two lips above the corolla before anthe- 

sis, rounded, ca. 0.5 mm wide, stigmatic surface mi- 

croscopically (40x) papillose; ovary-galls not found. 

Capsules globose to subglobose, obtuse at base, apex 

shallowly rounded, 2-2.5 x ca. 2 mm, dark brown 

without lenticels, glabrous throughout; disk black; 

disk loculicidal dehiscence present in old capsules. 

Seeds 0.58-0.87 x 0.48-0.77 mm, more or less un- 

gulate, yellow. 


secondary seasonal forests (pluvisilvas de montafia), 

in acid soils, on low mountain ranges, 700-1000 m. 

Endemic to Cuba. Flowering specimens were col- 

lected in January, February, and May. Fruiting speci- 

mens were collected in July and August. 

Conservation status. This species is known from 

relatively few collections and appears to be endan- 

gered. Some attempt should be made to protect the 

localities where it occurs. 

Specimens examined. CUBA. GRANMA: Nuevo Mundo, 

vic. of Rio Macaguanigua, 13 Jan 1972 (fl), Del Risco 27343 

(HAC); Rio Jaguani, 3 Feb 1973 (fl), Del Risco 27395 

(HAC); Sierra Maestra, Salida Cala, 17 May 1948, Acuna 

s.n. (HAC[2]); Toa, 19 Mar 1952, Smith 396 (HAC); Sierra 

Maestra, vic. of Sevilla, 800-900 m, 18 Jul 1918 (st), Ekman 

9417 (G). GUANTANAMO: Finca La Prenda, Jul 1922, 

Hioram & Maurel 6066 (HAC, NY); Camino La Melba, km 

26,31 Mar 1972, Del Risco 27458 (HAC). HOLGUIN: Sierra 

de Nipe, Loma Colorado, vic. of Rio Piloto, 10 Jul 1919 (fl), 

Ekman 9692 (G, NY). PINAR DEL Rio: Banks of Rio Santa 

Cruz, 1861, Wright 1622 (BR, G[2], HAC[2]); San Crist6bal, 

6 Dec 1915, Roig 6419 (HAC); San Diego, Finca Cortina, 

Mar 1910, Fors 142 (HAC); Valley of Taco-taco River, 

Rangel, Aug 1926, Le6n 12813 (HAC, NY). SANTIAGO DE 

CUBA: Pico Turquino, Loma del Sabici, 1000 m, Jul 1922, 

Le6n 11011 (HAC, NY); Loma del Gato, Cobre Range of 

Sierra Maestra, 900 m, Jul-Aug 1921, Le6n et al. 9941 

(HAC, NY). 

Local names and uses: Cuba: jilacho (Ekman 

9692), hilacho (see below), penda (pers. comm.), roble 

amarillo (pers. comm.), cera (see below). Roig and 

Mesa (1928) report the following (free translation 

from the original): "Cera: so it is called a tree that


grows in the mountains of San Cristobal, from which dark green above, pale green below, semi-coriaceous; 

is extracted a hard and valuable wood, of the same drying brown above, olive-green below, semi-coriaintense 

yellow as wax [cera]" and "Hilacho: a name ceous; glabrous above and below; primary and secgiven 

to a tree of the Sierra Maestra and of the family ondary veins glabrous, secondary veins 8-9 each side; 

Rubiaceae [C. cubensis]." 

tertiary veins starting subparallel and reticulate in the 

center; petioles 10-25 mm long, 1.5-2 mm thick, 

Of the three species of Chimarrhis endemic to the subterete to adaxially concave, minutely winged, gla- 

Caribbean Islands, this is distinct in having smaller brous; tuft-pit domatia often present, with a tuft of 

leaves, which are yellow erect-pubescent below. The densely yellowish-sericeous hairs obscuring the pit 

petioles and central veins of this species are minutely just below surface, randomly present at axils of secwinged, 

forming "pocket-domatia" in the axils of secondary and tertiary veins (Fig. 58C). Inflorescences 

ondary veins, never forming tufted domatia. Because 2(-3) per node, corymbose with opposite decussate 

of this I treat this taxon as separate, as did Steyermark branches, tertiary branchlets opposite, densely cymose 

(1965). 

at distal end of each branchlet; (10-)12-22 

Grisebach (1862) identified Wright 1262 and 1622 

(both from Monte Verde, Cuba) as C. cymosa Jacq.; 

the former collection is here selected as the lectotype 

of C. cubensis. Grisebach (1862) reported the flowers 

of these specimens to be "dioecious-dimorphic, 

either with exserted stamens and included stigma, or 

exserted style and included anthers." Barely reproducing 

Wright's specimen labels, Urban (1899) reported 

the flowers of this species as "floribus heterostylis (ex 

Wr.)." What were reported as dioecious-dimorphic 

(Grisebach, 1862) or heterostylous (Urban, 1899) 

flowers are instead two successive stages of anthesis. 

4. Chimarrhis jamaicensis (Urban) Steyermark, 

Mem. New York Bot. Gard. 12: 184. 1965. 

Chimarrhis cymosa Jacquin var.jamaicensis (Urban) 

Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 

192. 1936. Chimarrhis cymosa Jacquin subsp. 

jamaicensis Urban, Symb. Ant. 1: 411. 1899. 

Type. Jamaica. Saint James: Mount James, 330 

m, 25 Jul 1895 (fr), Harris 5810 (lectotype G, here 

selected). Figs. 17F, 58C, 63, 64A-F 

Trees to 18 m tall, single-stemmed trees profusely 

branching, with small buttresses; bark pale brown. 

Leafy branchlets 4-7 mm thick, woody (not succulent), 

terete (not costate), pale brown, glabrous; older 

branches grayish to pale brown; lenticels not found. 


with many parallel veins departing from the 

base, glabrous outside and inside, with sparse basal 

colleters inside, 20-25 x 8-9 mm, reddish brown, 

readily caducous, leaving a pale brown scar encircling 

the stem, ca. 0.5 mm wide and very evident in older 

branches. Leaves (9-)1 1-18.5 x 5.5-8 cm, L/W = 2:1, 

mostly obovate, acute-decurrent at base, obtuse and 

mucronate at apex, deltoid acumen ca. 5 mm long; 

x (6-)9- 

11 cm, 1-2 pairs of lateral branches, basal portion of 

axis not branched (5-)6.5-11 cm long; rachis 

decussately compressed to subterete, reddish brown, 

rachis and branches yellow short-puberulent; distal 

bracts absent, reduced to a line just below the branching 

point, or minute deltoid scales to 1 mm long; 

bracteoles subtending flowers absent. Flowers sessile 

to short-pedicellate; pedicels to 2 mm long, glabrous 

to sparsely minute-puberulent; hypanthium turbinate, 

1-1.5 x ca. 1.5 mm, glabrous to sparsely puberulent. 

Calyx reduced, truncate or with wavy margin, 0.5-1 

x 1.5-2.5 mm, glabrous. Corolla funnelform, deeply 

lobed, 4-5(-6) mm long, white; tube short cylindrical, 

ca. 1 x 2-2.5 mm, glabrous outside and inside; 

lobes 5, 2-3.5 x 1-1.5 mm, oblong-ovate, rounded to 

truncate at apex, glabrous outside, white-villous at 

medial zone, the remaining basal and distal portions 

glabrous inside. Stamens 5, exserted well above the 

corolla, equal, attached 1.5-2 mm from the base of 

the tube; filaments 4-4.5 mm long, terete, basally flattened, 

connate to throat, densely white-villous at base; 

anthers narrowly elliptic, rectilinear, 1.5-2 x ca. 0.3 mm, 

dorsifixed near the base, reversed at maturity, base 

rounded, with a membranaceous truncate ligula at 

apex. Pollen exine irregularly reticulate-foveolate. 

Style exserted, 4-5 mm long, as long as the corolla, 

glabrous, minutely costate (to winged) toward distal 

portion; style branches extruding as two lips above 

the corolla before anthesis, rounded to widely elliptic, 

ca. 0.5 mm wide, stigmatic surface microscopically 


obovoid to short-obovoid, obtuse at base, shallowly 

obtuse at apex, 4-5 x 3-3.5 mm, dark brown without 

lenticels, glabrous, brown below the disk; disk glabrous, 

black; disk loculicidal dehiscence present in old 

capsules. Seeds not seen. 

Distribution (Fig. 63) and ecology. Rain forests 

on limestone or bauxitic soils, on mountain slopes and


FIG. 64. Chimarrhisjamaicensis (A-E from Proctor 19818, LL; F from Harris 5810, G). A. Habit of inflorescences. 

B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style. F. Mature capsule.


low hills, 300-1400 m, of Jamaica and Hispaniola 

(Haiti). Flowering specimens were collected in March, 

June, July, August, and October. The single fruiting 

specimen known has been collected in July. 

Conservation status. This species is in danger of 

extinction, since it is known only from the heavily 

disturbed Massif de la Hotte (Haiti) and on the much 

decimated mountain slopes of Jamaica. Its habitat is 

in urgent need of protection. 

Champ Flore, on mtns. at low elev., Nov 1867 (fl- 

fr), Hahn 429 (neotype, P, here selected; isoneo- 

types, BR, G[5], K[2]). 

Figs. 30, 9D, 17G,H, 58D, 63, 65A-F 

Macrocnemum longifolium A. Richard, Mem. Fam. 

Rubiacees 199. Dec 1830 (reimp. Mem. Soc. Hist. 

Nat. Paris. ser. 3, 5: 279. 1834). Chimarrhis corymbosa 

A. Richard, orth. var. [of C. cymosa], Mem. 

Fam. Rubiace6s 199. Dec 1830. Type. Martinique. 

Locality unknown, 1820, A. Plee s.n. (lectotype, P, 

here selected). 

Specimens examined. JAMAICA. CLARENDON: Along 

trail between Sandy River and John Vale Forest Reserve, 700 Trees to 10 m tall, to 30 cm dbh; single-stemmed 

m, 12 Oct 1959 (fl), Proctor 19818 (DAV, LL, NY). PORT- trees, with slender trunk; bark pale brown; hardwood 

LAND: John Crow Mtns., 1.6 km S of Ecclesdown, 400 m, yellowish. Leafy branchlets 10-15 mm thick, mark- 

17 Jun 1959 (fl), Webster et al. 8341 (DAV[2], G, RB), edly costate, herbaceous, semi-succulent, green to pale 

Proctor 19742 (HAC, NY); Youthanside, 13 Jul 1908 (fl), 

brown, with a wide 

Moore 10648 (NY); near Troy, 650 m, Aug 1904 (fr) 8783 

pith, glabrous; older branches 

(NY), 16 Sep 1906 (fl) Harris 9449 (NY); Troy Valley, 13grayish 

to pale brown; lenticels not found. Stipules 

18 Sep 1906 (fr), Britton 610 (NY); Silver Hill, 1260 m, 5 

free at base, contorted in bud, narrowly triangular, 

Aug 1895 (fl-fr), Harris 5828 (NY); Bachelor's Hall, near acuminate, with many parallel veins departing from 

Bath, 300 m, 17 Sep 1908 (fr), Harris & Britton 105 76 (NY); base, glabrous outside, with a band of sericeous hairs 

Rodney Hall, 350 m, 28 Mar 1961 (fl), Adams 9289 (M). at the margins, glabrous and with sparse basal colleters 

SAINT ANN: Chippenham Park, rd. Moneague-Brown's inside, 40-60 x 12-15 mm, reddish brown, readily 

Town, 26 Jul 1850 (fl), Lemann 473 (K). SAINT JAMES: Gar- caducous, leaving a pale brown scar encircling the 

lands distr., 4-5 km SSW of Flamstead Gardens, 700 m, 7 stem, ca. 0.5 mm wide and very evident in older 

Aug 1962 (fl), Proctor 22591 (DAV, LL). WEST MORELAND: branches. Leaves 22-49 x 10.5-20 cm, L/W = 2:1, 

Copse Mt. Woods, 1.6 km SW of Rat Trap, 400-450 m, 23 

obovate, acute-decurrent at base, obtuse and mucr- 

Oct 1960 (fr), Proctor 21467 (NY). 

onate at 

HAITI. Massif de la Hotte, western group, Jeremie, 

apex, deltoid acumen 5-15 mm long; dark 

More Pain-de-Sucre, 1400 m, 22 Jul 1928 (fl), Ekman green above, pale green below, thick-foliaceous; dry- 

10398 (G, K); Fairfield, 1849 (fl-fr), Wullschaegel 1344bis ing brown above, olive-green below, stiff-charta- 

(M[2]). 

ceous; glabrous above and below; primary and secondary 

veins glabrous, secondary veins (11-)12-15 

Local name and use. Jamaica: wild fiddle wood each side; tertiary veins starting subparallel and re- 

(Adams 9289). This species has been reported to be a ticulate in the center; petioles 7-12(-20) mm long, (2good 

construction wood. 

)3.5-5 mm thick, subterete to adaxially concave and 

minutely winged, glabrous; tuft-domatia (not pitted) 

This species is unique in the genus in having leaf often present, but only at axils of primary with secblades 

with very distinct tuft-pit domatia (Fig. 58C) ondary veins, the hairs of the domatia departing from 

in the axils of secondary and sometimes tertiary veins. the veins at the axil zone, not from the blade (Fig. 

Chimarrhis jamaicensis is similar to C. cubensis in 58D). Inflorescences 2 per node, corymbose with 

having narrowly elliptic anthers 1.5-2 mm long, but opposite decussate branches, laxly cymose at distal 

differs in having much larger inflorescence, larger leaf end of each branch, tertiary branchlets alternate to 

blades with tuft-pit domatia (vs. "pocket-domatia"), subopposite; 15-24 x (7-)11-17 cm, (1-)2 pairs of 

and obovoid-oblong capsules (vs. globose to sub- lateral branches, basal portion of axis not branched 

globose). 

(5-)8-14 cm long; rachis decussately compressed, 

brown, rachis and lateral branches glabrous, terminal 

branchlets yellow short-puberulent (curled hairs); distal 

bracts absent or reduced to deltoid scales up to 2 

5. Chimarrhis cymosa Jacquin, Select. stirp. amer. mm long; bracteoles subtending flowers absent. Flowhist. 

61. 1763. Chimarrhis cymosa Jacquin subsp. ers sessile to short-pedicellate; pedicels (when 

genuina Urban, nom. inval., Symb. Ant. 1: 410. present) to 1.5 mm long, sparsely minute-puberulent; 

1899. Chimarrhis cyrrhosa Stend., Nom. ed. 2. ii. hypanthium turbinate, 1-1.5 x 1.5-2 mm, glabrous to 

88, in syn., orth. var. [ofC. cymosa]. Type. Martinique. sparsely puberulent. Calyx reduced, truncate or with


r~~~~ 

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?' . . 

?? 

?? o 

, 

\ 

ii 

... 

?~~~~~~~ B 

F'IG. 65. Chimarrhis cymosa (A and F fTom Wilbur el al. 813 7, LL; B-E from Webster 134 78, DAV). A. Habit of inflo- 

men. E. Apical portion of style. F. Mature capsule.


wavy margin, ca. 0.5 x 2-2.5 mm, glabrous. Corolla 

funnelform, deeply lobed, 3.5-5 mm long, white; tube 

short-cylindrical, ca. 1 x 1.5-2 mm, glabrous outside 

and inside; lobes 5, 3-4 x 1-1.5 mm, narrowly ob- 

long, rounded at apex, glabrous outside; densely 

white-villous at basal-medial zone, the distal portion 

glabrous inside. Stamens 5, exserted well above the 

corolla, equal, attached ca. 1 mm from the base of the 

tube; filaments 4-6 mm long, terete, basally flattened, 

connate to throat, with the basal 2 mm densely white- 

villous; anthers elliptic, ca. 1 x 0.3-0.4 mm, dorsifixed 

near the base, base rounded, apex acute, microscopi- 

cally (40x) mucronate. Pollen exine reticulate 

(densely reticulate-foveolate at poles). Style exserted, 

3.5-5 mm long, as long as the corolla, glabrous, mi- 

nutely costate (to winged) toward distal portion; style 

branches extruding as two lips above the corolla be- 

fore anthesis, rounded to oblate, ca. 0.3 mm wide, 

stigmatic surface microscopically (40x) papillose; 

ovary-galls not found. Capsules globose, obtuse at 

base, apex shallowly obtuse, 3-4 x 2.5-3 mm, dark 

brown without lenticels, glabrous to sparsely short- 

puberulent below the disk; disk black, glabrous, ob- 

viously exceeding the calyx; disk loculicidal dehis- 

cence present in old capsules. Seeds 0.56-0.78 x 

0.56-0.67 mm. 

Distribution (Fig. 63) and ecology. Rain forests, 

cloud forests, and their disturbed edges, on mountain 

slopes, 450-650 m, in the islands of the Lesser Antilles 

(Antigua, Guadeloupe, Dominica, Martinique, St. 

Lucia, and St. Vincent). On one specimen label (Duss 

345) it was noted that the species in Guadeloupe flow- 

ers from March to July. Flowering specimens were 

collected in February, March, and June, A single 

specimen (from Martinique) was collected in late 

blooming-early fruiting at the end of July. Fruiting 

specimens were collected in January, August, and 

October. 

Representative specimens examined. WINDWARD 

ISLANDS. GUADELOUPE: Close to river, 28 Feb 1895 (fl), 

Duss 345 (P); Ermitage a Trois Rivieres, 450 m, 29 Sep 1936 

(fr), Stehle 3015 (NY, P); 1944-46, Bena 1066 (P); Camp- 

Jacob, Trois Rivieres, 1892 (fl), Duss 2548 (NY), 1896 (fl), 

3424 (NY); Basse Terre, Col des Mamelles, 600 m, 26 Oct 

1990 (fr), Billiet & Jadin 5010 (BR); 1824, Perrottet 283 

(G). DOMINICA: St. Paul, ca. 1.6 km NE of Pont Casse on rd. 

to Rosalie, 650 m, 23 Jun 1965 (fl), Webster 13478 (DAV); 

Morne Colla Anglais, Sylvania, 610-732 m, 10-23 Aug 

1938 (fl), Hodge 695 (NY[2]); St. Joseph Parish, 9 km N of 

St. Joseph area of owner of Castaways, 15?24'N, 61?26'W, 

521 m, 23 Aug 1992 (fl), Lee 49 (NY[2]); 1 km SW of Millet, 

on the Mittel River, 20 m, 22 May 1984 (St), Slane 140 

(NY); St. David Parish, Imperial Hwy., near Pont Cass6, vic. 

of Grete Palmiste, ca. 560 m, 14 Mar 1993 (fr), Hill 24770 

(NY); St. David Parish, Bois Diable, 3 km E of Point Casse, 

on rd. to Rosalie, 15?22'N, 61 ?18'W, 24 Apr 1992 (fl), Tuxill 

et al. 63 (NY); 3 km from Pont Casse on rd. to Rosalie, 1 

Aug 1964 (fr), Wilbur et al. 8137 (CAS, LL); 30 Mar 1916 

(fl), Fishlock H338-16 (K); St. Joseph, Roche d'Or Estate, 

NE of Salisbury, 650 m, 1 Aug 1966 (fl), Stern & Wasshausen 

2582 (B). MARTINIQUE: Camp Flore area, 13-18 Mar 

1979 (fr), Howard & Howard 18846(NY, US); Borde de Miere, 

trail to St. Pierre, 150 m, 13 Jul 1936 (fl), Stehle 987 (NY); 

Gros Morne, 480 m, 11 Sep 1937, Stehle 2298 (US); Le Mont 

Vert, 21 Jul 1942 (fl), Stehle 5068 (US); Mome Calabasse 

area, 3 km NE of More Rouge toward Ajoupa-Bouillon, 

500-600 m, 23 Jul 1959 (fl-fr), Webster et al. 9205 (DAV[2]); 

1857, Belanger 195 (G), 196 (G); Martinica, Siebers.n. (M); 

Morne Rouge, Boullon, Duss 1452 (NY), 1880 (NY), 1883 

(NY); Camp-Jacol, Gompagne, Tres-Rivieres, Duss 2548 

(NY), 3424 (NY). ST. LUCIA: Matouba, 17-27 Apr 1979 (fl), 

Howard & Howard 19487; vic. of Mt. Gimie, 500 m, 2 Aug 

1959 (fr), Webster et al. 9403 (DAV); Canaries River approaching 

Mome Gimie, 28 Jan 1985 (fr), Howard et al. 

19930 (G, NY); St. Jacques, 8 Oct 1888 (fl), Ramage s.n. 

(K). ST. VINCENT: Between Three Rivers and Silver Spoon, 

1-7 Apr 1950 (fr), Howard 11143 (NY); Smith & Smith 889 

(NY[2]), Mar 1890 (fr), 1245 (K); Anderson s.n. (K). 

Local names and uses. Guadeloupe: r6solu de 

riviere (Duss 345), resolu de montagne, bois des bamsjaunes. 

Dominica: bois riviere (Fishlock H338-16), 

quinoyer. Martinique: bois de riviere (Hahn 429), bois 

riviere r6solu (Stehle 6626), river wood (Howard 

18846). St. Vincent: waterwood. On the specimen 

Duss 345 (Guadeloupe) it is reported that this species 

gives a much sought-after construction wood. 

This species is unique in the genus in having very 

thick, semi-succulent, strongly costate leafy branch- 

lets (Fig. 65A), with large spongy central pith, which 

makes the terminal branchlets very brittle (pers. obs.). 

The domatia in C. cymosa are commonly a sparse tuft 

of hairs attached on the central and secondary veins 

(Fig. 58D), and not on the blade itself (where mites 

and scale-insects are usually found). 

This is the type species of the genus, and is based 

on specimens collected by Jacquin in Martinique, in 

proximity of a river. No specimens collected by 

Jacquin, however, could be found (not at BM, not at 

P, not at W, LE?). The neotype should be selected 

from among the specimens listed by Urban (1899) 

under C. cymosa subsp. genuina (an invalid subspe- 

cific name, by which Urban referred to the "genuine" 

C. cymosa from the Lesser antilles). The best candi- 

date is Hahn 429 because it was collected in 

Martinique, fits the original description by Jacquin, 

and it has both flowering and fruiting material. The 

specimen of Hahn 429 preserved at P is here selected


as the neotype of C. cymosa and as lectotype of C. 

cymosa subsp. genuina until material from LE is examined 

or other possible type material is found. 

Jacquin (1763), in the original description, erroneously 

reported that the capsules of this species have 

one seed per locule. Achille Richard (1830) noted this 

error and treated Chimarrhis (which he erroneously 

reported as "Chimarrhis corymbosa") as synonymous 

with Macrocnemum. Richard (1830) established 

Macrocnemum longifolium (= C. cymosa) using material 

collected in Martinique (not Jacquin's types), 

but without accurate description and specimen citation. 

One specimen has been found at P annotated as: 

"Le Bois de Reviere - (Chimarrhis cymosa) - 

Martinique 1820 - A. Plee." Because of its collection 

locality, and the possibility that it was studied by A. 

Richard, I have selected this specimen as lectotype of 

M. longifolium. 

Urban (1899), the first worker to recognize the 

existence of three taxa on the Caribbean Islands, called 

this species C. cymosa subsp. genuina, because 

Jacquin's description was based on material collected 

in the Lesser Antilles. 

6. Chimarrhis glabriflora Ducke, Bol. Tecn. Inst. 

Agron. N. 4: 26. 1945. Type. Brazil. Amazonas: 

Esperanca, Rio Solim6es, at mouth of Rio Javari, 

"non rara in silva non inundabili," 15 Mar 1944 (fl), 

Ducke 1618 (holotype, R; isotypes, A, F, NY, 

US[2]). Figs. 3P, 4J,K, 9E,F, 58F, 66A-G, 

67A-D, 70, 77A 

Trees to 35 m (one individual reported 55 m tall, 

Bensman 331), to 50 cm dbh, much-branched at 

crown, with buttresses to 3 m high, extending several 

meters from trunk; bark red-brown, fibrous, soft, longitudinally 

fissured, brown-cream striate; inner bark 

pink, striate; wood yellow, very hard. Leafy branchlets 

decussately compressed, 3-4 mm thick, 9-12 mm 

wide', succulent to semi-succulent, subterete to 

decussately compressed, glabrous, youngest 2-3 

nodes sparsely to densely minute-puberulent; older 

branches glabrate, grayish; lenticels not found. 


acuminate, sparsely minute-puberulent outside, 

glabrous with sparse small colleters at base inside, 

(18-)20-53 x 5-11 mm, with many parallel veins 

departing from base, reddish green, readily caducous, 

leaving a white scar encircling the stem 1-2 mm wide. 

Leaves (15-)18-28 x long; dark green and shiny above, pale green-yellowish 

below, semi-coriaceous; drying rust-brown above, 

olive-green below, stiff-chartaceous; glabrous above 

and below; primary and secondary veins glabrous (to 

minute-puberulent), prominent below, secondary 

veins (10-)12-16 each side; tertiary veins starting subparallel 

and reticulate in the middle; petioles (10-)18- 

55 mm long, (1-)2.5-4 mm thick, terete to adaxially 

flattened, glabrous; domatia present or absent, usually 

a tuft of sparse hairs (white or yellow), 0.1-0.3 mm 

long, or barbellate at axils, or a few hairs surrounding 

a round slightly depressed glabrous area and converging 

toward its center (Fig. 57F). Inflorescences 

2 per node; corymbose, laxly branched, with opposite 

to subopposite decussate branches, cymules fairly 

compacted at distal end of each branchlet, forming a 

3-lobed flowering plane, (10-)13.5-20 

(7.5-)8-12.5 cm, L/W = ca. 2:1, 

obovate (to elliptic-obovate), acute-decurrent at base, 

obtuse at apex, sometimes with a deltoid acumen 0.5 cm 

x (8-)9.5-13 cm; 

lateral branches 3-4 pairs, basal portion of axis not 

branched (5.5-)8.5-10 cm long; L/A = 2:1 to 2.5:1; 

rachis decussately compressed, rachis and branches 

glabrous to puberulent sparsely short-pubescent; distal 

bracts absent or reduced to deltoid scales up to 2 x 

0.7 mm; bracteoles subtending flowers absent. Flowers 

sessile to short-pedicellate; pedicels to 7 mm long, 

glabrous to sparsely minute-puberulent; hypanthium 

obconical, 0.4-0.8(-1) x 0.4-0.8 mm, glabrous (to microscopically 

puberulent). Calyx extremely reduced 

and short-lobed, 0.4-0.5 x ca. 1 mm, glabrous; lobes 

deltoid to shallowly triangular, 0.1-0.5 mm long, often 

ciliate. Corolla funnelform, deeply lobed, 2.5-3.5 

mm long, white to cream-white; tube short cylindrical, 

1.5-1.8 x ca. 1 mm, glabrous outside and inside; 

lobes 5, 1.5-2.6 x 0.7-1.2 mm, oblong, rounded at 

apex, glabrous outside, with a tuft of white shaggy appressed-pilose 

hairs at medio-basal zone, with distal zone 

glabrous inside. Stamens 5, exserted, equal, attached 

1.5-1.8 mm from the base of the tube; filaments 2.6- 

3.1 mm long, terete, basally flattened, connate to 

throat, with a basal tuft of white-pilose hairs for half 

the filament length, the hairs decreasing in length toward 

medial zone; anthers yellow, elliptic, 0.5-0.6 x 

ca. 0.4 mm, dorsifixed at medial zone, base rounded, 

apex shortly apiculate (rounded to truncate). Pollen 

exine foveolate-reticulate. Style exserted, 2.6-3.5 mm 

long, as long as the corolla, terete to square, not costate; 

style branches extruding as two lips above corolla 

before anthesis, rounded, 0.3-0.4 mm wide, stigmatic 

surface microscopically (40x) papillose; ovary-galls 

not found. Capsules obovoid (to subglobose), with 

very thick pedicels, acute at base, shallowly hemispherical 

at apex, 1.5-2.5 x ca. 1.5 mm, rust-brown, 

without lenticels; disk black, glabrous, not exceeding 

the calyx; disk loculicidal dehiscence present in old 

capsules. Seeds 0.48-0.68 x 0.33-0.45 mm.


-[--------------------I---------------------------------------..... 

+rrl _rD 

~II ~C? - -c~B~q 

'': ' "'! 

tJ: ii ..~~~~~~~~~~~~~~~~ 

FIG. 66. Chimarrhis glabriflora (A-F from Ducke 1618, GH, isotype; G from Delprete & Verduga 6424, TEX). A. 

Habit of inflorescences. B. Flower with exserted anthers. C. Close up of glabrous disk. D. Detail of corolla, inside view. E. 

Stamen. F. Apical portion of style. G. Mature capsule. 

i 

~~ ntr~~~~'T 

II~~~ ~~~ dame~.


,'1 

4 :..- ^ ,,tli? ;i.... \? 

" ~ 

:'::~? ~ e 

' .~ 

~ 

i~~~~~~ic? 

'' ~ 

:'I''t"' ~' 

ifi'!i,i' i::," 

'~]gii!11.~1"i 

: 

I 

FIG.~~~~~~~~~* 

7. Ciari lbiir.A Toyugidvda c.30c b) .Flaeadinlrsecs .Tuko 

oldr ndviua (c. 0 m bh. . Dtal f hetrnk.(Pots akn a Jtfn aca iolgialSttin. 

FIG. 67. Chimarrhis glabrflora. A. Two young individual (ca. 30cm dbh). B. Foliage and inflorescences C Trunk of 

older individual (ca. 50 cm dbh). D. Detail of the trunk. (Photos taken at Jatun Sacha Biological Station.) 

Distribution (Fig. 77A) and ecology. Primary and 

secondary lowland (non-inundated) to upland rain 

forests or respective remnants, usually growing on 

well-drained lateritic soils (rarely in clay soil), in proximity 

of streams, of Amazonian Colombia, Ecuador, 

Peru, and Brazilian v&rzea forests. Flowering specimens 

were collected in January, February, March, 

April, May, October, and December. Specimens at 

intermediate stage between flowering and early fruiting 

have been collected in January, February, March, 

April, May, and August. Fruiting specimens were 

collected in all months of the year except March, 

April, and December. The cream-white, fragrant flowers 

have been reported to be visited by bees (Ecuador, 

Neill et al. 6220). 

Reproductive biology. In my several visits to 

Jatun Sacha Biological Reserve (Amazonian Ecua- 

dor), I observed a very interesting seed dispersal strat- 

egy of both C. glabriflora and C. hookeri, which co-


exist in this lowland forest. A mature individual of C. 

glabrifora (as most other species) during the flowering 

period(2-3 weeks) produces thousands of inflorescences, 

and their minute fragrant flowers are visited and 

most probably pollinated by bees. Approximately one 

month after fertilization the infructescences reach 

maturity and some of the capsules open, releasing their 

minute seeds. Most infructescences of the individuals 

observed were instead dropped (almost simultaneously) 

on the forest floor with many capsules still 

closed, forming a dense circle of the same diameter 

of the individual's crown. Most of these capsules open 

shortly after, releasing their seeds, and in about two 

weeks the young seedlings start sprouting in large 

circles just below the crown of the individual (sometimes 

directly from their capsules). Since the inflorescences 

are axillary on the terminal nodes, the 

branchlets resume their vegetative growth shortly after 

the infructescences were abscissed. 

The minute seeds of Chimarrhis have been thought 

to be wind-dispersed, which is probably the main 

means of dispersal in most species, but at least in this 

species (and C. hookeri) the mature infructescences 

seem to be a primary unit of dispersal. This dispersal 

behavior could also explain why Chimarrhis in the 

Amazon basin has been reported to have a tendency 

of clumping (see ecology discussion of C. turbinata, 

and Boom & Campos, 1991). 

N bank of Rio Napo, 200 m, 75?52'W, 00?36'S, 7-20 Jan 

1991 (fl), Bensman 331 (MO); Taisha, 500 m, 15 Feb 1962 

(fl), Cazalet & Pennington 7789 (K, NY, US); Coca, Palm 

Oriente Concession, 9 Sep 1983 (fl), Lescure 2060 (CAY, 

QCA); Puerto Misahualli, Reserva Biologica Jatun Sacha, 

01?04'S, 77?37'W, 450 m, 20 Aug 1994 (fl-fr), Delprete & 

Verduga 6423 (AAU, COL, G, MO, NY[2], QCA, QCNE, 

TEX), 6424 (AAU, CAS, COL, DAV, F, G, K, MO, NY[2], 

P, QCA, QCNE, TEX[3], UPS, US); Puerto Misahualli, 

Reserva Biologica Jatin Sacha, 9 Sep 1996 (fr), Delprete 

6526 (NY, QCNE). PASTAZA: Via Auca, 110 km S of Coca, 

10 km from Rio Tigiiifio, Sector Cristal, 01 15'S, 76?55'W, 

320 m, 7 Jan 1989 (fl), Palacios et al. 3380 (AAU, MO, NY, 

QCNE, TEX); Cton. Pastaza, "Ramirez" oil well, 20 km S 

of Curaray, 76051'W, 01?32'S, 300 m, 21-28 Feb 1990 (flfr), 

Zak & Espinoza 5130 (MO, TEX); Lorocachi, Rio 

Curaray, 75?58', 01 38'S, 200 m, 29 May 1980 (fl), Jaramillo 

et al. 31361 (AAU, MO). 

PERU. AMAZONAS: Rio Cenepa, vic. of Huampami, ca. 

5 km E of Chavez Valdivia, 78030'W, 04?30'S, 200-250 m, 

10 Aug 1978 (fr), Ancuash 1372 (F, MO, NY). HuANuco: 

Prov. Pachitea, Dtto. Honoria, Bosque Nacional Iparia, Rio 

Pachitea, 300-400 m, 18 Jan 1967 (fl), Schunke 1527 (F, G, 

GH, K, MO, NY, US); Prov. Pachitea, Dtto. Puerto Inca, 

Bosque Nacional de Iparia, 400-500 m, 19 Dec 1968 (fl), 

Schunke 2901 (F, NY, VEN); Prov. Pachitea, Dtto. Puerto 

Inca, 14 km from Rio Pachitea, 74?58'W, 093 I'S, 350 m, 

14 Apr 1982 (fl), Smith 1311 (MO[2]); Prov. Pachitea, Dtto. 

Puerto Inca, Yuyapichis, Unidad de Modelo y Producci6n 

Forestal DANTAS, 09?40'S, 75?02'W, 270 m, 1-15 Oct 1990 

(fr) Tello 313 (G), 16-31 May 1991 (fr) Tello 1995 (G). 

Representative specimens examined. COLOMBIA. JUNIN: Prov. Jauja, Reserva Forestal Granja, Satipo, 750 m, 

AMAZONAS: Mun. Leticia, Parque Nacional Amacayacu, trail 7 May 1963 (fr), Vdsquez 14-CBV(MO); Hda. Genova, 1600 

to Mata-Mata, 03?47'S, 70?15'W, 120 m, 21 Aug 1991 (fr), m, 8 Jul 1962 (fr), Woytkowsky 7380 (MO). LORETO: Dtto. 

Rudas et al. 3016 (MO). CAQUETA: Venecia, margins of Rio Mazan, Gamitanacocha, Rio Mazan, 100-125 m, 1 Mar 1935 

Ortegueza, 400 m, 31 Mar 1940 (fl-fr), Cuatrecasas 8947 (fl), Schunke 331 (A, F, NY, US); Dtto. Mazan, Rio Mazan, 

(F, NY, US). PUTUMAYO: Rio Putumayo, Puerto 28 

Espina, Apr 1977 (fr), Rimachi 2973 (F, NY); Prov. Alto Ama- 

00?10'N, 75?50'W, 23-26 Mar 1953 (fl), Schultes & Cabrera zonas, Shucushuyacu (Rio Huallanga), 75?50'W, 06?02'S, 

18929 (U, US). 

250 m, 12 Sep 1981 (fr), Vdsquez & Jaramillo 2431 (MO, 

ECUADOR. EL ORO: Lim6n-Playa, near Rio Dumari, NY, TEX); Prov. Maynas, Maniti, Recreo, 115 m, 72?50'W, 

03?29'S, 79?45'W, 500 m, 14 Oct 1993 (fl), Cornejo 582 03?42'S, 14 May 1988 (fl), Vdsquez 10631 (NY); Prov. 

(GUAY, QCA). MORONA-SANTIAGO: Centro Shuar Yukutais, Maynas, Quistococha, 9 May 1977 (fr), Revilla & Froehner 

W of Pedro Kunkumas house, 02?30'S, 78?08'W, 900 m, 31 2449 (F); Prov. Maynas, Quebrada Yanomomo, Rio 

Mar 1989 (fl), Bennett & Gomez-Andrade 3638, (QCNE); Amazonas above mouth or Rio Napo, 130 m, 15 Nov 1979 

Centro Shuar Yukutais, 11 Mar 1990 (fl), Bennett et al. 4028 (fr), Gentry & Jaramillo 28093 (MO). PASCO: Rio Palcazu 

(QCNE); Bomboiza, vic. of Salesian Mission in Shuar camp, Valley, second demonstration strip, 09?50'-10?45'S, 68?00'- 

03?25'S, 78?35'W, 800 m, 8-10 Jan 1986 (fl), Zaruma & 30'W, 300-600 m, 10 Mar 1986 (fl), Hartshorn et al. 2906 

Arguello 451 (K, NY, QCA, QCNE, TEX); Santiago, Rio (MO, NY, TEX); Rio Palcazu Valley, Samuel Ponce's pas- 

Santiago, 03?02'S, 77?58'W, 300 m, 16-17 Oct 1975 (fr), ture, 300-600 m, 15 May 1986 (fr), Hartshorn et al. 2925 

Little et al. 763 (MO). NAPO: Estaci6n Esperimental de (NY, TEX), and 20 Jun 1986 (fr), 2632 (MO); Prov. 

INIAP, San Carlos, 6 km E of Los Sachas, 250 m, 4 Apr 1985 Oxapampa, Rio Palcazu Valley, Rio San Jose, 10?09'S, 

(fl), Neill et al. 6220 (AAU, MO, NY, QCA, QCNE); San 75?20'W, 400 m, 13 May 1983 (fl-fr), Smith 4019 (MO[2], 

Jose Payamino, 40 km W of Coca, 00?30'S, 77?20'W, 300- NY). SAN MARTIN: Prov. Mariscal Caceres, Dtto. Tocache 

600 m, 19 Jan 1984 (fr), Irvine 634 (F, NY[2], QCA); 3.5- Nuevo, Quebrada Cafiuto, 520 m, 2 Jan 1979 (fl-fr), Schunke 

4.8 km E of Rio Conejo on rd. to Lago Agrio, 340 m, 1 Apr 10653 (F, K, MO, NY). 

1972 (fl), MacBride & Dwyer 1408 (MO, QCA); La Joya BRAZIL. ACRE: Cruzeiro do Sul, Rio Jurua & Rio Moa, 

de los Sachas, 76?37'W, 00?25'S, 250 m, 22 Aug 1992 (fl), Serra da Moa Village, 26 Apr 1971 (fl-fr), Prance et al. 

Gudino 1707 (NY); Cton. Aguarico, Chiro Isla community, 12462 (GH, MO, NY[2], U[2], US).


Local names and uses. Ecuador: mincha caspi 

(Quichua, = wick tree) (Irvine 634, Napo, Irvine 585), 

mecha caspi (Quichua, Bensman 331), jatun mincha 

caspi (= large wick tree) (Napo, Irvine 204). Peru: 

itauba (Tello 1995), itauba amarilla (Tello 313), cascarilla 

masha (Quichua, Schunke 1527, 2901; 10653), 

palo palillo (Junin, C. B. Vasquez 14-CBV), pablo 

manchana, pampa remocaspi (Quichua, = 

that detached easily in irregular pieces. The tree architecture 

of this species follows the Aubreville Model 

(cf. Halle et al., 1978). 

7. Chimarrhis hookeri K. Schumann in Martius, Fl. 

Bras. 6(6): 259. 1889. Type. Peru. San Martin: 

paddlewood), and yero prueba (the last three names Tarapoto, in tall forest, tree ca. 25 m tall, Jan 1857 

reported by Duke & Vasquez, 1994). 

(fl), Spruce 4930 (B*; lectotype, K, selected by 

This species is reported by many foresters and in- Delprete, 1999b; isolectotypes, BR, C-n.v., F[2], 

digenous tribes to have very hard wood G, GH, K, NY, P; 

(with copiphoto-C 

at F, GH, MO, NY, 

ous watery orange juice when cut), providing excel- VEN). Spruce annotated these specimens as "Conlent 

timber for construction of houses (Peru, Schunke 

daminea glabrata DC. [unpubl.] aff., a curious 

331). Duke and Vasquez (1994) reported that the wood thing!" Figs. 9G, 18C, 58G, 68A-H, 69A-D, 70, 77B 

of this species is used for rural construction, firewood, Chimarrhis williamsii Standley, Publ. Field Columbian 

and living fences. Bensman 331 bears the following Mus., Bot. Ser. 8: 162. 1930. Type. Peru. Loreto: 

notes: "tree 55 mm high and over 50 cm at DBH, with Rio Nanay, 23 May 1929 (fl), LI. Williams 409 (holarge 

buttresses, yellowish wood with thin bark, used lotype, F, N. 601986; frag-F at G). 

as construction wood." 

Trees to 35 m tall, 50 cm or more dbh (exception- 

Chimarrhis glabriflora is sympatric with C. ally 50 m tall, 1 m dbh, Baker et al. 6000), much 

hookeri (Fig. 77A-B), with which it is commonly branched at crown, with tall buttresses; bark fissured, 

confused, but from which it differs in having larger white-grayish, peeling off in long fibrous strips exand 

rather fleshy leaves, which are deep green and posing the reddish bark underneath; wood yellow, 

shiny above, semi-succulent leafy branchlets (Figs. very hard. Leafy branchlets 3-4(-5) mm thick, 

66A, 68B, 70), a larger inflorescence, glabrous cap- woody (not succulent), subterete to decussately comsules, 

and glabrous disks. The leaf veins in C. pressed, youngest 2-3 nodes densely appressed-canesglabriflora 

are always glabrous and the domatia are cent; older branches glabrate, terete, grayish to pale 

rather variable in either being represented by a small brown; lenticels not found. Stipules free at base, contuft 

of hairs, or a peculiar area of hairs attached on torted in bud, narrowly triangular, acuminate, densely 

the veins' axil and perpendicular to them (resembling canescent (rarely puberulent or rarely glabrous) outa 

small brush), or completely absent. 

side, glabrous with sparse small colleters at base in- 

When Ducke (1945) described this species, he side, 12-18(-30) 

added the following notes: "This well defined species 

is easily distinguished from the two remaining species 

of Brazilian Amazonia (Ch. turbinata and Ch. 

barbata), by various botanical characters, chiefly by 

the very small calyx and by the entirely glabrous corolla. 

It is a rather frequent tree of the upland forest 

of the western limit of Brazil, and will certainly be 

observed in the neighboring parts of Peru and Colombia. 

The fragrant flowers appear at the climax of the 

rainy season. The tree yield good fire wood." 

Strangely Ducke (1945) did not compare this species 

to its close relative C. hookeri, from western Amazon 

(and with which it is sympatric), which is well 

described in Flora brasiliensis (Schumann, 1889). 

The largest individuals of Chimarrhis that I have 

personally seen belong to this species (at Jatun Sacha 

Biological Station, Ecuador). One individual had large 

buttresses (4-5 m long), the trunk about 1.5 m in diameter 

at about 4 m above ground (above the buttresses) 

and ca. 45 m tall, and rust-brown fibrous bark 

x 3-5 mm, gray-canescent, with 

many parallel veins departing from base, readily caducous, 

leaving a white scar encircling the stem, 0.5- 

1 mm wide. Leaves 6-16(-20) x 3-6.5(-9) cm, L/W = 

2:1, elliptic to elliptic-obovate, acute at base, obtuse 

at apex, sometimes with a deltoid acumen 0.5 cm long; 

dark green above, pale green below, semi-coriaceous; 

drying dark brown above, olive-green below, stiff-chartaceous; 

juvenile leaves with blade, primary and secondary 

veins sparsely appressed pubescent above, 

mature leaves glabrous above, appressed pubescent 

below; primary and secondary veins ascending-pubescent 

to canescent (rarely glabrous), secondary veins 

8-9 each side; tertiary veins starting subparallel and 

reticulate in the middle; petioles 8-20 mm long, 1- 

1.5 mm thick, terete to adaxially flattened, glabrous 

to puberulent to canescent; domatia represented by a 

few hairs surrounding a round slightly depressed glabrous 

area and converging toward its center (Fig. 

57G), hairs 0.4-0.7 mm long. Inflorescences 2 (rarely 

4) per node; corymbose with opposite to subopposite


_ .. 

DI II - 

FIG. 68. Chimarrhis hookeri (B-G from Spruce 4930, K, lectotype; H from Delprete & Verduga 6421, TEX). A. Archi- 

tecture and dimensions of mature individual (from pers. obs.). B. Habit of inflorescence. C. Flower with exserted anthers. 

D. Close up of pubescent disk. E. Detail of corolla, inside view. F. Stamen. G. Apical portion of style. H. Mature capsule.


I 

? :B' . 

g 

N,~~~~. 

r~~~~~~ 

~~~~~~~~~~~~~~~~~~~~~~~~?u 

I ? -.:'. ' :' 

" 

~~~~~~rC? . ....~ ~ Q~?X ~ ~ ~ *F ~ ~ ~ ~ ''~.:~i B 

... 

P l~~~~~~ip~~~~~" 

, -, 

t,: 

. ! ' * , 

,' 

.. 

I, ? ' 

,. " 

'' i.@. 

', pv":o ',:....,,,i{!'.. 

..: , . ,,.',.:~,: 

?~~~~~~~~~~, 

.:. i,?" 

t~~~~~~~~~~~~~~~~~~~~~~~~~~.., 

! 

Stat , E: 'd 

... 

: . 

~~~~~~~~~~~~~~.., 

}~,, ..~:.,:.1:, :. 

~.'/~i,~;. . 

g tt l i / yF~~~~~~~~~~~~~~~~~~~~';,. 

. 

r. tr~tz i 

i ?~~~~~~~~~~~ ~ ~ ~ ~ ~ I~~~~~~~~~. 

tM, 

, ~.,_.;. 

i F, ,?i ??; ; 

| ~ I i 

_l \ i;'9 S2 

_ r Y J i t s i t 't | ?' ,1\1 i.- ." ;' 

';' 

, 

i 

:.. ??;::'":4i.: 

FIG. 69. Chimarrhis hookeri. A. Tree climber reaching the c:rown of..-- a:youn. indiidual. Inf'l-~e.c:en.es.w,'h," 

;., a B. 

- t.re 

[i. I! 

-, 

,~!~ 

fruits~~~' (se figrfrsae.C. Trun of anodenivda (ca~~..",:', bh.: D.Dt.ail of.the'trunk.-? (Pooae tJti 

c4 

: 

40 4 

i ~ .'?, ~~~~~ ~ "!~'. ~ ~~~~~~~~~~~~~; 

: ,~~ '; :i 

1 "?'? _ ~ ~ S ~ i ~ ~ l v.,,.' ~ : i, :r ': 

; \91. , ; 

r'~' '' 

? 

| [f; \ l 

F'~~~ ~~~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~~". 

FIG. 69. Chimarrhis hooleri. A. Tree climber reaching the crown of a young individual. Infiorescences B. 

with mature 

Sacha Biological Station, Ecuador).


:.... .,. e . .. .... 

H? ? 

_s __? 

cal Station, Ecuador). 

FIG. 70. Terminal leafy branchlets of Chimarrhis glabriflora (left) and C hookeri (right). Note larger leaves and semi- 

cal Station, Ecuador). 

decussate branches, cymules fairly compacted at dis- 

tal end of each branchlet, forming a continuous flow- 

ering plane, 8-10(-12) x (4-)5.5-7.5 cm, lateral 

branches 2(-3) pairs, basal portion of axis not 

branched 4-5.5(-8) cm long; L/A = 2.5:1 to 2.7:1 

(rarely 3:1); rachis decussately compressed, rachis and 

branches sparsely yellow-puberulent; distal bracts 

reduced to scales or linear, 1-3 x 0.3-0.5 mm; 

bracteoles subtending flowers absent. Flowers sessile 

to short-pedicellate; pedicels to 1 mm long, glabrous 

to sparsely pubescent (to short-villous); hypanthium 

obovate, 0.9-1.2 x 0.8-1.1 mm, glabrous, reddish 

brown. Calyx extremely reduced, truncate to short- 

lobed, 0.3-0.5 x 1-1.5 mm, glabrous; lobes shallowly 

triangular, connected by a membrane, 0.1-0.6 mm 

long, often ciliate. Corolla funnelform, deeply lobed, 

2.5-3.5 mm long, cream-white to yellowish white 

(turning pale yellow at maturity); tube short cylindri- 

cal, 1-1.5 x 0.7-1 mm, glabrous outside and inside; 

lobes 5, 1.5-2 x 0.5-0.7 mm, oblong, rounded at apex, 

glabrous outside, with a tuft of white thin-sericeous 

hairs at basal zone with medial and distal zones gla- 

brous inside. Stamens 5, exserted well above the co- 

rolla, equal, attached 1-1.5 mm from the base of the 

tube; filaments 2-2.5 mm long, terete, basally flat- 

tened, connate to throat, with a dense tuft of white- 

sericeous hairs for half the length at base; anthers el- 

liptic, 0.5-0.7 x 0.4-0.5 mm, dorsifixed at medial 

zone, base rounded, apex rounded, not mucronate. 

Pollen exine foveolate-reticulate. Style exserted, 2.5- 

3.5 mm long, as long as the corolla, terete to square 

(not costate), often with few sparse hairs on distal 

portion; style branches extruding as two lips above 

corolla before anthesis, rounded, ca. 0.5 mm wide, 

stigmatic surface microscopically (40x) papillose; 

ovary-galls not found. Capsules subglobose to obo- 

void, obtuse at base, shallowly hemispherical at apex, 

2-2.5 x 1.5-2 mm, capsule dark brown, without len- 

ticels; disk black, sparsely to densely pubescent, not 

exceeding the calyx; disk loculicidal dehiscence 

present in old capsules. Seeds 0.5-0.63 x 0.35-0.5 mm. 

Distribution (Fig. 77B) and ecology. Primary and 

secondary rain forests, and respective remnants, in 

riverside forest, in alluvial, lateritic, sandy to clay 

soils, from low to high fertility, 150-800 m, of Ecua-


dorian and Peruvian Amazon. Flowering specimens (fl), Gutierrez 32 (MO); Prov. Pachitea, Dtto. Honoria, rd. 

were collected in January, February, May, June, Septo 

Turavista, 24 Jan 1963 (fl), Lao Magin 44 (F, K, NY), 

tember, and December. Specimens at intermediate 31 May 1963 (fr), 51 (F, NY, US). LORETO: Upper Rio 

stage between flowering and early fruiting have been Marafion, mouth of Rio Santiago, 160 m, 3 Dec 1924 (fl), 

Tessman 4668 

collected in April and 

(B, F, NY); Prov. Maynas, Pto. Allianza, 

August. Fruiting specimens 

72?55'W, 04?08'S, 160 m, 29 May 1981 (fr), Vdsquez & 

were collected in February, April, May (many), June, Criollo 1855 (F, MO, NY, TEX); Prov. Maynas, Yanomono, 

September, October, and December. 

Explorama Tourist Camp, between Indiana and mouth of Rio 

Napo, 130 m, 18 Feb 1981 (fr), Gentry et al. 31384 (F, G, 

Representative specimens examined. ECUADOR. MO, NY); Rio Nanay, 23 May 1929 (fl), Williams 369 (F, 

ESMERALDA: Canton Esmeralda, San Lorenzo-Lita, trail to K, US); Bosque Nacional Alexander Von Humboldt, km 91 

Punta, 00?49'N, 78?26'W, 200 m, Macias & Burbano 7788 on rd. Pucallpa-Lima, 300-500 m, 12 May 1975 (fr), Hart- 

(QCNE). NAPO: Estaci6n Experimental INIAP, San Carlos, shorn 1687 (F, MO[3]). MADRE DE Dios: Tambopata, 

6 km SE of Los Sachas, 250 m, 4 Apr 1985 (fr), Palacios et 12?49'S, 69?18'W, 280 m, 26 Feb 1984 (fr), Gentry et al. 

al. 240 (AAU, F, MO, NY, QCNE, TEX, US); Reserva 45661 (MO, NY), 46255 (MO). UCAYALI: Prov. Coronel 

Biol6gica Jatun Sacha, 8 km downstream from Puerto Misa- Portillo, Pucallpa-Tingo Maria Hwy., 75?00'W, 08?41'S, 

hualli on Rio Napo, 01?04'S, 77?36'W, 450 m, Palacios et 250m, 11 Feb 1981 (fr), Gentryetal. 31219(F, MO); Prov. 

al. 365 (AAU, F, MO, NY, QCNE, TEX); Rio Aguarico, near Coronel Portello, Campo Verde, km 78 C.F. Basadre, 250the 

confluence of Rio Pavayacu, Sep 1981 (fr), Bravo & 270 m, 13 Mar 1989 (st), Chavez 265 (MO). 

G6mez 306 (QCA); Amazon primary forest, 77?28'W, 

01?20'S, 520 m, 19 May 1976 (fr), Oldeman & Arevalo 16 Local names and uses. Ecuador: intacchi 

(QCA); Parque Nacional Yasuni, road-oleoduct Maxus in (Quichua, Napo, Baker et al. 6000). Peru: itauba (Tello 

construction, km 46 to Capiron Well, primary forest, 862), itauba amarilla (Tello s.n.), itahuba (Pachitea, 

00?41'S, 76?29'W, 244 m, 6-12 Sep 1993 (fl), Dick 290 M.A. V 42), tuwara (Ll. Williams 369,409), yacu-caspi 

(QCNE); 5 km SE of Las Sachas, 300 m, 13 Apr 1985 (fr), (Tessman 4668), huacapu masha (Hartshorn 1687), 

Baker et al. 6000 (MO, NY); Parque Nacional Yasuni, La 

quillo-bordon, purma-caspi (Quichua, = yellow- 

Joya de Los Sachas, rd. Maxus oleoduct, km 15, 00?25'S, 

wood), 

76?37'W, 250 m, 27 Feb 1993 (fl), Gudino & 

chollachaqui 

Grefa 2336 

caspi blanco (Ucayali, Chacvez 

(TEX); Canton Archidona, SE of Volcan Sumaco, rd. 265), sacha jagua (Leoncio Prado, Gutierrez 32), 

Hollin-Loreto, km 65, 00?44'S, 77028'W, 600 m, 26 Jun papelillo caspi (Huanuco, Lao Magin 44 and 51), 

1989 (fr), Alvarado 292 (AAU), Hurtado 2365; Cton. Tena, 

aromuhe (Maynas, Paitan 3b), pampa remo caspi 

8 km E of Puerto Misahualli, Reserva Biologica Jatun Sacha, (Maynas, Flores 61 [Quichua 

01?04'S, 77?36'W, 400 m, 20 Aug 1994 (fl-fr), Delprete & 

Verduga 6421 (AAU, F, G, K, MO, P, NY[2], QCA, QCNE, 

TEX, UPS[2], US), 6422 (COL, NY, QCA, TEX); Puerto 

Misahualli, Reserva Biologica Jatun Sacha, 9 Sep 1996 (fr), 

Delprete 6527 (NY, QCNE), 6528 (NY, QCNE). PASTAZA: 

Rio Curaray, 75040'W, 01?36'S, 200 m, 5 Jun 1980 (fl), 

Brandbyge & Asanza 31615 (AAU, MO, QCA, US); 

Lorocachi, 2 km S of village, left margin of Rio Curaray, 

75?58'W, 01?38'S, 200 m, 29 May 1980 (fl), Jaramillo et 

al. 31631 (AAU, QCA, US); rd. PETRO-CANADA, Via 

Auca, 115 km S of Coca, 4 km S of Rio Tigiifio, 01?15'S, 

76?55'W, 320 m, 22-28 Feb 1989 (fl), Zak 3979 (G, GB, 

MO, QCNE, TEX); Montalvo, 1 km NE of military camp, 

76058'W, 02005'S, 250 m, 17-19 May 1979 (fl), Lojtnant & 

Molau 13513 (AAU). SUCUMBIOS: Cant6n Lago Agrio, Parr. 

Tarapoa, 1 km from Mariann 3 oil well toward Reserva 

Faunistica Cuyabeno, 76022'W, 00?08'S, 240 m, 27 Apr 1990 

(fl-fr), Cer6n & Ayala 9656 (MO, TEX, QCNE). 

PERU. AMAZONAS: Rio Santiago Valley, 77?40'W, 

03?50'S, Quebrada Caterpiza, 3 Dec 1979 (fl), Tunqui 205 

(MO); Rio Cenepa, Quebrada Sasa, 30 May 1973 (fr), 

Ancuash 504 (F, MO, NY). HUANUco: Prov. Puerto Inca, 

Dtto. Yuyapichis, Unidad de Modelo y Producci6n Forestal 

DANTAS, 09?40'S, 75002'W, 270 m, 1-15 Dec 1990 (fr) 

Tello 862 (G); Prov. Huamalies, Cordillera Azul, Fundo 

Sinchono, km 209, 1600 m, 12 Oct 1944 (fr), Hodge 9 (F); 

Prov. Leoncio Prado, Hda. Laconyope, 870 m, 17 Jan 1962 

= paddlewood]), 

mukugd (Huambisa, Amazonas, Tunqui 945), pablo 

manchana, yemo prueba, and palo perro (the last three 

nemes reported by Duke & Vasquez, 1994). 

The label of Baker et al. 6000 reports: "Tree 50 m 

tall, I m dbh, with a crown roundish and flattened on 

top; its wood is very hard and the indigenous people 

use it to build their houses." Additional uses are reported 

as: "Its bark is used to make ropes, and its wood 

is of potential value for production of particle boards" 

(Lao Magin 44 and 51), and "used for construction of 

boats" (M.A. V. 42). 

This Amazonian species is recognizable by its 

small leaves, thin and readily woody leafy branchlets 

(Fig. 70), small capsules, and pubescent disks; leaf 

blades sparsely puberulent to glabrous below; leaf 

veins ascending-pubescent to canescent (rarely glabrous); 

and domatia often present as a tuft of pilose 

hairs converging toward the center of the axils. When 

the blades and the veins are glabrous the domatia are 

absent (Fig. 58G). Mature individuals of C. hookeri 

have a tree architecture that follows the Aubreville 

Model (cf. Halle et al., 1978). 

Chimarrhis hookeri is often confused with C. 

glabriflora (with which it is sympatric), the former


differing from the latter in having smaller, thinner 

leaves (Figs. 66A, 67B, 70), with 8-9 pairs of secondary 

veins (vs. 12-16 pairs of veins), slender woody 

(vs. semi-succulent and thick; see Fig. 70) branchlets, 

smaller, densely appressed-pubescent to canescent 

(vs. sparsely puberulent) stipules, and a pubescent (vs. 

glabrous) capsule disk. 

Chimarrhis williamsii was established by Standley 

(1930b) from material collected in the proximity of 

Rio Nanay (Prov. Loreto, Peru). He distinguished it 

from C. hookeri by erroneously stating that the latter 

has glabrous stipules. He also admitted that he did not 

see any material of the latter species. The disk and the 

stipules of C. williamsii (Ll. Williams 396 and 409) 

are sparsely pubescent, although the rest of the vegetative 

and reproductive characters closely match 

those of C. hookeri. The two taxa are here treated as 

synonymous. 

8. Chimarrhis microcarpa Standley, Bull. Torrey 

Bot. Club 53: 471. 1926, non Chimarrhis cymosa 

Jacquin subsp. microcarpa Urban, Symb. Ant. 1: 

411. 1899 (= C. cubensis Steyermark), non 

Chimarrhis cymosa Jacquin var. microcarpa (Urban) 

Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 

192. 1936 (= C. cubensis Steyermark), Chimarrhis 

microcarpa var. microcarpa, Mem. New York 

Bot. Gard. 12(3): 183. 1965. Type. Trinidad. 

Maraval, 1904 (fr), Dannouse 6946 (holotype, 

NY). Figs. 71A-G, 73 

Chimarrhis longistipulata Bremekamp, Recueil Trav. 

Bot. Neerl. 31: 260. 1934. Type. Surinam. Kabalebo 

River, Avanero Falls, 3 Sep 1920 (fr), Pulle 379 (holotype, 

U, photo-U at NY, VEN; isotypes, K, US). 

Trees to 40 m tall, 100 cm or more dbh, canopy 

trees with lobed bole, small spreading crown, with 

buttresses to 3.35 m high; bark fissured, grayish 

brown; wood yellow, very hard. Leafy branchlets 

2.5-5(-6) mm thick, subterete to decussately compressed, 

green to brown, glabrous; older branches terete, 

grayish to pale brown; lenticels not found. 


acuminate, sparsely appressed-puberulent 

(rarely glabrous) outside, glabrous with basal colleters 

inside, (20-)25-35 x (4-)5-7 mm, reddish brown, 

membranous, with many parallel veins departing from 

base, readily caducous, leaving a scar encircling the 

stem ca. 1 mm wide. Leaves (10-)16-22 x the deltoid acumen 0.5-1.8 cm long; dark green above, 

pale green below, thick-foliaceous; drying brown to 

olive-green, stiff-chartaceous; glabrous above, glabrous 

to sparsely appressed-puberulent below; primary 

and secondary veins glabrous to appressed-puberulent, 

secondary veins 8-11 each side; tertiary 

veins subparallel, laterally jointed in the center; petioles 

(13-)18-31 mm long, 1-1.5(-2) mm thick, adaxially 

flattened to concave, glabrous; tuft-domatia 

always present, tuft of hairs departing from the veins 

and converging toward the center of the axil (Fig. 

58E), only at the axils of primary with secondary 

veins, hairs 0.7-1 mm long, white to golden. Inflorescences 

2 per node, corymbose with opposite to 

subopposite decussate branches, cymose, very compacted 

at distal end of each branchlet, forming a continuous 

flowering plane, (8-)10-12(-15) 

(5-)6.5- 

9.5 cm, L/W = ca. 2:1, elliptic to obovate, acute-decurrent 

at base, acute to obtuse and mucronate at apex, 

x (4-)5-7 

(-9) cm, lateral branches 2(-3) pairs, basal portion of 

axis not branched (5-)6.5-10.5 cm long; L/A = ca. 

3.1; rachis terete to decussately compressed, rachis 

and branches short puberulent; distal bracts usually 

absent or narrowly triangular, to 4 x 3 mm; bracteoles 

subtending flowers absent. Flowers sessile to shortpedicellate; 

pedicels to 1 mm long, glabrous to sparsely 

puberulent; hypanthium obconical, 0.8-1 x 0.8-1.2 

mm, glabrous, reddish brown. Calyx extremely reduced, 

truncate to faintly undulate margin, 0.3-0.6 x 

1-1.4 mm, glabrous. Corolla funnelform, deeply 

lobed, 4-5 mm long, cream-white to greenish white 

(turning pale yellow at maturity); tube short cylindrical, 

0.9-1.5 x 0.6-1.1 mm, glabrous outside and inside; 

lobes 5, 1.8-3.2 x 0.4-0.6 mm, narrowly oblong, 

acute (rarely rounded) at apex, glabrous outside, 

sparsely white-villous at basal zone, with medial and 

distal zones glabrous inside. Stamens 5, exserted well 

above the corolla, equal, attached 1-1.7 mm from the 

base of the tube; filaments 3.5-5.5 mm long, terete, 

basally flattened, connate to throat, with a tuft of 

densely white-pilose hairs at base; anthers elliptic, 

0.6-0.7 x 0.3-0.4 mm, dorsifixed at medial zone, reversed 

at maturity, base rounded, apex shortly apiculate, 

truncate to rounded, dehiscing by longitudinal 

slit. Pollen exine foveolate-reticulate. Style exserted, 

3-5 mm long, as long as the corolla, glabrous, minutely 

costate to minutely winged; style branches 

extruding as two lips above corolla before anthesis, 

rounded to oblate, ca. 0.3 mm long, stigmatic surface 

microscopically (40x) papillose; ovary-galls not 

found. Capsules subglobose, obtuse at base, rounded 

at apex, 1.9-2.5 x 1.3-1.7 mm, dark brown without 

lenticels, glabrous throughout; disk black, exceeding 

the calyx; disk loculicidal dehiscence present in old 

capsules. Seeds 0.68-73 x 0.53-0.7 mm.


0 

FIG. 71. Chimarrhis microcarpa (B and G from Dannouse 6946, NY, holotype; C-F from Smith 3433, F). A. Architec- 

ture and dimensions of mature individual (from pers. obs.). B. Habit of inflorescences. C. Flower with exserted anthers. D. 

Detail of corolla, inside view. E. Stamen. F. Apical portion of style. G. Mature capsule. 

ic



secondary rain forests, and respective remnants, mixed 

forests, and recent regrowth vegetation, on lateritic 

soils (often on red clay), usually in proximity of creeks 

and streams, on slopes at low elevation, 20-600 m, 

of eastern Venezuela, Trinidad, Guyana, French 

Guiana, Surinam, and northern Brazil. Flowering 

specimens were collected in January, February, 

March, April, and October. Fruiting specimens were 

collected in January, February, March, April, June, 

August, September, and December. 

Representative specimens examined. VENEZUELA. Local names and uses. Venezuela: carutillo (Delta 

ANZOATEGUI: Between Rio Le6n ofQuebrada Danta and Rio Amacuro, Marcano Berti 241), totumo amarillo (Mar- 

Zumbador, NE of Bergantin, 500-600 m, 24 Feb 1945 (fl), cano Berti 1426), amarilla (Anzoategui, Steyermark 

Steyermark 61136 (A, F[2], NY, US, VEN). BOLIVAR: Si- 

61136), agua fresca (Merida, Bernardi 5943). Guyana: 

erra de Lema, source of Rio Chicanan, 80 km SW of El white calabash (McDowell 4472). Steyermark (1974) 

Dorado, 60?05'N, 62?W, 500 m, 27 Aug 1961 (fr), 

Steyermark 89514 (NY, VEN); Reserva Forestal La 

reported that the hard wood of this species is used to 

Paragua, make 

margins of Rio Asa, Jun 1970 (fr), Blanco 822 (F, NY, 

wheel-spokes, beams, and posts. 

VEN). 

TERRITORIO FEDERAL DELTA AMACURO: Close to border with 

Estdo. Bolivar, E of Rio Grande, ENE of El Palmar, 21 Jun This species is unique in having narrow-funnel- 

1964 (fr), Marcano Berti 241 (F, K, MO[4], NY, VEN); form corollas 4-5 mm long, small, compactly 

Lower Orinoco, May 1896 (fl), Rusby & Squires 126 (A, F, branched inflorescences (L/A = 3:1), slender woody 

G, GH, K[2], MO, NY[2], US[2]); vic. of Rio Grande, E of leafy branchlets, and leaves with 8-1 1 secondary veins 

Upata, 61?44'W, 08?14'N, 7 Apr 1967 (fl), Brujin 1646 (F, (Fig. 71B). Chimarrhis microcarpa is a rather com- 

M, MO, P, NY, US, VEN), 1647 (F, M, MO, P, NY, US, mon species in the lowland rain forests of northeast- 

VEN), 1651 (F, M, MO, P, NY, US, VEN). LARA: Dto. ern South America, reaching dimensions of large 

Iribarren, vic. of Laguna Negra, Loma de los Naranjos, 

Montafia de Macanillal and Fila de San Esteban, 10-19 km 

canopy-trees (40-50 m tall and to 1.5 m diam.; see 

S of Rio Claro, 1300-1500 m, 24-25 Mar 1975 Fig. 71A), and often described as trees with lobed 

(fl), 

Steyermark et al. 111625 (MO, NY, VEN). MONAGAS: boles, large buttresses, and small flattened crowns. 

Montafia de Aguacate, along Quebrada de Pajarral, 

The tree 

tributary 

architecture of this species is the Aubreville 

of Rio Caripe, NE of Alto de Aguacate, 600-900 m, 19 Apr 

Model (Terminalia branding; cf. Halle et al., 1978). 

1945 (fr), Steyermark 62185 (A, F, NY, VEN). SUCRE: Its bark is shallowly fissured and grayish brown; when 

Manacal, NW of Irapa, 800 m, 10 Dec 1966 (fl), Marcano freshly cut its sapwood is creamy-white to yellowish 

Berti et al. 1426 (VEN). 

white, and its heartwood pale-brown to dark-brown. 

TRINIDAD AND TOBAGO. TRINIDAD: Heights of Its leaves are thick-papery, dark-green and shiny 

Aripo, 10-26 Jan 1922 (fl), Broadway 10011 (NY, US); above and pale-green below, and its flowers are sweet- 

Blanchisense rd., 11 m post, 31 Dec 1926 (fr), Broadway 

6464 

fragrant. 

(K). 

Chimarrhis 

GUYANA. Kanuku Mtns., NW slopes, drainage of 

microcarpa differs from C. speciosa 

Moku-moku Creek, 150-400 m, Mar-Apr 1938 (fl), Smith 

in the larger funnelform corolla, slender and woody 

3433 (A, B, F, G, IAN, MO, NY, P, U, US); Rupununi, near leafy branchlets, and the much smaller, more compact 

Moco-Moco, 03?20'N, 59?35'W, 100 m, 29 Oct 1979, Maas inflorescences. The capsules in C. microcarpa and C. 

& Westra 3912 (U, VEN); Barima-Waini Region, Barima speciosa are similar in size and shape, and have gla- 

Head, along Barima River, 07?35'N, 60035'W, 91-122 m, brous disks; but in C. microcarpa the pedicels are 

15 Apr 1991 (fl), McDowell et al. 4411 (G, U, US); Barima- longer and more slender, while in C. speciosa the 

Waini Region, Matthew Ridge to Blue Mtn., 07?26'N, capsules are subsessile (vs. pedunculate). 

6011' W, 100-400 m, 28 Apr 1991 (fl), McDowell 4472 

Many herbarium specimens of C. microcarpa have 

(MO); Lower Essequibo River, Groete Creek, 19 Apr 1943 

(fl), Fanshawe F1249 (3985) (F, K[2], NY[2], U); Potaro- 

Siparuni region, 4-5 km N of Surama village, confluence of 

Burro-Burro and Surama rivers, 04?10'N, 59003'W, 75 m, 1 

May 1992 (fl), Hoffmann 1532 (US). 

FRENCH GUIANA. SAUL: Grand Boeuf Mort, 

03?37'N, 53?12'W, 250-300 m, 15 Jun 1988 (fl), Mori & 

Gracie 18943 (CAY); Petit BoefMort circuit, 21 Sep 1974 

(fr), Granville B-5167 (CAY, U, US); Monts La Fum6e, 

03?37'N, 53?12'W, 200-400 m, 2 Oct 1982 (fl), Mori et al. 

15109 (CAY, VEN), 15016 (P, US); Monts La Fumee, 200- 

400 m, 13 Oct 1982 (st), Boom & Mori 2037 (CAY), 2041 

(CAY), 2043 (CAY); layon toward Limonade bend, ca. 2 

km from village (crique Douille), 17 Jan 1976 (fr), Granville 

B-2671 (CAY, U); trail of Batardeau, 4 Mar 1977 (fr), 

Granville B-2799 (CAY, U); Mont Galbao, E sect., 600 m, 

03?35'N, 53020'W, 30 Jan 1986 (fr), Granville et al. 9041 

(B, CAY, US). 

BRAZIL. RORAIMA: Serra Tepequem, 1200 m, 17 Feb 

1967 (fr), Prance etal. 4452 (F, IAN, M, MG, NY, R, U, VEN). 

been misidentified as C. cymosa, but the latter has 

thick costate succulent branchlets, a much larger in- 

florescence, larger leaf blades (Fig. 65A-F), typical 

tuft-domatia (Fig. 58D), and is endemic to the Lesser 

Antilles (Fig. 63). As already noted by Steyermark 

(1965), C. longistipulata (from Surinam) is synony- 

mous with C. microcarpa.


9. Chimarrhis speciosa (Steyermark) Delprete, sp. 

et stat. nov. Chimarrhis microcarpa Standley var. 

speciosa Steyermark, Mem. New York Bot. Gard. 

12(3): 184. 1965. Type. Venezuela. Aragua: 

Rancho Grande, 18 Sep 1951, Garcia 147 (holo- 

type, VEN). Figs. 9H, 18A,B, 72A-G, 73 

Trees commonly 15-20 m tall, 30-40 cm dbh [exceptionally 

30 m tall, 80 cm dbh, Pittier 15275], with 

lobed bole, sometimes with small buttresses; bark 

grayish brown; wood yellow, very hard. Leafy 

branchlets (4-)5-8 mm thick, subterete to decussately 

compressed, greenish brown, often herbaceous, 

glabrous; older branches terete, grayish to pale brown; 

lenticels not found. Stipules free at base, contorted 

in bud, narrowly triangular, acuminate, sparsely appressed-puberulent 

(rarely glabrous) toward base and 

at margins outside, glabrous with basal colleters inside, 

25-30 x 5-7 mm, reddish brown, membranous, 

with many parallel veins departing from base, readily 

caducous, leaving a scar encircling the stem ca. 1 mm 

wide. Leaves (12-)18-27 x (5-)9-12 cm, L/W= 1.5:1 

to 2:1, widely elliptic to obovate, acute-decurrent at 

base, obtuse and mucronate at apex, the deltoid acumen 

0.5-1.8 cm long; dark green above, pale green 

below, semi-coriaceous; drying olive-green, stiffchartaceous; 

glabrous above and below; primary and 

secondary veins glabrous to sparsely appressed-puberulent, 

secondary veins (9-)11-12 each side; tertiary 

veins subparallel and laterally jointed in the center; 

petioles (10-)18-36 mm long, (1-)2-3 mm thick, 

adaxially flattened to concave, winged, glabrous; tuftdomatia 

always present, only at axil of primary with 

secondary veins, tuft of hairs departing from the veins 

and converging toward the center of the axil, hairs 0.4- 

0.6 mm long, golden-yellow (sometimes the hairs 

dense, straight, velutinous, white, starting before axil). 

Inflorescences 2 per node, corymbose with lax opposite 

to subopposite decussate branches, with small, 

very compacted cymules at distal end of each branchlet, 

not forming a continuous flowering plane but a rather 

sparse inflorescence, (10-)13-18(-22) x 7-13.5 cm, 

lateral branches (2-)3-4 pairs, basal portion of axis 

not branched (5-)7-13 cm long; L/A = ca. 2:1; rachis 

terete to decussately compressed, rachis and branches 

sparsely short-puberulent; distal bracts usually absent 

or deltoid to narrowly triangular, to 2 x 0.5 mm; 

bracteoles subtending flowers absent. Flower sessile 

to short-pedicellate; pedicels to 0.7 mm long, glabrous 

to sparsely puberulent; hypanthium obconical, 0.8- 

1.4 x 0.9-1.3 mm, glabrous, reddish brown. Calyx 

extremely reduced, truncate to obviously lobed, 0.3- 

0.6 x 1-2 mm, glabrous; lobes absent to rounded, to 

0.3 mm long. Corolla cyathiform, deeply lobed, 1.6- 

2.4 mm long, cream-white to greenish white (turning 

pale yellow at maturity); tube short cylindrical, 0.8- 

1 x 1.2-1.5 mm, glabrous outside and inside; lobes 5, 

0.8-1.3 x 1-1.6 mm, ovate to oblong-ovate, rounded 

to obtuse at apex, glabrous outside, with a dense tuft 

of white-villous hairs at basal zone, with medial and 

distal zones glabrous inside. Stamens 5, exserted well 

above the corolla, equal, attached ca. 1 mm from the 

base of the tube; filaments 2.4-3.7 mm long, terete, 

basally flattened, connate to throat, with a tuft of dense 

white-pilose hairs at base; anthers elliptic, 0.5-0.7 x 

0.4-0.5 mm, dorsifixed at medial zone, base rounded, 

apex rounded to obtuse-mucronate. Pollen exine fo- 

veolate-reticulate. Style exserted, 2-2.8 mm long, as 

long as the corolla, glabrous, minutely costate to mi- 

nutely winged; style branches extruding as two lips 

above the corolla before anthesis, rounded to oblate, 

0.2-0.3 mm long, stigmatic surface microscopically 


subglobose, obtuse at base, rounded to hemi-ovoid at 

apex, 1.8-2.3 x 1.5-2 mm, dark brown without len- 

ticels, glabrous throughout; disk black, exceeding the 

calyx; disk loculicidal dehiscence present in old cap- 

sules. Seeds 0.7-0.95 x 0.5-0.75 mm. 

Distribution (Fig. 73) and ecology. Restricted to 

cloud forests on steep slopes, usually in rich mixed 

evergreen forest, 1000-1500 m (500 m in Guatopo 

National Park), of the coastal Cordillera of northeast- 

ern Venezuela (states of Carabobo, Aragua, Lara, and 

Miranda). Flowering specimens were collected only 

in January, while specimens at late blooming-early 

fruiting stage have been collected in March, Septem- 

ber, and October. Fruiting specimens were collected 

March, May, July, October, and November. 

Specimens examined. VENEZUELA. ARAGUA: Parque 

Nacional Henry Pittier, Quebrada Palo Vaco, trail up Periquito 

Parque Nacional Henry Pittier, 1300-1400 m, 25 Oct 

1961 (fr), Steyermark 89885 (NY, VEN[2]); Parque Nacional 

Henry Pittier (Rancho Grande), 16 Jul 1963 (fr), Uferide 13 

(VEN); rd. Rancho Grande-Gran Regresiva, 18 May 1984 

(fr), Field 424 (K[2]); Rancho Grande, S slope, 1000 m, 8 

Jan 1937 (fl), Pittier 13854 (F, NY, US, VEN); Rancho 

Grande, N slope, 1100 m, s.d. (fl), Pittier 14138 (F, NY, US, 

VEN); vic. of Alto de Rancho Grande, 29 Oct 1946 (fl-fr), 

Pittier 15275 (US, VEN); without locality, 30 Dec 1946 (fl), 

Lasser 2264 (VEN). CARABOBO: Between La Toma and 

"Capuchinos," S of Borburata, above Rio San Gian, 750- 

1000 m, 28 Mar 1966 (fr), Steyermark & Steyermark 95211 

(F, NY, U, US). FALC6N: Dto. Acosta, Mun. Jacura, Cerro 

La Mina, 350-650 m, 17 Feb 1961 (fl), Ruiz Teran 488 

(VEN[2]). MIRANDA: Parque Nacional de Guatopo, 48 km 

NNW of Altegracia, 500 m, 11 Sep 1960 (fl-fr), Steyermark


0~~~~ 

~~~~~~~~~~ 

' It 

FI.72 hiarhssecoa AF rm iter185,NY fo ah 33,NY.A.Hbt finlrecnc n 

mature!leaf B. Flower wih exserted anthers. C. ariation in calyx lobes.D. Detail of corolla, iside view. E. Stamen. F 

A 

o i. 

,,.~\:: ? 

:..:' 

'~..__.....~~ 

~~' '? ~ ~ ??? 

FIG. 72 Chimarhis speiosa (AF from ittier 3854, Y; G frm dahn 339, NY. A. Hait of iflorescnce an 

Apical portion style. of 

G. Mature capsule.~;T;'


*6' 

FIG. 73. Distribution of Chimarrhis microcarpa (circles) 

and C. speciosa (stars). 

87104 (NY, US, VEN). WrrmoIT LOCAurT: Aguacatal, 1000 

m, Jul 1919 (fr), Jahn 1339 (NY). 

Local name. Venezuela: santonino (Jahn 1339). 

This species is unique in having the smallest corolla 

(1.6-2.4 mm long; see Fig. 72D) in the genus 

(followed by C. parviflora of Central America), shortcyathiform 

corolla tubes, large laxly branched inflorescences 

(Fig. 72A), with irregularly separated distal 

units of compacted cymules, and semi-succulent 

leafy branchlets. 

Chimarrhis speciosa is closest to C. microcarpa 

in having capsules similar in size and shape (subsessile 

in the former), glabrous disks, and leafdomatia as tuft 

of pilose hairs at the vein axils; the former differing 

from the latter in being much larger (rain forest 

canopy) trees with bigger buttresses, longer corollas, 

smaller inflorescences, thinner woody leafy branchlets, 

and longer pedicels. The two species also occupy 

two different altitudinal ranges: C. microcarpa occurs 

in lowland (50-500 m) rain forests (on lateritic soils) 

of northeastern South America, and C. speciosa is 

restricted to steep-sloped cloud forests (1000-1400 m) 

of the Venezuelan coastal Cordillera. 

As already noted by Steyermark (1965, 1974), 

there are several collections that are transitional between 

C. microcarpa and C. speciosa, which were 

collected in the state of Miranda (where the two species 

come into contact) at intermediate elevations (50- 

500 m) in Guatopo National Park and Cerros del 

Bachiller. These specimens have transitional morphological 

characters: inflorescences not laxly branched 

and of intermediate length, leafy branchlets of intermediate 

thickness, capsules short-pedicellate (neither 

subsessile nor long-pedicellate). None of these tran- 

sitional specimens were collected in flower, so the 

dimensions and shapes of their corollas are unknown. 

Steyermark 87104 from Guatopo, which was reported 

as transitional by Steyermark (1965, 1974), is a good 

representative of C. speciosa. The transitional speci- 

mens between Chimarrhis microcarpa and C. 

speciosa are as follows: VENEZUELA. MIRANDA: 

Parque Nacional Guatopo, rd. Santa Teresa- 

Altagracia de Orituco, Rio Santa Cruz, 520 m, 23 Nov 

1961 (fr), Steyermark 89961 (NY, US[2], VEN); 

Bosque del Guatopo, 450-600 m, 29 Nov 1956 (fr), 

Bernardi 5943 (G[2], NY, VEN); Cerros del 

Bachiller, S of Santa Cruz, 10009'N, 65?48'W, 20-65 

m, 16 Mar 1978 (fr), Steyermark & Davidse 116278 

(MO, NY), 20-26 Mar 1978 (fr), 116885 (MO, VEN). 

CHIMARRHIS JACQUIN SUBGEN. 

PSEUDOCHIMARRHIS 

Chimarrhis Jacquin subgen. Pseudochimarrhis 

(Ducke) Delprete, subgen. et stat. nov. 

Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio 

de Janeiro 3: 255. 1922 (and Arch. Jard. Bot. 

Rio de Janeiro 4: 177, pl. 23. 1925). Type spe- 

cies. Pseudochimmarhis turbinata (A. P. de Can- 

dolle) Ducke (= Chimarrhis turbinata A. P. de 

Candolle). 

10. Chimarrhis turbinata A. P. de Candolle, Prodr. 

4: 404. 1830. Pseudochimarrhis turbinata (A. P. 

de Candolle) Ducke, Arch. Jard. Bot. Rio de 

Janeiro 3: 255. 1922. Chimarrhis duckei Rizzini, 

nom. superfl., Rev. Bras. Biol. 7: 277. 1947. Type. 

French Guyana. Ile de Cayenne, date unknown, 

Patris s.n. (lectotype, G-DC, here selected). 

Figs. 3Q, 10C, 18D, 58L, 74A-F, 75A-E, 

76A,B, 77A 

Elaeagia brasiliensis Standley in Gleason & Smith, Bull. 

Torrey Bot Club 60: 395. 1933. Type. Brazil. Para: 

Tapajos River region, Boa Vista, Fordlandia, 6 Sep 

1931 (fr), Krukoff 1018 (holotype, NY; isotypes, 

K, U). 

Pseudochimarrhis difformis (Benoist) Benoist, Arch. 

Bot. 5 (Mem. 1): 264. 1933. Bathysa difformis 

Benoist, Bull. Mus. Hist. Nat. Paris 26: 185. 1920. 

Type. French Guiana. Vic. of St-Jean-du-Maroni, 8 

May 1914 (fl-fr), Benoist 1190 (holotype, CAY; 

isotype, P).


g1 

FIG74Cimrrhsurbnaa(-EroIin a544,N; from Mo&Cardoso1712,GH). 

FIG. 74. Chimarrhis turbinata (A-E from Irwin et al. 55443, NY; F from Mori & Cardoso 17128, GH). A. Habit of 

inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style (note short antrorse 

pubescence). F. Mature capsule.


Trees to 40 m tall, to 170 cm dbh, bole irregular, 

much-branched at crown, fluted trunk, with buttresses 

to 3 m tall and extending several meters from trunk; 

bark deeply sulcate, lamellate, grayish white; wood 

yellow, very hard. Leafy branchlets (4-)7-11 mm 

thick, semi-succulent to woody, quadrangular to 

decussately compressed, terminal 2-3 nodes sparsely 

puberulent to densely pubescent; older branches 

glabrate, exfoliating in narrow strips, showing the redbrown 

bark just below; lenticels not found. Stipules 

free to connate at base, more or less contorted in bud, 

often adnate to petioles, deltoid to narrowly triangular, 

strongly acuminate, golden-pubescent outside, 

often with ciliate margins, glabrous with sparse small 

colleters at base inside, 9-20 x 6-8 mm, subcaducous, 

reddish green, leaving a scar encircling the stem, adnate 

to the petiole scar 1-3 mm wide. Leaves 9-19 

(-33) x 6-10.5(-16) cm, L/W = 1.5:1 to 2:1, oblong 

to elliptic to obovate, acute (rarely decurrent) to obtuse 

at base, obtuse to shallowly acute at apex, acumen 

usually absent; dark green, often shiny above, 

pale green-yellowish below, thick-chartaceous to 

semi-coriaceous; drying olive-green above, rustbrown 

below, thick-chartaceous to leathery; glabrous 

above, glabrous to puberulent (rarely pubescent) below; 

primary and secondary veins glabrous to puberulent 

(rarely pubescent), prominent below; secondary 

veins 8-12(-16) each side; tertiary veins alternatively 

subparallel and reticulate, quaternary veins finely reticulate, 

very distinctive from other species; petioles 

4-15(-22) mm long, 1-2.5 mm thick, terete to adaxially 

flattened or concave, (glabrous-) puberulent to pubescent; 

domatia absent or present, usually a tuft of 

sparse yellow hairs, puberulent to pubescent (Fig. 

57L). Inflorescences 2 per node, corymbose, laxly 

branched, with opposite to subopposite decussate 

branches, cymules lax to fairly compacted at distal end 

of each branchlet, (8.5-)11-13.5(-32) x cm long. Corolla narrowly funnelform, deeply lobed, 

the lobes reflexed at anthesis, 3.5-6 mm long, white 

to yellowish white; tube long-cylindrical, 1.5-3 x 0.7- 

1 mm, glabrous outside and inside; lobes (4-)5, 2.5- 

3.5 x ca. 0.5 mm, oblong with acute apex and round 

tip, glabrous outside, with a basal area pilose to tomentose 

(ca. 1 mm wide), with distal zone glabrous 

inside. Stamens 5, exserted well above the corolla, 

equal, attached 1-2.5 mm from the base of the tube; 

filaments 3.5-5.5 mm long, terete, minutely costate, 

basally flattened, connate to throat, sparsely white 

long-pilose at base, the hairs decreasing in length toward 

medial zone; anthers elliptic, 0.8-1.5 x 0.3-0.5 

mm, dorsifixed near medial zone, base rounded, apex 

rounded or minutely mucronate. Pollen exine foveolate-reticulate. 

Style exserted, 4-6 mm long, as long 

as the corolla, terete, densely antrorse-strigose; style 

branches extruding as two lips above the corolla before 

anthesis, oblate to oblong, 0.3-0.5 mm long, stigmatic 

surface microscopically (40x) papillose; ovaries 

often infested by insect, causing production of 

pyriform fruit-galls. Capsules turbinate, acute at base, 

rounded at apex, 3.5-6 x 2.5-3.3 mm, rust-brown, 

without lenticels, sparsely to densely puberulent below 

the disk; disk densely white-pilose, not exceeding 

the calyx; disk loculicidal dehiscence present in 

old capsules. Seeds 0.75-1.95 x 0.75-1.25 mm. 

Distribution (Fig. 77A) and ecology. Primary and 

disturbed non-inundated rain forests (terra firme), and 

riverine forests, in lateritic soils, clay to sandy-clay 

soils, lowland to mild slopes, 50-400 m, of Guyana, 

Surinam, French Guiana, and Brazil. Flowering specimens 

were collected in March, June, July, August, 

September, and October. Fruiting specimens were 

collected in all the months from June through Janu- 

5-10(-18) 

cm, lateral branches 1-2(-4) pairs, basal portion of 

axis not branched 5-8(-15) cm long; L/A = 2:1 to 

2.8:1; rachis basally terete, distally decussately compressed, 

rachis and branches sparsely to densely yellow-puberulent; 

distal bracts from reduced deltoid 

scales 0.2-0.3 mm long to linear bracts to 7 x 1-2 mm, 

sometimes leaf-like, to 7 x 2.7 cm, glabrous to puberulent, 

the margin sometimes ciliate; bracteoles 

subtending flowers absent. Flowers sessile to shortary. 

Boom and Campos (1991) reported that C. 

turbinata "is ecologically the most important Rubiaceae 

in the BDFF [Biological Dynamics of Forest 

Fragments; located north of Manaus] reserves" and 

that its "distribution... suggests a tendency of clumping. 

Geographic distribution is mostly eastern Amazonian; 

this species' occurrence in this study are the 

westernmost records we have seen. Ecologically, it 

is restricted to terra firme, at elevations of up to 350 

m. Flowering: Jun-Oct. Fruiting: Jun-Jan." 

pedicellate; pedicels to 1.5 mm long, densely puberulent; 

hypanthium obovoid-turbinate, 1.5-2.5 x 1-1.6 

mm, densely puberulent. Calyx extremely reduced, 

truncate to short-lobed, 0.4-0.7 x 1-1.6 mm, glabrous 

to sparsely puberulent; lobes shallowly triangular, to 

0.2 mm long; rarely in some flowers one of the calyx 

lobes enlarged into a pale-green calycophyll, to 1.5 

Specimens examined. GUYANA. Gunns Strip, 

Essequibo River, 01?39'N, 58?38'W, 240-260 m, 30 Sep 

1989 (fr), Jansen-Jacob et al. 1899 (B, U). 

SURINAM. Forest reserve, section 0, tree n. 577, Forestry 

Bureau 1151 (U), 3 Jun 1916 (fr) 2040(U), 7 Sep 1916 

2415 (A, BR, IAN, U), 31 Oct 1916 (fr) 2464 (G, MO, U),


FIG. 75. Leaf and inflorescence variation in Chimarrhis turbinata (A from Boom et al. 8756, NY; B and D from Schultz 

7289, G; C from unknown collector 2907 [Surinam], MO). A, B. Inflorescence range. C-E. Leaf range. All drawn to the 

indicated scale bar.


11 Jan 1917 (fr) 2585 (K, NY, U, US), 5 Jun 1917 (fl) 2907 

(G, MO, U), 10 Oct 1920 (fl) 4952 (NY, U, US); vie. of Camp 

8, 17 Aug 1955 (fl), Schulz 7289 (G, NY, U); Komaramaraballi, 

1 Nov 1944 (st), Stahel 261 (A[2], B, MO, NY, U), 

Jun 1945 261a (U); Lucie River, below confluence of Oost 

River, 03020-32'N, 56?26-49'W, 225 m, 7 Sep 1963 (fl), 

Irwin et al. 55443 (F, GH, MO, NY, US); vic. of Nassau, 18 

Mar 1949 (st), Lanjouw & Lindeman 2810 (U). 

FRENCH GUIANA. CAMOPI: Trois Sauts, Mitake, 18 

Sep 1977 (fr), Grenand 1500 (CAY). CAYENNE: Approague 

River, confluence with Arataye River, Parare waterfalls, 25 

Nov 1985 (st), Barrier 5060 (CAY); Orapu, rd. to Fourgessi6, 

km 8,200 m, 15 Oct 1966 (fl), Oldeman B-674 (CAY, 

P); Sinnamary River, Petit Saut, 05?03'N, 53?03'W, 26 Jul 

1988 (fr), Sabatier & Prevost 2116 (CAY, US); Sinnamary 

River, above Petit Saut, between Petit Crique and Plomb and 

Crique Tigre, 87 m, 05?00'N, 53001'W, 6 Sep 1993 (st),Mori 

et al. 23645 (NY); Sinnamary-Counamama River Basin, 

Piste de Saint-Elie, 05020'N, 53?00'W, 13 Aug 1991 (st), 

Sabatier & Prevost 3731 (CAY, US). MARIPA-SOULA: Maroni 

River Basin, Grand Inini River, 03?40'N, 53?50'W, 19 Jul 

1990 (st), Sabatier & Prevost 3356 (CAY). SAOL: Bellevue, 

250 m from bridge, left shore of Crique Bellevue, 1.2 km 

from Me Madere Victor, 6 Dec 1956 (fr), Bena 1262 (CAY); 

Gallion, km 30, Cascade, 13 Sep 1956 (fl), Bena 1187 

(CAY); rd. to Belizon, between Eaux Claires and Layon 

Biche, 03037'N, 53?12'W, 200-400 m, 4 Aug 1993 (fl), Mori 

et al. 23098 (CAY); Montagne du Chevaux, near Route de 

l'Est, 04?45'N, 52?28'W, 50 m, 27 Jun 1988 (fr), Gentry & 

Morawetz 63183 (CAY, MO); Limonade, ORSTOM, 15 Oct 

1977 (fr), Granville 5377 (CAY, U, US); rd. Charvein- 

Acarouany, km 1, 11 Mar 1954 (fl-fr),BAFOG 166M(CAY, U). 

BRAZIL. AMAPA: Rio Araguari, 01009'-26'N, 51?52'- 

Isabel, Jun 1908 (fl), Museu Goeldipersonnel 9425 (F, G), 

9713 (F, G, NY); Belem, Bosque Municipal, 17 Jun 1943 

(fl), Ducke 1235 (MG19119) (IAN, MG); Belem, Bosque 

Municipal, 16 Jul 1947 (fl), N. T. Silva 51 (B); Belem, 

Utinga-Providencia, 20 Aug 1914 (fr), Ducke s.n. 

(MG15757) (MG); Belem, 31 Aug 1922 (fr) and 23 Jun 1923 

(fl), Ducke s.n. (RBI 7356) (RB, U, US); Belem, Lago Catu, 

10 Sep 1914 (fr), Ducke s.n. (MG15480) (MG); Belem, 

Reserva do Mocambo, 16 Aug 1982 (fr), Ramos & Rosario 

15 (MG); Itacaiunas, affluent of Rio Tocantins, Serra 

Buritirama, 05?31'S, 50013'W, Sep 1970 (st), Pires & Belem 

13006 (IAN); Belterra, 26 Jun 1947, Black F8 47-955 (US); 

Melgaco, Reserva Nacional de Caxiufla, site of the Ferreira 

Penna Scientific Station, 2-15 Feb 1991 (st), Chagas et al. 

2217 (MG); Mun. Oriximina, Serra da Preciosa, 21 km N of 

Rio Cumina-Mirim, 130 m, Martinelli 6835 (NY); Carajas, 

Serra Norte, Maraba, 7 Aug 1982 (fr), Maciel et al. 776(IAN, 

MG, NY); Belem, Mata da Utinga, 15 Dec 1945 (st), Fr6es 

20777 (IAN, NY); rd. Belem-Brasilia, km 2, 28 Sep 1959 

(fr), Kuhlmann & Jimbo 310 (IAN, TEX), 329 (IAN, TEX); 

Planalto de Santarem, Currupiru, loc. Gato, 22 Aug 1954 (fr), 

Fr6es 31049 (UB); Santar6m, km 70 on rd. to Cachoeira do 

Palhao, of Rio Curua Una, 5 Oct 1966 (fr), Cavalcante & 

M. Silva 1598 (IAN, MG); Santar6m, km 35 on Palhao rd., 

Igarape Curupira Camp, 28 Aug 1969 (fr), M. G. Silva & 

Rosdrio Souza 2420 (MG, US), 2 Sep 1969 (fr), 2475 (CAS, 

MG, US); Mun. Mez, confluence of Rio Pinaticaua and Rio 

Jaracu, 19 Sep 1955 (fr), Fr6es 32097 (UB); Mun. Curralinho, 

between Rio Mapua and Rio Piria, 18 Jul 1950 (fl), 

Black et al. F8 50-9831 (IAN); Moju, rd. to Acara, km 60, 

15 Aug 1975 (fr), N. T. Silva 3924 (IAN); Fazenda Uriboca, 

Pirelli, Jun-Aug 1958 (fr), Pires 6826 (UB), 6922 (UB), 

6969 (UB), 6978 (UB), 7107 (UB). 

58'W, 11 Sep 1961 (fl), Pires et al. 50853 (G, GH, IAN, M, 

MG, U, US); Serra do Navio, 26 Sep 1961 (fl), Pires et al. 

51224 (IAN, NY, U, US); rd. Calcoene-Oiapoque, 60 km 

SSE of Oiapoque, 03? 18'N, 51039'W, 2 Dec 1984 (fr), Mori 

Local names and uses. French Guiana: bois 

pagaie (Bena 1262, Granville B5377), bois chapelle 

(Bena 1262),citronelle (BAFOG 166-M), samaati- 

& Cardoso 17128 (F, GH, MG, MO, NY, US); Rio Oiapoque, 

3 km E of mouth of Rio Matura, 02?34'N, 52?31'W, 22 

Sep 1960 (fl), Irwin et al. 48440 (GH, IAN, MG, NY, U, 

US); Rio Jari, Monte Dourado, 9 Sep 1968 (fl), N. T. Silva 

905 (CAS, NY), 17 Sep 1968 (fl), 999(NY, US), 17 Oct 1968 

(fr), 1246 (NY); Rio Jari, Planalto A, km 11, 2 Aug 1969 

(fl), M. Silva 2568 (IAN, NY, TEX, UB); Porto Platon, Perimetral 

Norte, Cupixi, 14 Sep 1976 (fl), M. Silva 2794 

(INPA); Rio Jari, Monte Dourado, 4 Dec 1967 (fr), Oliveira 

3836 (NY), 29 Jan 1968 (fr), 4036 (NY); Camaipi, 

EMBRAPA Reserve, 0010'N, 51037'W, 3 Sep 1983 (fr), 

palioudou (BAFOG 166-M), citronelle rouge (commercial 

name, Bena 1262, BAFOG 1262). Surinam: 

samaati-palioudou (Paramaka, BAFOG Service 

Forestiere 7608), walalu (Way pi, Grenand 1500), 

sinjahoedoe (Sur., Lanjouw & Lindeman 2810), koemaramaraballi 

(Arow., Lanjouw & Lindeman 2810), 

kaire-e (Car., Lanjouw & Lindeman 2810), sienjahoedoe 

(Schultz 8581). Brazil: pau de remo (Para, 

Irwin et al. 48440, Oliveira 3854, Pires 48902)), ramal 

do pau rosa (Para, Silva & Rosario 3732), carapa- 

Mori et al. 15786 (MG, NY[2]). AMAZONAS: Rd. Manaus- 

Caracari, km 146,25 Sep 1973 (fr), Bisby et al. P18095 (US); 

Mun. Manaus, rd. ZF-3, km 24, Reserva N0 1202, 17 Jun 

1980 (fr), Lima & Zimmerman 519 (NY, U); rd. Manaus- 

Itacoatiara, km 165, 16 Oct 1965 (st), Rodrigues 8098 

(INPA), km 140, 4 Oct 1965 (st), 8099 (INPA), km 125, 4 

Feb 1970 (st), Rodrigues 8692 (INPA); rd. between Rio 

Araqa and Lago Castanho, toward Careiro, 13 Jul 1972 (fl), 

M. Silva 5073 (US). MARANHAO: Rio Alto Turiacu, Nova 

Esperanca, 02?55'S, 45?45'S, 29 Nov 1978 (fr), Jangoux & 

Bahia 86 (MG, NY, UB); PARA: Rd. Belem-Braganca, Santa 

nauba (Para, Black F8-4 7-955), carapanauba cinzeiro 

(Para, Black F8-47-955), paraka'y (MaranhAo, Balee 

& Ribeiro 431), paraku-'iran-'y (Maranhao, Balee & 

Ribeiro 2826). 

This species offers pale-yellow hard wood of first 

quality for construction of houses. In French Guiana, 

the Saramacas use the wood to make paddles, toys, 

combs, wooden spoons and benches, and as construction 

wood for various purposes; the bark is also used 

by the same tribe to make ropes and floral artifacts

174 

A 

? 

.? 

. 

. 

iii.=izi'i' 

B ? .. . 

!f ' 

t..... 

*D?? L ,,: ? . i 

A''! 

.....:; i?" ''?.. 

B ~~~X.~~S~B~ i~ ~ .;t~, 

FIG. ?6. Chiman'6i~ 

' 


turbina~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~a. 

A. Inflorescence. B. Trunk with bnsnl Boom.)~~~~~~~~~~~~~~~~sf~i 

buttresses. (Photog~~~~~~~~~~~~~~~~~~~~ 

by Brinn


for decoration of their houses. Various indigenous 

tribes in Brazil and in the Guianas use the plank wood 

from the buttresses (without cutting down the tree) to 

make paddles, from which the common name is derived 

["Pau de remo" (Portuguese), "Bois Pagaie" 

(French) = fore it belonged to the Cinchoneae. Bremekamp 

(1934) considered this separation unacceptable and 

returned them to Chimarrhis. Pseudochimarrhis is 

here treated as a subgenus of Chimarrhis, characterized 

by persistent, deltoid stipules. 

paddle wood]. 

Benoist (1920) described Bathysa difformis (using 

his collection from French Guiana), which he later 

This species is a frequent canopy tree, and is rec- (Benoist, 1933) transferred to Pseudochimarrhis. 

ognizable by its large, turbinate capsules (hence the Although this name is based on material with immaname), 

pubescent disk not exceeding the calyx, and ture capsules, it is clear that these specimens belong 

ascending-puberulent slender style. The leaf blades to C. turbinata. 

and the inflorescences are very variable in size and The trunk of C. turbinata (Fig. 76B) is usually irshape 

(Fig. 75A-E) even within the same individual. regular, alveolate and fluted, with bark deeply sulcate, 

In some inflorescences of C. turbinata (Mori & rust-brown and fibrous, sapwood pale-yellow, and 

Cardoso 17128; Pires et al. 51224) the distal bracts heartwood dark brown. Color photographs of this 

subtending secondary branches are foliose, delicate, species can be seen in Parrotta et al., 1995 (306-307). 

and pale-green, which might be superficially confused 

with calycophylls, but these are growing at the inflo- Dubious collection: FRENCH GUIANA. 

rescence nodes, or more rarely at the tips of inflores- CAMOPI: Trois Sauts, Aft iti, 17 Apr 1976 (fr), 

cence branches (but not calyx lobes, and with pinnate Grenand 1238 (CAY[3]). The young branches are 

venation). Also, as observed by Ducke (1922), very sterile and the leaves very much resemble those of 

rarely one of the calyx lobes expands into small, pale C.turbinata; the infructescences are detached and 

green calycophyll (with palmate venation). probably collected from the forest floor. The fruits are 

Leafvestiture in this species is rather variable, but much smaller than those of C. turbinata, and similar 

in most cases the blades are glabrous. In a few col- to those of C. microcarpa. Since both species occur 

lections (Boom et al. 8756, Mori et al. 16248, Balee in this area, it is uncertain whether these specimens 

& Ribeiro 2826, and Pires 48902) the blades are 

represent a mixed collection or a chance hybrid bedensely 

golden soft-hirtellous (velvety to the touch) tween these two species. 

beneath, exceptionally large and often asymmetrical 

(see below). Domatia in C. turbinata are absent (on 

the pubescent blades a few short hairs are present at Deviating collections. A few collections (see bethe 

axils). 

low) have distinct morphological traits, as compared 

Chimarrhis turbinata is similar to C. barbata in with typical C. turbinata, having stipules free at base, 

having large capsules and nonglabrous styles; C. bar- not contorted in bud; obovate capsules (2.5-)3.5-8 

bata differs from the former in having smaller flowers, 

smaller anthers, ascending-pubescent thicker and 

shorter styles, and capsules with minute-puberulent 

disks exceeding the calices. 

Chimarrhis turbinata was first described by A. P. 

de Candolle (1830) based on material collected by 

Patris in Cayenna (French Guiana). One specimen 

preserved at Geneva (G, Candolle Herbarium) has 

been found and selected to be the lectotype. Elaeagia 

brasiliensis, as already pointed out by Steyermark 

(1965), is synonymous with C. turbinata. Rizzini 

(1947), declaring C. turbinata of dubious identity, 

proposed C. duckei (a superfluous name), which is 

here treated as synonymous with C. turbinata. 

Ducke (1922) separated the genus Pseudochimarrhis 

from Chimarrhis so as to include C. turbinata and 

C. barbata [as P. barbata], stating that the general 

morphology of the flowers and the vertically inserted 

seeds resembled those of Calycophyllum, and there- 

x 

2-5 mm, golden-puberulent; uncommonly large 

(23.5-34 x 10-16 cm, L/A = 2:1) often asymmetrical 

leaf blades, densely golden-hirsute; and uncommonly 

large inflorescences (ca. 33 cm long). Flowers, pollen, 

and seeds of these specimens are still unknown, 

and future collections might show that these samples 

deserve nomenclatural recognition. 

Specimens examined. BRAZIL. AMAPA: Camaipi, 

EMBRAPA reserve and vic., 00?10'N, 51?37'W, 17 Sep 

1983 (st), Mori et al. 16248 (NY). AMAZONAS: Distrito 

Agropecuario, Reserva 1501 of the WWF/INPA Project, 50- 

125 m, 6 Dec 1988 (fr), Boom et al. 8756 (INPA, K, NY, 

US); Distrito Agropecuirio da SUFRAMA, Mun. Manaus, 

rd. BR-174, 02?25'S, 59?51'W, 13 May 1985 (fr), INPA/ 

WWF 1109.25 (NY). MARANHAO: Mun. Carutapera, Gurupiuna, 

Ka'apor Indian Reserve, affluent of Rio Gurupi, 

02?43'S, 46026'W, 4 Nov 1986 (st), Balee & Ribeiro 2826 

(NY). PARA: Belem, grounds of Instituto Agronomico do 

Norte, 1 Nov 1960 (st), Pires 48902 (NY, SP).


11. Chimarrhis barbata (Ducke) Bremekamp, 

Recueil Trav. Bot. Neerl. 31: 260. 1934. 

Pseudochimarrhis barbata Ducke, Arch. Jard. Bot. 

Rio de Janeiro 4: 177, pl. 23. 1925. Type. Brazil. 

Para: Tapajoz River, between Boim and Serra 

Humayta, 8 Apr 1924 (fl-fr), Kuhlmann s.n. (RB 

17384) (holotype, R; isotypes, B*, U, US; photo- 

B at MO, F, NY). Figs. 77B, 78A-F 

Trees to 30 m tall, bole irregular; bark deeply fissured; 

wood yellow, very hard. Leafy branchlets 2- 

4 mm thick, thin-woody, glabrous to sparsely puberulent; 

older branches glabrate, exfoliating in narrow 

fibrous strips, dark brown; lenticels punctiform to lin- 

ear, to 2 mm long. Stipules connate at base, adnate to 

the petioles, not contorted in bud, deltoid to narrowly 

triangular, mucronate, puberulent outside, often with 

ciliolate margins, glabrous and with sparse colleters 

at base inside, 4.5-15 x 4.5-7 mm, subcaducous, leaving 

a scar encircling the stem. Leaves 6-16 x 4-7 cm, 

L/W = 1:1 to 2:1, elliptic, obovate, to widely ovate, 

acute to obtuse at base, acute to obtuse at apex, sometimes 

short mucronate, semi-coriaceous; glabrous 

above and below, drying olive-green; primary and 

secondary veins glabrous, prominent below, secondary 

veins 6-10 each side; tertiary veins subparallel and 

reticulate; petioles 6-20 mm long, 1-3 mm thick, 

adaxially flattened or concave, glabrous to minutely 

puberulent; domatia absent or few sparse hairs present. 

Inflorescences 2-4 per node, corymbose, short pedunculate, 

5-22 x 4.5-12 cm, lateral branches one 

pair; rachis basally terete, decussately compressed 

distally; L/A = 2.1:1 to 2.5:1; rachis and branches 

sparsely puberulent; distal bracts extremely reduced, 

1-5 x ca. 1 mm, linear to deltoid, rarely foliose, to 

1.5 cm long, narrowly elliptic; bracteoles subtending 

flowers deltoid scales ca. 0.5 mm long. Flowers 

sessile to short-pedicellate; pedicels to 0.5 mm long, 

sparsely short-puberulent; hypanthium obconical, 

1.5-2.5 x 1-1.5 mm, golden short-puberulent. Calyx 

extremely reduced, truncate to short-lobed, 0.5-1 x 

1.5-2.5 mm, glabrous to sparsely puberulent; lobes 

absent to shallowly triangular, to 0.5 mm long; rarely 

one or two flowers per inflorescence with one calyx 

lobe expanding into a small greenish white 

calycophyll. Corolla funnelform, deeply lobed, the 

lobes reflexed at anthesis, 3-4.5 mm long, white to 

greenish white; tube short cylindrical, 1-1.8 x 1-2 mm, 

glabrous outside and inside; lobes 5, 2-3.5 x 0.7-1.5 

mm, oblong with round apex, glabrous outside, with 

a basal area white pilose and distal zone glabrous inside. 

Stamens 5, exserted well above the corolla, 

equal, attached 1-1.5 mm from the base of the tube; 

filaments 2.5-3 mm long, terete, basally flattened, 

densely white pilose at base; anthers elliptic, 0.8-1 x 

0.3-0.5 mm, medially dorsifixed, base rounded, apex 

rounded or minutely mucronate. Pollen exine foveolate-reticulate. 

Style exserted, 4-5 mm long, terete, 

densely strigose, appressed-retrorse; style branches 

protruding as two lips above corolla before anthesis, 

oblate to oblong, reflexed, 0.6-1 mm long, stigmatic 

surface minutely papillose. Capsules pedicellate, 

pedicels to 2 mm long, narrowly turbinate, acute at 

base, rounded at apex, 5-9 x 2.5-5 mm, rust-brown 

to black-brown, without lenticels, sparsely goldenpuberulent 

below the disk; disk short white-puberulent, 

obviously exceeding the calyx; disk loculicidal 

dehiscence not seen. Seeds 1-1.5 mm long. 0.8-1.25 

mm wide. 

Distribution (Fig. 77B) and ecology. Primary, 

secondary, and fragmented non-flooded Amazon for- 

est (terra firme), in the Brazilian states of Para (Rio 

Tapajoz, Boim, and Itaituba) and Amazonas (Manaus, 

Tef6, and Villa Braga). Flowering specimens were 

collected in October, November, and December. Fruit- 

ing specimens were collected in April and August. 

Boom and Campos (1991) reported that the distri- 

bution of C. barbata "within the BDFF reserves is 

sporadic. Geographic distribution is strictly central 

Amazonian. The species is rare and/or rarely col- 

lected. Ecologically, is known only from terra firme 

forest." Prance et al. (1976) cited C. barbata in the 

Rubiaceae encountered in a hectare plot 30 km north 

of Manaus. 

Specimens examined. BRAZIL. AMAZONAS: Villa 

Braga, 1924 (fr), Ducke s.n. (RB17358) (B*, NY, U, frag- 

F); Manaus, Mindu, 22 Oct 1929 (fl), Ducke s.n. (RB22847) 

(K, U, US); Manaus, Villa Municipal, 11 Nov 1931 (fl), 

Ducke 38 (Yale 20719) (A, F, US); Manaus, Villa Munici- 

pal, 26 Dec 1941 (fl), Ducke 860 (F[2], IAN, MG, NY, R[2]); 

Rio Negro, SAo Gabriel, 16 Feb 1936 (young fr), Ducke s.n. 

(RB35055) (K, RB); upper Rio Solimfes, Mun. Tef6, Villa 

Nogueira, Xelele, 02?18'S, 64?48'W, 10 Nov 1986 (fl), Daly 

et al. 4312 (NY); rd. Manaus-Itacoatiara, km 26, Reserva 

Florestal Ducke, 02?53'S, 59?58'W, Reserva Florestal 

Ducke, tree N. 3220-15, 26 Sep 1995 (fl), Sothers et al. 568 

(INPA 185102) (INPA, NY); Reserva Florestal Ducke, tree 

N. 3220-15, 9 Feb 1996 (fr), Campos et al. 468 (INPA 

185019) (INPA, NY); Reserva Florestal Ducke, tree N. 3357- 

15, 12 Oct 1995 (fl), Sothers et al. 625 (INPA 185105) 

(INPA, NY); Reserva Florestal Ducke, tree N. 3357-15, 17 

Jan 1996 (fr), Sothers et al. 750 (INPA 185110) (INPA, NY); 

Reserva Florestal Ducke, tree N. 4384-15, without date, 

Assuncao 369 (INPA); Itaituba, Parque Nacional do Tapaj6s 

(IBDF), km 85 on rd. to Itaituba, 14 Nov 1978 (fl), M.G. 

Silva & Rosdrio 3732 (MG). PARA: Santar6m, Reserva For-


so 70 s So / 0 

16 l 

0t 

hooker FIG. (circles), 77. Distribution C barba ofChimarrhis. (solid strs), A. C. glabrflora (op (circles), strs), C. turbinate C. (diamonds), (i and ls). C. brevipes (stars). B. C. 

FIG. 77. Distribution of Chimarrhis. A. C glabrflora (circles), C turbinate (diamonds), and C brevipes (stars). B. C 

hooleri (circles), C. barbata (solid slurs), C. dulckeana (open sic-s), and C. gentryana (Uriangles). 

estal Curuana (SUDAM), right margin of Rio Curuana, 

Barreirinha, 19 Aug 1988 (fr), Rosdrio et al. 852 (MG), 858 

(IAN, MG). 

Local name Brazil: pau amarelo (Prance et al., 1976). 

Chimarrhis barbata is distinguished by its large capsules 

with puberulent to minutely puberulent disks, which 

exceed the calices (Fig. 78F). Chimarrhis barbata is a 

rare species of large canopy-trees endemic to the Brazilian 

Amazon, with deeply fissured, fibrous, reddish 

brown bark, extremely hard pale-yellow wood, semileathery 

leaves, and cream-white to greenish white, 

very fragrant flowers visited by bees. Because of its 

scanty collections, this is probably the least known 

species of Chimarrhis from the Amazon basin. 

Chimarrhis barbata is similar to C. turbinata in 

having large capsules, the latter differing from the 

former in having capsules with pubescent disk not 

exceeding the calyx, longer corollas, larger anthers, 

longer and thinner style with shorter style branches. 

At least one specimen of C. barbata [Ducke s.n. 

(RB53055)] has flowers with obvious calycophylls, 

about 4.5 x 2 cm, and greenish white when fresh (ac- 

cording to the label-data). 

Chimarrhis barbata is easily distinguishable from 

C. duckeana by its smaller leaves that turn pale ol- 

ive-green (to light-brown) when dry, capsules with 

ovoid disks much exceeding the calices, longer and 

slenderer antrorse-pubescent style, slender leafy 

branchlets exfoliating in small pieces, and smaller 

flowers.


Ic 

FI. Chimrribabr(ADfoDue224,U;EfoDayeal43,N;FfoKhman18,U, 

8 

isoyp). . abi oinforscnce. . Fowr wthexsrtd nthrs C.Deailofcorll, nsie iew D Stme. E Syl 

(note long antrorse-pubescence). 

F. Mature capsule.?? 

FIG. 78. Chimarrhis barbata (A-D from Ducke 22847, US; E from Daly et al. 431, NY; F from Kuhlmann 17384, US, 

isotype). A. Habit of inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style 

(note long antrorse-pubescence). F. Mature capsule.


12. Chimarrhis duckeana Delprete, sp. nov., non 

oblong with round apex, glabrous outside, with basal 

Chimarrhis duckei Rizzini, nom. superfl., (based area densely white-pilose and distal zone glabrous 

on de Candolle's description of C. turbinata), inside. Stamens 5, exserted well above the corolla, 

Revista Brasil. Biol. 7: 277. 1947 . Type. Brazil. 

equal, attached ca. 1 mm from base of tube; filaments 

Amazonas: Manaus, Villa Municipal, non-flooded 4-5 mm long, terete, basally flattened, densely pilose 

secondary forest, 2 Jan 1930 (fl), Ducke s.n. (RB at base; anthers elliptic, 0.8-1 x 0.2-0.4 mm, medi- 

22846) (holotype, RB; isotypes, K, P, RB, U). ally dorsifixed, base rounded, apex rounded. Pollen 

Figs. 10B, O1F, 18E, 77B, 79A-F 

exine foveolate-reticulate. Style exserted, 2.5-3 mm 

long, minutely costate, densely retrorse white puberu- 

Arbores Chimarrhi barbatae similes, sed ab illa foliis lent; style branches protruding as two lips above comajoribus 

ambitu ellipticis ovatisve basi cordatis cordatisve, rolla before anthesis, ovate, reflexed, 0.3-0.5 mm 

brevius petiolatis, lamina in statu sicco nigrescenti (nec long, stigmatic surface minutely papillate. Capsules 

olivacea), stylo breviori crassiori minutim costato, secus 

pedicellate, pedicels to 2 mm long, turbinate to widely 

costas tantum puberulo (nec gracili tereti ubique retrorsus 

turbinate, acute at base, 

strigoso), necnon styli ramulis brevioribus diversa. 

oblong-rounded at apex, 6- 

12 x 4-6 mm, dark-brown, without lenticels, microscopically 

puberulent throughout; disk obviously exceeding 

the calyx; disk loculicidal dehiscence not 

seen. Seeds 1.15-2.38 x 0.83-1.5 mm. 

Trees 20-25 m tall, 30-55 cm dbh, mid-canopy 

trees; bark fissured, brown. Leafy branchlets 

decussately compressed, 4-6 mm thick, (4-)5-8 mm 

wide, semi-succulent, glabrous; older branchlets gla- 

brous, smooth (not exfoliating in narrow strips), gray- 

ish brown; lenticels linear, to 3 mm long. Stipules free 

or connate at base, not contorted in bud, narrowly tri- 

angular to deltoid, acuminate, glabrous outside (with- 

out ciliolate margins), glabrous and with sparse 

colleters inside, 7-19 x 7-10 mm, dark green, 

subcaducous to persistent. Leaves (8-) 18-21(-32) x 

(7-)10-18 cm, L/W = 1.1:1 to 1.2:1, widely elliptic 

to widely ovate, obtuse to cordate at base, rounded to 

obtuse at apex, drying dark-brown to black; primary 

and secondary veins glabrous, prominent below; sec- 

ondary veins 8-10 each side, widely spaced (2-3 cm 

intervein); tertiary veins subparallel, faintly evident 

above; petioles 25-43 mm long, 2-5 mm thick, ad- 

axially concave, glabrous; domatia absent. Inflores- 

cences 2-4 per node, corymbose, short to long pedun- 

culate, with opposite decussate branches terminating 

in dense cymules, (6.5-)12.5-22 x (3-)6-12 cm, lat- 

eral branches 1-4 pairs, basal portion of the axis not 

branched 5-14 cm long; L/A = 2:1 to 3:1; rachis ba- 

sally terete, decussately compressed distally, rachis 

and branches glabrous to minutely puberulent; distal 

bracts extremely reduced, 0.5-4 x 0.5-1 mm, deltoid 

to lanceolate (no foliose bracts observed); bracteoles 

subtending flowers deltoid scales ca. 0.5 mm long. 

Flowers pedicellate; pedicels 1-2 mm long, puberu- 

lent; hypanthium obconical, ca. 2 x 1.5-2 mm, pu- 

berulent. Calyx extremely reduced, short-lobed, 0.5- 

1.5 x 1.5-2.5 mm, glabrous to sparsely puberulent; 

lobes deltoid, ca. 0.5 mm long; flowers with 

calycophylls not found. Corolla funnelform, deeply 

lobed, the lobes reflexed at anthesis, 3-4.5 mm long, 

white; tube short-cylindrical, 1-1.5 x 1-1.5 mm, gla- 

brous outside and inside; lobes 5, ca. 2.5 x 1-1.5 mm, 

Distribution (Fig. 77B) and ecology. Primary, 

secondary, and fragmented forests in non-flooded 

Amazon (terra firme), in clay or sandy lateritic soil. 

Several specimens were collected in the surroundings 

of Manaus, but one specimen was collected on the Rio 

Negro. Flowering specimens were collected in January 

and February. Fruiting specimens were collected 

in March, July, August, September, and October. 

Specimens examined. BRAZIL. AMAZONAS: Rd. 

Manaus-Itacoatiara, km 26, Reserva Florestal Ducke, 

02?53'S, 59?58'W, parcel 22, tree N. 2938,22 Mar 1966 (fr), 

Rodrigues & Coelho 7600 (INPA 16903) (US); Reserva 

Florestal Ducke, tree N. 3553-15, 12 Dec 1995 (fl), Ribeiro 

& Assuncdo 1776 (INPA 185097) (INPA, NY); Reserva 

Florestal Ducke, 10 Aug 1976 (st), Oliveira s.n. (INPA 

60564) (INPA); Reserva Florestal Ducke, tree N. 1613 

(inventario florestal), 4 Sep 1968 (fr), W. Rodrigues 8573 

(INPA 21522) (INPA); Reserva Florestal Ducke, 16 Jul 1976 

(st), Adair s.n. (INPA 72795) (INPA); Manaus, estrada da 

Raiz, between Cachoerinha and Mind6, 30 Jul 1929 (fr), 

Ducke s.n. (RB 22845) (RB) [mixed collection]; Manaus, 

Campus Universitario do INPA, estrada do Japiim, 7 Dec 

1977 (st), Falcao & Coelho 209 (INPA); Manaus, Campus 

Universitario do INPA, estrada do Aleixo, 21 Feb 1977 (fl), 

Leite 2 (INPA); Manaus, margem do igarape do BuiAo, 10 

Jan 1955 (fl), J. Chagas 486 (MG25131) (IAN, INPA, MG); 

Dto. Agropecuario da SUFRAMA, rd. 174, km 64, Fazenda 

Esteio, Reserva N. 1109, tree N. 302, 02?25'S, 59049'W, 20 

Aug 1981 (fr), Nascimento et al. s.n. (INPA/WWF 1109.302) 

(NY); Manaus, Ponta Negra, Hotel Tropical, tree N. 17, 19 

Jul 1977 (fr), Rodriguez 9954 (INPA), 9976 (INPA); Rio 

Negro, Paduary, Mata do Tucano, 14 Oct 1947 (fr), Fr6es 

s.n. (RB 22596) (NY[2], SP, U, US). 

Local names and uses. Amarelinha 

(Chagas 486), 

pau amarelo (Adair s.n., INPA 72795). Label data


IN 

A~ 

Pe 

~~~~~~~~~~~'5* 

r~~~~~~~~~~~~~ ?. 

?~~~~~~~~ 

..: ~-; 

i "i' '"' ~... 

-.~~~~~~ o 

. 

i :'~~~~~~' 

~~~~~~~~~~~~~~.~:~ 

.~~~r 

?? 

!~~~~~. 

..... 

'.?'? ? B, 

FIG 7. hiarhi dckan (-Efrm uce 286,P,istye;F ro Nscmetoetal IPA! 09$0, Y) A 

pubescence inlniuiaros.FMauecpl. 

~ ~ ~ 

FIG. 79. Chimarrhis duckeana (A-E from Ducke 22846, P, isotype; F from Nascimento et aL INPA/1 109.302, NY). A. 

Habit of inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style (note antrorse 

pubescence in longitudinal rows). F. Mature capsule. 

o 

A 

i .o


from Chagas 486 reports that the wood of this spe- petioles, deltoid, glabrous outside and inside, densely 

cies is used in sawmills. 

covered with colleters throughout inside, 6-9 x 7-9 

mm, dark green, subcaducous, leaving a scar encir- 

Chimarrhis duckeana is distinguishable from C. cling the stem 1-2 mm wide. Leaves 17-23 x 11.5barbata 

by its larger ovate to cordate leaves with 22 cm, L/W = 1:1 to 1.5:1, widely pyriform, rounded 

shorter petioles; blades that turn black-brown when to subcordate at base, rounded to obtuse at apex, acudry 

(vs. olive-green to light-brown in C. barbata); men absent; dark green above and below, semi-coriashorter 

and stouter minutely costate style, puberulent ceous; drying dark olive-green above, reddish brown 

only on the costae (vs. slender terete [non costate], below, semi-leathery to leathery; glabrous above and 

and appressed-retrorse throughout in C. barbata); and below; primary and secondary veins glabrous, promishorter 

style branches. Nascimento et al. s.n. (INPA/ nent below, secondary veins 5-6 each side; tertiary 

WWF 1109.302) was reported to have been collected veins subparallel throughout; petioles (4-)7-11 mm 

from a tree 25 m tall and 55 cm dbh. The morpho- long, 3-4 mm thick, terete to adaxially flattened or 

logical variation in size and shape of the leaves and concave, glabrous; domatia absent. Inflorescences 2- 

architecture of the inflorescences of this species (as 4 per node, corymbose, densely branched distally, 

already reported in C. turbinata) makes it difficult to with opposite to subopposite decussate branches teridentify 

sterile specimens. 

minating in cymules, forming a divided flowering 

Fr6es 22596 has atypically long-pedunculate in- plane, (6.5-)9-15 

florescences and ovate leaves, but these two characters 

alone are, in my opinion, not sufficient for the 

establishment of a new taxon; future collections with 

flowers in anthesis may prove that this collection deserves 

at least varietal recognition. 

Ducke s.n. (RB 22845) sheet A (RB) is a mixed 

collection of C. barbata and C. duckeana. The specimen 

mounted on the upper portion of the sheet is C. 

duckeana, with blackish brown shiny leaves, stout 

inflorescences with mature capsules, and three calycophylls. 

The specimen in the lower portion of the sheet 

is C. barbata, with light-brown opaque leaves, smaller 

slenderer inflorescences in anthesis, and no calycophylls 

present. The presence of calycophylls is not 

uncommon in this species, which were also observed 

in Chagas 486 (IAN, INPA, MG). 

Adolpho Ducke (1876-1959) is the botanist that 

most contributed toward the understanding of the systematics, 

taxonomy, and phenology of the Amazonian 

species of Chimarrhis (see Delprete, 1996b: 37). It is 

a great honor to dedicate the epithet of this species to 

such a famous Amazonian botanist. 

13. Chimarrhis brevipes Steyermark, Mem. New 

York Bot. Gard. 12: 179. 1965. Type. Venezuela. 

Amazonas: Cerro Sipapo (Paraque), forest near 

Base Camp, 125 m, 25 Dec 1948 (fr), Maguire & 

Politi 27971 (holotype, NY; isotypes, U, US, 

VEN). Figs. 10A, 581, 77A, 80A,B 

Small trees. Leafy branchlets glabrous; older 

branches smooth, not exfoliating in narrow strips, 

grayish pale brown; lenticels not found. Stipules connate 

at base, not contorted in bud, often adnate to 

x 4.5-7.5 cm, lateral branches one 

pair, basal portion of axis not branched (3.5-)6-9.5 

cm long; L/A = 2.5 to 3:1, rachis basally subterete, 

distally decussately compressed, rachis and branches 

sparsely puberulent; distal bracts extremely reduced, 

deltoid to lanceolate, to 2.5 x 1 mm; bracteoles subtending 

flowers absent or reduced to microscopic 

scales. Flowers not seen, sessile to short-pedicellate, 

puberulent (observed from young fruits). Pollen not 

seen. Calyx extremely reduced, truncate to minutely 

lobed, 0.5-1 mm long, minutely puberulent; lobes 

barely distinguishable, to 0.5 mm long; rarely one 

calyx lobe expanded into a calycophyll, usually 

present on aborted flowers. Calycophylls ca. 5.5 x 

ca. 3.5 cm, blades obovate, with peltate venation, 

whitish; stalks ca. 1.8 cm long. Capsules obovoid, 

acute at base, rounded at apex, 4.5-6 x 2.5-3.5 mm, 

pale brown, without lenticels, minutely white puberulent 

below the disk; disk white-pubescent, not exceeding 

the calyx; disk loculicidal dehiscence present in 

old capsules. Seeds 0.85-1.05 x 0.63-1 mm. 

Distribution (Fig. 77A) and ecology. This species 

is known only from a few collections from lowelevation 

forests, on lateritic soil, 125-190 m, in the 

Venezuelan Amazon. Fruiting specimens were collected 

in February, April, and December. 

Specimen examined. VENEZUELA. TERRITORO FED- 

ERAL AMAZONAS: Dpto. Atabapo, plains of Rio Padamo, 

03?20'N, 65?14'W, 190 m, Feb 1990 (fr), Marin 740 (MO, 

NY); San Carlos de Rio Negro, 30 Apr 1982 (fr), Dezzeo 73 

(VEN). 

Dubious specimen. BRAZIL. RONDONIA: 

Projecto RADAM, Ponto 26-SC-20-XD, 30 Aug 1975 

(fr), Cordeiro 676 (VEN). This specimen has leath-


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FIG. FIG. 80. Chimarrhris 

Chimarrhis brevipes brevipes (Maguire (Maguire &r & Politi 27971, 27971, US, isotype). A. Habit of infiorescences 

inflorescences and mature 

leaf. 

B. Mature capsule. 

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ery, pandurate leaves, and subsessile fruits, as in C. brevipes, 

but the fruits of this specimen are abnormally large 

and the collection locality is outside the expected range. 

brown; lenticels not found. Stipules connate at base, 

not contorted in bud, often adnate to petioles, deltoid, 

not acuminate, glabrous outside, microscopically 

(40x) ciliolate, glabrous with a basal triangular area 

Chimarrhis brevipes resembles C. gentryana in of colleters inside, 8.5-9 x 6.5-10 mm, reddish green, 

having leathery, pandurate leaves (usually much larger 

in C. brevipes), but the latter differs from the previous 

in having much smaller and narrowly obconical 

subcaducous, leaving a scar encircling the stem and 

connected to the petiole scars, 1-2 mm wide. Leaves 

(8-)13-24 

capsules. 

It is one of the three species of Chimarrhis (with 

C. microcarpa and C. speciosa) present in Venezuela 

(Chimarrhis venezuelensis is a Dioicodendron dioicum, 

q.v.). One isotype specimen (Maguire & Politi 

27971, NY) has an evident calycophyll on an aborted 

flower, not mentioned by Steyermark (1965, 1974). 

This calyx-lobe expansion is a calycophyll proper 

because it has peltate venation. In the other collections 

(Marin 740, MO; Dezzeo 73, VEN), no calycophylls 

are present; therefore, this feature is sporadic 

and of no taxonomic significance in C. brevipes. The 

occurrence ofcalycophylls in Chimarrhis is very rare, 

being sporadically present in this species and C. barbata, 

and commonly present only in C. gentryana. 

Chimarrhis brevipes is similar to C. turbinata in 

having puberulent capsules with small pubescent disks 

not exceeding the calices, the former differing from 

the latter in having leathery broadly pandurate leaves, 

and smaller obovoid capsules. From C. barbata this 

species differs in having smaller obovoid capsules 

with pubescent disk not exceeding the calices (vs. turbinate 

capsules with puberulent disk exceeding the 

calyx), and pandurate leaves. 

Chimarrhis brevipes differs from C. microcarpa 

and C. speciosa by: capsular disks white-pubescent 

(vs. glabrous in the latter two), domatia absent (vs. 

tufted domatia), bigger, obovoid capsules (vs. subglobose), 

and leaves usually pandurate (vs. elliptic to 

obovate). 

14. Chimarrhis gentryana Delprete, Brittonia 48: 

37. 196. Type. Peru. Loreto: Maynas, Mishana, 

Rio Nanay, halfway between Iquitos and Santa 

Maria de Nanay, ca. 150 m, 73?30'W, 03?50'S, 

upland forest mostly on white sand, 1 Mar 1980 

(fl), Gentry & Aronson 25307 (holotype, MO; 

isotype, NY). Figs. 10E, 58J, 77B, 81A-F 

Trees usually 15-20 m tall, exceptionally to 30 m 

tall; bark deeply fissured, fibrous; wood extremely 

hard. Leafy branchlets 4-8 mm thick, semi-succulent, 

terete to subterete, glabrous; older branches 

smooth (not exfoliating in narrow strips), grayish pale 

x (4-)8-13 cm, L/W = 1.5:1 to 1.7:1, 

pandurate, obtuse to rounded at base, concave at basal 

area, obtuse at apex, acumen absent; dark green above, 

pale green below, semi-coriaceous; drying reddish 

brown, semi-leathery to leathery; glabrous above and 

below; primary and secondary veins glabrous, prominent 

below, secondary veins 7-10 each side; tertiary 

veins subparallel throughout; petioles 6-12 mm long, 

2-3 mm thick, terete to adaxially flattened or narrowly 

concave, glabrous; domatia absent. Inflorescences 2 

per node, corymbose, densely branched distally, with 

opposite to subopposite decussate branches terminating 

in cymules, forming a continuous flowering plane, 

9.5-15 x 7.5-11 cm, lateral branches 1-2 pairs, basal 

portion of axis not branched 4.5-7.5 cm long; L/A = 

1.5 to 2:1; rachis basally subterete, distally decussately 

compressed, rachis and branches white or golden 

minute-puberulent; distal bracts absent or extremely 

reduced, deltoid to linear, to 4 x 1 mm; bracteoles subtending 

flowers absent or reduced to microscopic 

scales. Flowers sessile to short-pedicellate; pedicels 

to 1.5 mm long, puberulent; hypanthium narrowly 

obconical, 1-1.5 x ca. 1 mm, puberulent. Calyx extremely 

reduced, truncate to short-lobed, 0.4-0.8 x 

1.3-2 mm, golden puberulent, pale green; lobes barely 

distinguishable, to 0.5 mm long; in some flowers one 

calyx lobe expanded into a calycophyll. Calycophylls 

5-7 x 1.6-3 cm, blades ovate to elliptic, with peltate 

venation, pale green to white; stalks 1.5-2 cm long. 

Corolla funnelform, deeply lobed, the lobes reflexed 

at anthesis, 3.5-4.5 mm long, white to greenish white; 

tube short cylindrical, 1.5-2.5 x 0.5-0.7 mm, glabrous 

outside and inside; lobes 5, 2-2.5 x 0.6-0.8 mm, narrowly 

oblong with acute apex, glabrous outside, with 

a basal area white-pilose and with distal zone glabrous 

inside. Stamens 5, exserted well above the corolla, 

equal, attached 1-2 mm from the base of the tube; filaments 

3.5-5 mm long, terete, basally flattened, connate 

to throat, sparsely white pilose at base; anthers 

elliptic, 0.6-0.7 x 0.3-0.5 mm, medially dorsifixed, 

base rounded, apex acute to short mucronate. Pollen 

not seen. Style exserted, 5-6 mm long, terete, densely 

strigose, appressed-retrorse; style branches extruding 

as two lips above corolla before anthesis, oblate, 0.3- 

0.6 mm long, stigmatic surface microscopically (40x) 

papillose; ovary-galls present. Capsules narrowly 

obconical to turbinate, acute at base, rounded at apex,


A 

FIG. 81. Chimarrhis gentryana (A-E from Huashikat 2136, MO; F from Vdsquez & Arevalo 8969, TEX). A. Habit of 

inflorescence with mature leaf. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Style (note antrorse 

pubescence). E. Stamen. F. Mature capsule. (Reproduced with permission from Brittonia 48, fig. 1. 1996.) 

' o i' 

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FIG81Charhsgnraa(-frmHahkt23,M;Ffo sqe&Aral899TX)A.Hbto 

pu~scn?) E Samn.F Mtue 8,f.1.19. 

apul. Rerduedwih erisio ro Bitona


3.5-5 x 1.5-3.5 mm, rust-brown, without lenticels, 

densely golden puberulent below the disk; disk white- 

pubescent, not exceeding the calyx; disk loculicidal 

dehiscence present in old capsules. Seeds 0.95-1.13 

x 0.58-0.88 mm. 

Distribution (Fig. 77B) and ecology. Primary and 

secondary high-canopy non-flooded rain forests (terra 

firme) with high canopy, in white sand and clay soil, 

of low elevations, 115-390 m, of Amazonian Ecua- 

dor, Peru, and Brazil. Flowering specimens were col- 

lected in February and March. Fruiting specimens 

were collected in March, April, July, and August. 

Specimens examined. ECUADOR PASTAZA: UNOCAL 

petroleum exploration well site, Mazaramu, 01?27'S, 

76?54'W, 390 m, 28 Apr 1990 (fr), Beck et al. 1055 (paratype, 

NY, QCNE); Kapawi, Rio Pastaza, 76?48'W, 02?31'S, 

235 m, 14-20 Jul 1988 (fr), Lewis et al. 13543 (QCNE). 

PERU. AMAZONAS: Valley of Rio Sagrado, 65 km N of 

Pinglo, Quebrada Caterpiza, 200 m, 19 Feb 1980 (fl), 

Huashikat 2136 (MO). LORETO: Prov. Maynas, Mishana, Rio 

Nanay, between Iquitos and Santa Maria de Nanay, 73?30'W, 

03?50'S, 150 m, 25 Jul 1984 (fr), Vdsquez et al. 5405 (MO, 

NY); Prov. Loreto, Nauta, road Nauta-Iquitos, 73?35'W, 

04?29'S, 200 m, 28 Mar 1987 (fr), Vdsquez & Arevalo 8969 

(F, TEX). 

BRAZIL. AMAZONAS: Mun. Sio Paulo de Olivenca, 

basin of Bel6m creek, Oct-Dec 1936 (fr), Krukoff8782 (MO, 

NY, U). 

This species is easily recognizable by its numerous 

cream-white calycophylls (Fig. 81A), rust-brown 

puberulent capsules, and puberulent disk which does 

not exceed the calyx. The calycophylls usually darken 

and break off at capsule maturity. The leaf blades of 

C. gentryana are commonly pandurate, semi-leathery, 

and olive-green when fresh, turning reddish brown 

when dry. 

Chimarrhis gentryana is most similar to C. brevipes 

(of the Venezuelan Amazon), both possessing 

pandurate leaves and calycophylls; the former differs 

from the latter in having narrowly obconical capsules 

(vs. broadly obovoid), which are rust-brown (vs. 

beige-brown) and densely golden-puberulent (vs. 

sparsely white-puberulent). 

Chimarrhis gentryana is often confused with Calycophyllum 

obovatum (Ducke) Ducke, because of its 

white calycophylls, minute white flowers, and obovate 

to pandurate leaves (Fig. 81A-E). The genus 

Calycophyllum differs from Chimarrhis in having 

terminal inflorescences (vs. axillary and subterminal), 

seeds with strongly bipolar wings (vs. minute seeds 

with small concentric wing), and smooth thin bark 

exfoliating into long vertical strips (vs. soft-fibrose 

bark falling off in small irregular pieces). 

Chimarrhis gentryana is commonly encountered 

as medium-sized trees (15-20 m tall; but two emergent 

individuals were reported to be 25 and 30 m tall), 

growing below the high canopy Amazon forest (30- 

35 m high), with small buttresses, rust-brown fibrous 

bark, leathery leaves, cream-white fragrant flowers, 

and cream-white calycophylls. 

Excluded and Dubious Species 

Chimarrhis bathysoides Steyermark, Mem. New York 

Bot. Gard. 12(3): 181. 1965. Type. Venezuela. 

Amazonas: Cerro Sipapo (Paraque), forest along 

trail near Base Camp, 25 Jan 1949 (fl), Maguire 

& Politi 28626 (holotype, NY; isotypes, B, GH, 

US[2], VEN[2]) = Bathysa bathysoides (Steyermark) 

Delprete (see Delprete, 1996b). 

Chimarrhis clausicorollata J. H. Kirkbride, Bio- 

Llania, Edicion Esp. 6: 394. 1997. Type. Guyana. 

Distr. Brokopondo, Kentoegoni Island, S of 

Gansee, 21 Jul 1966 (fl), van Donselaar 1471 

(holotype, U; photo-U at NY). = Simira tinctoria 

Aublet. 

This taxon has been recently described by Kirk- 

bride (1997a), based on a single specimen with young 

flowers. The terminal inflorescence and the reddish 

tinge in the leaves and the wood of this specimen prove 

that this species belongs to the genus Simira (Chimarrhis 

has lateral inflorescences and yellowish wood). 

Additional collections from virtually the same site 

(van Donselaar 1033-U, 1297-U, and 1324-U) have 

convinced me that the C. clausicorollata is synonymous 

with Simira tinctoria Aubl. More specifically, 

the specimen van Donselaar 1297 (U) has flowers that 

match Kirkbride's description of C. clausicorollata. 

Also, label data of all the van Donselaar's collections 

report that the vernacular name for this species is 

"tjintji-oedoe" (Sranan), which means tinged wood 

(Lindeman, pers. comm.). 

Chimarrhis decurrens Steyermark, Ceiba 3: 18. 1952. 

Type. Panama. Cocle: El Valle de Anton, 600 m, 

28 Jun 1946 (fl), Allen 3566 (holotype, F; isotypes, 

F[2], MO) = Allenanthus erythrocarpus Standley. 

Chimarrhis dioica K. Schumann & K. Krause, Bot. 

Jahrb. Syst. 40: 312. 1908. Type. Ecuador. Loja: 

Rio El Vargro, shrubs growing on oriental Andean 

slopes, 3400-3600 m, Oct-Nov 1906 (fl), Lehmann


7718 (B*, photo-B at G, GH, MO; lectotype, K) = 

Dioicodendron dioicum (K. Schumann & K. 

Krause) Steyermark. 

Chimarrhisferruginea (Standley) Standley, N. Amer. 

Fl. 32: 6. 1918. Rustiaferruginea Standley, Contr. 

U.S. Natl. Herb. 18: 142. 1916. Type. Panama. 

Col6n: Along Rio Fato, in forest or thickets, 10- 

100 m, Aug 1911 (fl), Pittier 4201 (holotype, US). 

(not Chimarrhis, not Rustia) = Bathysa (?) sp. 

Dwyer (1980) did not cite this species in either Chi- 

marrhis or Rustia. This species, in my opinion, does 

not belong to either of the two genera under consid- 

eration, and the absence of fruiting material (and the 

depauperate inflorescence) does not allow its place- 

ment with certainty. Future flowering or fruiting col- 

lections will hopefully clarify its generic position. 

Chimarrhis goudotii Baillon, Adansonia 12: 307. 

1879. Sickingia goudotii (Baillon) Standley, Publ. 

Field Columbian Mus., Bot. Ser. 7: 27. 1930. Type. 

Colombia. Quinindi6, 1844 (fl), Goudot s.n. (lec- 

totype, P, here selected; B*, photo-B at F, GH, 

NY) = Simira goudotii (Baillon) Steyermark. 

Baillon (1879) described a number of species of 

Simira under Chimarrhis, which were treated by him 

as closely related genera. Chimarrhis is easily recognizable 

by its axillary inflorescences and peltate seeds 

with concentric wing, while Simira has terminal inflorescences 

and horizontal seeds with a lateral orbicular 

wing. He treated Sickingia [= Simira] as a section 

of Chimarrhis. Chimarrhis goudotii was described by 

Baillon (1879) as "C. (Sickingia) goudotii" using 

specimens collected by Goudot in Colombia. One 

specimen preserved at Paris (P) has two labels: 

"Sickingia - Chimarrhis goudotii Baill [Baillon handwriting]. 

- Quindiu [Goudot handwriting]- NC"e Grenade 

- M. Justin Goudot 1844," and the other (in 

Baillon handwriting) bears a description of the species, 

which declares it related to Simira longifolia, and 

a sketch of the flower. I select the specimen preserved 

at P as the lectotype of C. goudotii (= Simira goudotii). 

An additional specimen of Goudot's collection was 

preserved at Berlin (now destroyed, photos at F, GH, 

NY) was labeled as "Chimarrhis (Sickingia) goudotii 

Baill. - Quindiu et Tolima - leg Goudot" (Baillon's 

handwriting). This species was correctly transferred 

by Steyermark (1972) to Simira. 

Chimarrhis 

longifolia (Willdenow) Baillon, Adanso- 

nia 12: 306. 1879. Sickingia longifolia Willdenow, 

Gesellsch. Naturf. Freunde Schr. 3: 445. 1801. 

Type. Venezuela. "Prov. Caracas," Bredemeyer 

s.n. (no types found) = Simira longifolia 

(Willdenow) Bremekamp. 

Baillon (1879) transferred Sickingia longiflora to 

Chimarrhis. Bremekamp (1954a) concluded that 

Sickingia and Simira are congeneric and transferred 

C. longifolia to Simira. Inspired by Bremekamp's 

conclusions, Steyermark (1972) transferred all of the 

species of Sickingia known to him to Simira. 

Chimarrhis paraensis Baillon, Adansonia 12: 308. 

1879. Sickingiaparaensis (Baillon) K. Schumann 

in Martius, Fl. Bras. 6(6): 232. 1889. Type. Bra- 

zil. Para: Saint-Hilaire s.n. (ex herb. Lisboa), ([P 

not found]; B*, photo-B at F, GH, NY; fragment 

of isolectotype, F [ex herbario parisiensis]) = 

Simiraparaensis (Baillon) Steyermark. 

The specimen used by Baillon in P to describe C. 

paraensis was not found, and the specimen once pre- 

served at B is now destroyed. The photos of the Ber- 

lin type allow enough definition to conclude that also 

this specimen was seen by Baillon (his handwriting 

is on the label). A fragment (one leaf) with the stamp 

"Ex Herbario Musei parisiensis" and preserved at F, 

is labelled "Sickingia paraensis - Para - herb. 

lusitanicum [Lisboa, Portugal]"; this is probably a 

portion of the Paris type. 

Chimarrhisperijaensis Steyermark, Acta Bot. Venez. 

8: 248. 1973. Type. Venezuela. Zulia: Sierra Perija, 

forest near Rio Omira-Kuna, SW de Pishikako, 

1440-1460 m, 28 Mar 1972 (fl), Steyermark & 

Dunsterville 105658 (holotype, VEN) = Bathysa 

perijaensis (Steyermark) Delprete (see Delprete, 

1996b). 

Chimarrhis pisoniaeformis Baillon, Adansonia 12: 

307. 1879. Sickingia pisoniiformis (Baillon) K. 

Schumann in Martius, Fl. Bras. 6(6): 225. 1889. 

Type. Brazil. Para(?): Saint-Hilaire s.n. [ex herb. 

Lisboa] (lectotype, P, here selected; photo-P at A, 

F, MO, NY). Ixora podocarpa J. D. Hooker in 

Bentham & Hooker, Gen. PI. 2: 113. 1873. Type. 

Brazil. Rio Negro, above Barraroa, Spruce 1994 

(lectotype, K, here selected; photo-K at NY). = 

Simira pisoniiformis (Baillon) Steyermark. 

The selected lectotype of C. pisoniaeformis pre- 

served at P has not been directly seen, but its photo 

(at A, F, MO) is adequate to identify it. This speci-


men has two labels (Baillon handwriting): one with a 

sketch of a flower, and the other with the listing of its 

morphological characters. Steyermark (1974) noted 

that Ixora podocarpa (as "nomen subnudum") is synonymous 

with Sickingia pisoniiformis. I did not see 

the type specimen of Ixora podocarpa preserved at 

K, but its photo (NY) is adequate to recognize the 

characteristic shape of its capsules, and its label 

(Spruce handwriting) reports the collection locality as 

given in the original description; consequently the K 

specimen is selected as lectotype of I. podocarpa. 

Chimarrhis pittieri Standley, Publ. Field Columbian 

Mus., Bot. Ser. 8: 53. 1930. Type. Venezuela. 

Distrito Federal: Caracas, 1926 (fr), Pittier 11954 

(holotype, US, US-frag at F, US-photo at F, G; 

isotypes, G, NY, VEN) = Bathysa pittieri 

(Standley) Steyermark. (see Delprete, 1996b). 

Chimarrhis sessile Rusby, in herb., ined. Basis: Bo- 

livia, Yungas, 1890, Bang 418 (G, GH, NY). 

Bang 418 was annotated and distributed by Rusby 

as "C. sessile," a name that remained unpublished. 

Bang 418 is the type of Malanea grandis Rusby 

(1893), which was later transferred by Rusby (1895) 

to Elaeagia. About its identity, Steyermark (1965) 

stated "Standley [1931b] placed this species under 

Elaeagia in his Rubiaceae of Bolivia, but it is obvi- 

ously not this genus. The 5-6-lobed corolla, 5-6 sta- 

mens, conspicuously dentate 5-6-lobed calyx, and 

other characters would exclude it from Elaeagia, and 

the multi-ovulate cells of the ovary would exclude it 

from Malanea, with which it has also been filed in 

some herbaria." This species also does not belong to 

Chimarrhis because of its terminal inflorescences. To 

place this taxon in a genus, fruiting specimens are 

necessary. At the present time I can only suggest 

placement in either Bathysa or Warszewiczia. 

Chimarrhis venezuelensis Standley & Steyermark, 

Fieldiana, Bot. 28: 162. 1930. Type. Venezuela. 

Lara: Slopes of mtn. below Santo Domingo and 

Los Quebraditos, S of Las Sabanetas, above 

Humocaro Bajo, 2430-2475 m, 8 Feb 1944 (fl), 

Steyermark 55408 (holotype, F; isotypes, GH, NY, 

US, VEN). = Dioicodendron dioicum (K. 

Schumann & K. Krause) Steyermark (see Dioicodendron, 

below). 

DIOICODENDRON 

7. Dioicodendron Steyermark, Bol. Soc. Venez. 

Cienc. Nat. 25: 24. 1963. Schumann & Krause, 

Engl. Bot. Jahrb. 40: 312. 1908; Standley, Publ. 

Field Columbian Mus., Bot. Ser. 7: 203. 1931; 

Standley & Steyermark, Fieldiana, Bot. 28: 570- 

571, fig. 121. 1953; Steyermark, Bol. Soc. Venez. 

Cienc. Nat. 23-28. 1963; Steyermark in Lasser & 

Steyermark, Fl. Venez. 9(1): 253-256, fig. 40. 

1974; Delprete in Harling & Andersson, Fl. Ecuador 

(in press). Type species. Dioicodendron 

dioicum (K. Schumann & K. Krause) Steyermark. 

Shrubs to trees, much-branched, without buttresses; 

bark light brown. Stipules interpetiolar, free 

at base, not adnate to the petioles, persistent. Leaves 

short-petiolate, elliptic to ovate, short-acuminate at 

apex; petioles not thickened at base; domatia absent. 

Inflorescences terminal, frondose, pyramidal, laxly 

paniculate, with opposite to subopposite lateral 

branches terminating in corymbose cymules. Flowers 

dioecious, 4-merous, short-pedicellate, fragrant; 

hypanthium of male flowers shallowly obconical; hypanthium 

of female flowers oblong-elliptic. Calyx 

extremely reduced, with 4 narrowly triangular to ovate 

lobes, persistent. Corolla funnelform, deeply lobed, 

with reflexed lobes, white to greenish white; tube 

extremely reduced, entirely glabrous; lobes 4; aestivation 

valvate with contact zone. Male flowers whitish 

green, fragrant, with 4(-5) stamens alternate to 

the petals, exserted well above the corolla; filaments 

attached on the upper part of the tube, slender, basally 

flattened, entirely glabrous; anthers oblong-elliptic, 

dorsifixed, dehiscing by lateral slits. Pollen tricolporate, 

exine thinly reticulate. Female flowers 

whitish green, fragrant, with exserted style; style 

branches protruding above the corolla only after anthesis, 

arcuate, with the tips remaining in contact during 

maturity and pollination, ovate. Ovary 2-celled, 

turbinate to obovate, placentation peltate on the septum; 

ovules many in each locule, horizontally inserted. 

Capsules woody, obovoid to turbinate; dehiscing 

septicidally, disk-loculicidal dehiscence present in old 

capsules. Seeds obovoid to suborbicular, 3-5-angular, 

with a microscopic concentric wing. 

Dioicodendron 

is a unique genus of trees in the Rubi- 

aceae, having dioecious tetramerous sweet-fragrant 

flowers, whitish green to bluish green corollas, foliose- 

ovate spreading stipules, cinnamoneous (light-brown) 

bark, and capsules containing minute ovoid seeds.


Dioicodendron is similar to (and often confused Semiscandent shrubs 2-4 m tall to much 

with) Chimarrhis in having minute fragrant flowers 

and peltate placentation; the previous differing from 

the latter in having terminal, paniculate, frondose inflorescences 

(Fig. 82A; vs. lateral, subterminal), flowers 

4-merous and dioecious (vs. 5-merous and monoecious), 

ovate-foliose persistent stipules (Fig. 82B,C; 

vs. narrowly lanceolate and readily caducous), and 

minute, ovoid, non-winged seeds (vs. minute, flattened, 

and with concentric wing). Dioicodendron has 

also been confused with Chimarrhis, because the latter 

has been erroneously reported to be dioecious or 

heterostylous (Wright 1262 in label data; Grisebach, 

1862; Urban, 1899). 

The flowers of Dioicodendron (Fig. 83A-F) have 

valvate-reduplicate aestivation. Its female flowers 

(Fig. 83A-D) are short-pedicellate, with spreading 

corolla lobes, narrowly obovoid hypanthium, and the 

stigmas convergent at maturity. Its male flowers (Fig. 

83E,F) are, in contrast, long-pedicellate, with erect 

corolla lobes, shallowly obconical hypanthia, filaments 

curved toward the center, and anthers often in 

contact with each other. 

Dioicodendron is here treated as a monotypic genus, 

occurring at medium-high elevations of the South 

American Andes. 

1. Dioicodendron dioicum (K. Schumann & K. Krause) 

Steyermark, Bol. Soc. Venez. Ci. Nat. 25: 24. 1963. 

Chimarrhis dioica K. Schumann & K. Krause, Bot. 

Jahrb. Syst. 40: 312. 1908. Type. Ecuador. Loja: 

Rio El Vagro, shrubs growing in E Andean slopes, 

3400-3600 m, Oct-Nov 1906 (fl), Lehmann 7718 

(lectotype, K, selected by Delprete, 1999b; photo-K 

at VEN; isolectotype, B*, photo-B at G, GH, MO, 

VEN). 

Figs. 3R, 4L,M, 11A, 18G,H, 82A-E, 83A-F, 84 

Dioicodendron cuatrecasasii Steyermark, Bol. Soc. 

Venez. Ci. Nat. 25: 25. 1963. Type. Colombia. Valle: 

Cordillera Oriental, W slopes, riverbank of Rio 

Digua, left side, Piedra de Moler, 900-1180 m, 19- 

28 Aug 1943 (fr), Cuatrecasas 15192 (holotype, 

VEN; isotypes, F, US). 

Chimarrhis venezuelensis Standley & Steyermark, 

Fieldiana, Bot. 28: 570, fig. 121. 1953. Type. Venezuela. 

Lara: Slopes of mtn. between Santo Domingo 

and Los Quebraditos, S of Las Sabanetas, above 

Humocaro Bajo, 2430-2475 m, 8 Feb 1944 (male 

fl), Steyermark 55408 (holotype, F; isotypes, A, NY, 

US, VEN). 

branched trees 10-12 m tall, 10-20 cm dbh (rarely 

single-stemmed trees 20 m tall); trees with slender 

trunk, much branched at crown; bark smooth, thin, 

grayish brown (cinnamoneous). Leafy branchlets 

subtetragonal, 3-5 mm thick, pale brown, densely 

puberulent to pubescent (to rarely hirtellous); older 

branches glabrate, grayish to pale brown. Stipules free 

at base, foliose to subfoliose, widely to narrowly ovate 

(rarely deltoid), rounded at apex, sparsely to densely 

puberulent outside, sparsely pubescent with few basal 

narrow colleters inside, 3-8 x 4-7 mm, light green, 

persistent, splitting and breaking when old. Leaves 

(4-)7-16(-20) x (1.5-)3-8.5 cm, L/W= 1.6:1 to 2.5:1, 

lanceolate, oblanceolate to narrowly ovate, attenuate 

to rounded at base, acute at apex, gradually tapering 

into an acumen 0.5-2 cm long; dark green above, pale 

green below, foliaceous, drying brownish green 

above, yellowish green below; glabrous to sparsely 

puberulent above; glabrous, puberulent, or pubescent 

(rarely hirsutulous) below; the blade plane or often 

rugose (depressed secondary venation); primary, secondary, 

and tertiary veins sparsely to densely puberulent 

(to appressed-pubescent), secondary veins 5-9 

each side, ascending-arcuate, anastomosing 2-5 mm 

from the margin; tertiary veins subparallel-reticulate; 

petioles 3-20 mm long, 1-2(-3) mm thick, flattened 

to concave adaxially, glabrescent, puberulent to shortpubescent; 

Inflorescences terminal, frondose, laxly 

to densely paniculate, with opposite to subopposite 

lateral branches subtended by leaf-like bracts, terminating 

in dense corymbose cymules, pyramidal to 

umbelliform; 16-29 cm long, basal branches (10-)12- 

23 cm long, lateral branches 3-7 pairs; rachis 

subtetragonal, rachis and branches sparsely puberulent 

or erect to appressed-pubescent; flowers on distal 

branches in dense cymules; foliose bracts subtending 

the secondary, tertiary and quaternary branches 

leaf-like, decreasing in size toward the distal portion 

of the inflorescence, 8-13 x 3-7 cm at basal branches, 

gradually reduced to 1 x 0.5 cm at most distal 

branches; bracteoles subtending flowers 0.7-3 x 0.2- 

0.4 mm, linear, sparsely puberulent. Flowers shortpedicellate 

(female flowers) to pedicellate (male flowers). 

Calyx extremely reduced, 0.5-1 x 0.7-1 mm, 

pale green to bluish green; lobes 4, narrowly triangular 

to widely ovate, 0.7-1.2 mm long. Corolla funnelform, 

deeply lobed with reflexed lobes, 1.5-3.5 

mm long, entirely glabrous, white to greenish white; 

tube extremely reduced, 0.3-0.5 x 0.7-1.3 mm; lobes 

4, 1.5-2.3 x 1-1.4 mm, oblong-ovate and acute at 

apex. Male flowers with pedicels 1.5-2 mm long,


trt 

ALi 

N~, . . i .. 

FIG. 82. Dioicodendron dioicum. A. Habit of frondose inflorescence (Lozano & Diaz 3 734, COL). B, C. Stipules (Ham- 

mered et al 5233, NY). B. Lateral view. C. Frontal view. D, E. Mature capsule (Little 8505, US). D. Lateral view. E. Top 

view, with circular disk and central hole.


AI[ 

i ' & ,~~~~~~~~~~~~~~~~~~~~~~~~~ 

]~~~~~7 

FIG. 83. Dioicodendron dioicum. A-D. Female flower (Barclay et al. 3424, US). A. Flower in anthesis. B, C. Female 

flower bud. B. Side view. C. Top view. D. Close up of receptive stigma. E, F. Male flower (Cuatrecasas 8665, US). E. 

Male flower in anthesis. F. Close up of anther (note reticulate thecae).


00 '70 

'0 

cleared slopes, (450-)900-3500 m, of Colombia, 

Venezuela, Ecuador, and Peru. Flowering specimens 

were collected in January, February, March, April, 

May, July, August, October, and November, but 

mainly in October. Fruiting specimens were collected 

in February, April, May, July, August, October, No- 

vember, and December. 

Specimens examined. COLOMBIA. BOYACA: Cerro 

Berlin, between Arcabuco and La Palma, 1780-1800 m, 28 

Mar 1973 (male fl), Cuatrecasas et al. 28672 (COL, US[2]); 

Cordillera Oriental, Mun. Arcabuco, 2850 m, 20 Oct 1965 

(male fl), Huertas & Camargo 6282 (COL); Mun. Arcabuco, 

La Cumbre, 2400 m, 9 Oct 1980 (male fl), Lozano & Diaz 

3734 (COL); Cordillera Oriental, NE of Arcabuco, 2650 m, 

12 Oct 1966 (male fl), Garcia-Barriga 18763 (AMES, 

COL); Arcabuco, 2750 m, Nov 1965 (male fl), Uribe-Uribe 

5468 (COL, NY). CAQUETA: Cordillera Oriental, Ruidosa, 

5 km NW of camp Ruidosa, 2800 m, 11 Jul 1944 (fr), Little 

8205 (COL, NY[2], US, VEN). CAUCA: Cerro San Pedro, 

2700 m, 10 Sep 1944 (fr), Hutter 1215 (US). CHoc6: Mun. 

San Jose de Palmar, 11 Nov 1985 (fr), Lozano et al. 4885 

(COL); Mun. N6vita, Curund6, right bank of Rio Ingari, 

400-550 m, 12 Jan 1983 (male fl), Diaz-Pedrahita 3446 

(COL); Cieneguita, s.d., Triana 1869 (P, US). CUNDINA- 

FIG. 84. Distribution ofDioicodendron dioicum (circles) MARCA: Rd. Cundinamarca-Fusagasuga, km 38-39,2300 m, 

and Dolichodelphys chlorocrater (stars). 

17 May 1972 (female fl), Barclay et al. 3424 (COL, US); 

rd. Bogota-Fusagasuga, below San Miguel, Quebrada Malagana, 

28 km SW of Bogota, 2600 m, 30 Apr 1944 (female 

sparsely puberulent; hypanthium shallowly obconical, fl), Fosberg 21857 (NY, VEN). HUILA: Cordillera Oriental, 

0.5-1 x 0.7-1 mm, glabrous to sparsely puberulent; 

Vertiente Occidental, between Gabinete and Andalucia, 

stamens 4(-5) exserted well above the 2200-2300 

corolla, m, 25 Mar 1940 

equal, 

(male fl), Cuatrecasas 8665 

attached at the superior zone of the tube, alternate to (COL, US); Mun. La Plata, Vereda Arrabal, Turbera de 

Paramo, 2380 m, 1 Oct 1984 

the lobes; filaments 2.5-3 mm 

(female fl), Lozano et al. 4395 

long, terete, glabrous, 

(COL). META: Ridge between Rio Tigre and Rio Papamene, 

white; anthers oblong-elliptic, 0.6-1.2 x 0.3-0.5 mm, Colombia-Uribe trail, 03013'N, 74034'W, 1100 m, 23 Dec 

medially dorsifixed, reversed at maturity, microscopi- 1942 (fr), Fosberg 19527 (US). PUTUMAYO: Punto Buenos 

cally (40x) reticulate (Fig. 78F), base rounded; male Aires, Cerro Portachuelo, 2800 m, 25 Jul 1964 (male fl), 

flowers sometimes with a non-functional style (with- Soejarto 1086 (ECON, US). QuINDfo: Quindio, 24 Nov 1853 

out functional stigmas). Female flowers with pedicels (fl), Holton 410 (K, NY). VALLE: Mun. Argelia, vereda Las 

0.5-1 mm long, sparsely puberulent; hypanthium Brisas, 2050-2200 m, 21 Jan 1983 (male fl), Diaz-Pedrahita 

oblong-elliptic, 2-2.5 x 1.3-1.6 mm, glabrous to 3817 (COL); Cordillera Occidental, N of Las Brisas, 

sparsely puberulent; style exserted, (2.5-)3-3.5 mm 

Gibraltar, 2100-2200 m, 25 Oct 1946 (male fl), Cuatrecasas 

22516 

long, glabrous, terete; style branches arcuate, with the 

(US); Cordillera Occidental, mt. above Alto de Mira 

(between Tabor and Carrizales), 2100-2350 m, 23 Oct 1946 

tips connected at maturity, thick, widely ovate, 0.7-1 

(fr), Cuatrecasas 22458 ([2]US); Cordillera Occidental, 

mm long, stigmatic surface microscopically (40x) Serrania de Los Paraguas, Mun. El Cairo, Cerro del Ingl6s, 

papillose. Capsules obovoid to turbinate, acute at 2260 m, 5 Jan 1987 (male fl), Silverstone-Sopkin 3001 (US). 

base, apex rounded, 3-4.5 x 2-3.5 mm, dark brown WITHOUT LOCALITY: 1760-1808, Mutis 2194 (US). 

without lenticels, glabrous throughout; disk donut- VENEZUELA. LARA: Las Sabanetas, SW slopes of 

shaped, with a hole at the center (caused by the cadu- Aposentos, above Humocaro Bajo, 2500-2530 m, 3 Feb 

cous style); loculicidal dehiscence present in old 1944 

cap- 

(fr), Steyermark 55220 (US); SW-facing slopes at Los 

sules. Seeds obovoid to orbicular, 

Aposentos, above Las Sabanetas, 2500-2530 m, 3 Feb 1944 

3-5-angular, 

(fr), Steyermark 55223 (VEN). TRUJILLO: Dtto. Bocon6, 20 

0.5-0.9 mm long, testa reticulate. 

km E of Bocon6, Guaracamal, 09?14'N, 70011'W, 1900- 

2300 m, 7 Feb 1987 (female fl), Aymard et al. 5233 (NY, 

Distribution (Fig. 84) and ecology. Rare shrubs VEN). 

and trees of understory in seasonal dry forest and its ECUADOR. CARCHI: Cant6n Tulcan, above Maldoremnants, 

rarely in seasonal cloud forest, also on nado, Chilma, 78002'W, 005 1'N, 2000 m, 20 May 1991 (fr),


Palacios & Rubio 7290 (MO, QCNE, TEX); Espejo, El 

MOLOPANTHERA 

Guantal, slopes of Cerro Golondrinas, 00?51'N, 78?07'W, 

8. 

2450 m, 21 Aug 1994 (fr), Palacios & Clark 12633 

Molopanthera Turczaninow, Bull. Soc. Imp. 

(QCNE). 

LOJA: Parque Nacional Podocarpus, above Nudo de 

Naturalistes Moscou 21: 580. 1848. Forsteronia 

Cajanuma, 

trail to Mirador, 3000-3500 m, 79?10'W, 04?05'S, 10 Casaretto ex K. Schumann, in synon., in Martius, 

Apr 1989 (fr), Madsen 85961 (QCNE); rd. Loja-Malacotas, Fl. Bras. 6(6): 200. 1889; Turczaninow, Flora 31: 

Nudo de Cajanuma, 10 km S of Loja, 2670 m, 4 Oct 1944 711-712. 1848; Miquel, Linnaea 22: 804-805. 

(female fl), Wiggins 10880 (DS, NY, TEX, US, VEN); be- 1849; Walpers, Ann. bot. syst. 2: 799. 1852; J. D. 

tween Tambo Cachiyacu (La Entrada) and Nudo de Hooker in Bentham & Hooker, Gen. P. 2: 41-42. 

Sabanillas, 2500-3500 m, 7 Oct 1943 (female fl), Steyermark 1873; Baillon, Hist. P1. 7: 346-347, 491. 1880; 

54450 (NY, US, VEN). MORONA-SANTIAGO: Rd. Gualaceo- Schumann in Martius, Fl. Bras. 6(6): 198-200, fig 

El Lim6n, 2500 m, 15 Aug 1989 (fl), Van der Werff & 

109. 1889; Schumann in 

Gudinfo 11098 

Engler & Prantl, Nat. 

(QCNE). 

Pflanzenfam. 

PERU. AMAZONAS: Prov. Chachapoyas, 1-5 km W of 

4(4): 54.1891. Type species. Molop- 

Molinopampa, Jalca, 2400-2450 m, 18 Jul 1962 

anthera 

(fr), 

paniculata Turczaninow. 

Wurdack 1383 (K, NY, UC, US, VEN); Shanyan, Nov (fl), 

Jelski 376 (paratype of C. dioica [frag-W at F]). 

Trees; bark longitudinally fissured, brown; sapwood 

pale-yellow, very hard; heartwood light-brown. 

Local names and uses. Venezuela: quina blanca 

Stipules interpetiolar, free at base, adnate to petioles, 

(Steyermark 55220). The bitter bark is used as an alpersistent. 

Leaves elliptic to narrowly elliptic, acumiternative 

source of quinine to cure malaria (Standley nate at apex; not pellucid punctate; petioles slightly 

& Steyermark, 1953). 

thickened at base; domatia absent. Inflorescences 

terminal, frondose, pyramidal, laxly paniculate, sec- 

This species was first described by Schumann and 

ondary branches subtended by leaf-like bracts, tertiary 

Krause (1908a) as Chimarrhis dioica, based on speci- branchlets thyrsoid. Flowers zygomorphic (flower 

mens collected in Ecuador and Peru; of its two syn- buds and stamens curved upward), protandrous, 

types, I studied Lehmann 7718 preserved at Kew, sessile to short-pedicellate, fragrant; hypanthium obowhich 

was selected as lectotype by Delprete (1999b). void. Calyx extremely reduced, with minute deltoid 

A duplicate specimen was once preserved at Berlin 

lobes, persistent. Corolla rotate, curved upward, 

(probably the original type, but now destroyed), of 

deeply lobed, with reflexed unequal lobes, glabrous 

which I studied the photos (G, GH, MO). 

throughout, white to cream-white; tube short-cylin- 

Standley and Steyermark (1953) described C. 

drical; lobes (4-)5; aestivation imbricate. Stamens 5, 

venezuelensis, distinguishing it from C. dioica, the alternate to petals, unequal (the lower ones longer than 

previous having longer spreading hairs, larger cap- the upper ones), exserted; filaments attached near the 

sules, and shorter filaments. 

base of the tube, slender, basally flattened, glabrous; 

Steyermark (1963) founded the genus Dioico- anthers narrowly elliptic, dorsifixed near base, dehiscdendron, 

the name meaning dioecious tree, to which 

ing by lateral slits. Pollen tricolporate, exine densely 

he transferred C. dioica, and described D. cuatrereticulate. 

Style exserted, elongating after anther decasasii. 

Chimarrhis venezuelensis was reported by hiscence; style branches narrowly oblong, reflexed at 

Steyermark (1963) to be synonymous with D. diomaturity. 

Ovary 2-celled, obovoid, bilobed, with a 

icum. The characters (vestiture, size of capsules, and 

globose placenta supported by a transverse stalk atsize 

and shape of calyx lobes) used by Steyermark tached to the septum; ovules many per locule. Cap- 

(1963) to separate D. cuatrecasasii from D. dioicum 

sules thin-woody, strongly bilobed, the two sides subare 

solely a geographical variation of secondary morglobose; 

dehiscing loculicidally, disk septicidal 

phological features, and the two taxa are synonymous. dehiscence absent. Seeds with irregular outline, with 

The collections Holton 410 (GH, NY) annotated a concentric deeply fringed wing. 

by Standley as "Molopanthera colombiana Standl" 

(unpublished name), are D. dioicum. 

Molopanthera is a rare genus of trees endemic to 

Dioicodendron is a rare genus most commonly the forests of eastern Brazil, easily recognizable by 

found in the Colombian cordilleras at 450-3500 m. its minute curved flower buds (Figs. 4T, 85B), small 

Human pressure on these environments will likely bilobed-globose capsules, capitate placenta supported 

cause the species' demise. 

by a short stalk, seeds peltate to the placenta, umbilically 

attached, and concentric seed-wing with 

deeply fringed margins (Fig. 11B).


Genera with zygomorphic flowers are rare in the 

Rubiaceae. In Molopanthera, zygomorphy is more 

noticeable in its minute flower buds, which are curved 

upward (Fig. 85B), than in its open flowers (Fig. 85C). 

Nevertheless, when magnified, it is possible to appreciate 

the unequal length of the filaments and corolla 

lobes (longer in the lower portion of the corolla). The 

filaments remain bent upward in two couples with the 

two adjacent anthers connected to each other by their 

pointed extensions at both ends, forming two cupshaped 

units (Fig. 85C), the fifth filament remaining 

free. This particular morphology is correlated with its 

peculiar pollination syndrome. The anthers in couples 

release pollen in a spherical mass, which is thrown 

onto the pollinator (i.e., small bees) by the single anther 

(pers. obs.) 

Turczaninow (1848) in describing M. paniculata 

as having leaves glabrous throughout. The generic 

name is derived from the Greek gioXoWxV (molops = 

Trees 5-10 m tall, rarely to 30 m tall, to 80 cm 

dbh; bark shallowly longitudinally fissured. Leafy 

branchlets thin-woody, glabrous or densely ascending 

pubescent; lenticels sparse on older branches. 

Stipules free at base, broadly deltoid, long-acuminate, 

glabrous or densely pubescent outside, long-sericeous 

with few basal colleters at base inside, 3-4 x 4-5 mm, 

persistent, leaving a scar encircling the stem. Leaves 

5-16 x 2.5-5 cm, elliptic to narrowly elliptic, acute 

at base and at apex, sometimes short-acuminate, the 

acumen 0.5-1.2 cm long; stiff-foliaceous; glabrous or 

sparsely pubescent above, glabrous or densely golden 

pubescent below; primary and secondary veins glabrous 

or densely pubescent, prominent below, secondary 

veins 5-8 each side; tertiary veins subparallel-reticulate; 

petioles 0.7-1.5 mm long, 1-1.5 mm thick, 

adaxially concave, glabrous to densely pubescent; 

domatia absent. Inflorescences frondose, pyramidal, 

laxly paniculate, with opposite decussate branches, 

bruise or weal) and av9rTo (anteros = anther), mean- each lateral branch subtended by leaf-like bracts, tering 

bruised anthers, probably in allusion to the fact minating in tertiary thyrsoid branchlets, 13-18 cm 

that they are often fused (in couples) during anthesis. long; basal branches 9-13 cm long, lateral branches 

Molopanthera has been treated as incertae sedis 5-9 pairs; rachis basally terete, decussately com- 

(Turczaninow, 1848), tentatively placed in the Cinpressed 

distally, without lenticels, rachis and branches 

choneae (Hooker, 1873; Baillon, 1880; Schumann 

densely short-pubescent; flowers in tertiary thyrsoid 

1889, 1891), transferred to the Condamineeae (sensu branchlets single or in few-flowered units. Flowers 

Hooker) by Andersson and Pearsson (1991), and in- sessile to short-pedicellate; pedicels to 0.5 mm long, 

cluded in the "genera associated with Portlandia" by short-pubescent; hypanthium obovoid, faintly bilobed, 

Robbrecht (1993). Molopanthera is here treated as a 0.5-0.9 x 0.6-0.7 mm, densely pubescent; flower buds 

monotypic genus with the two varieties recognized by curved upward, zygomorphic. Calyx minutely 

Schumann (1889). 

cupular, short-lobed, 0.4-0.7 x 0.9-1.2 mm, pubescent; 

lobes deltoid, 0.3-0.5 mm long. Corolla rotate, 

deeply lobed, the lobes reflexed at anthesis, 3-4 mm 

long, glabrous throughout, white to cream-white; tube 

1. Molopanthera paniculata Turczaninow, Bull. Soc. 

short cylindrical, 0.3-0.5 x 0.5-0.6 mm; lobes (4-)5, 

Nat. Imperialistes Moscou 21: 581. 1848. 

2.5-3.5 x 0.4-1 

Molopmm, 

oblong-linear with round apex. 

anthera panaiculata K. Schumann, orth. var., in Stamens (4-)5, exserted well above the corolla, un- 

Martius, Fl. Bras. 6(6): 200, tab 109. 1889. equal, attached 0.1-0.2 mm from the base of the tube; 

Forsteronia panniculata Casaretto ex K. Schu- filaments 0.7-1.5 mm long, the lower ones longer than 

mann, in syn., in Martius, Fl. Bras. 6(6): 200. 1889. the upper ones (because of flowers buds curved up- 

Type. Brazil. Bahia: Serra de Jacobina, 1837, ward), slender-terete, basally flattened, glabrous 

Blanchet 2557 (lectotype, K, here selected; photo- throughout; anthers narrowly elliptic, 1-1.5 x 0.3-0.4 

K at NY; isolectotypes, F, G, MG, NY). Possible mm, dorsifixed near the base, with pointed dark exisolectotypes 

might be found at KW and LE. tensions at both ends, which are often fused in adja- 

Figs. 3T, 4N, 1 lB, 85A-D, 86 cent anthers. Pollen exine minutely foveolate-reticulate. 

Style exserted, elongating only after anthers 

Coffea floribunda Miquel, nom. superfl., Linnaea 22: 

dehiscence, 1-1.5 mm long, terete, glabrous; style 

804. 1849. Specimens cited: Brazil. Bahia: Serra de 

branches linear-oblong, reflexed, 0.5-1 mm long, stig- 

Jacobina, 1837, Blanchet 3282 (F, G-n.v., KW?, 

matic surface minutely papillose. Capsules short- 

LE?, MG, RB, U). 

Non Coffeafloribunda Martius, Flora 24, suppl. 2: 85. pedicellate, pedicels 0.5-1 mm long; strongly bilobed 

1841. Type. Brazil. Cuiaba: Morro do Rio da Casca, the two sides globose, rounded at base and at apex, 

collector unknown (BR?). [= Ixora densiflora 2-3.5 x 3.5-5 mm, golden to greenish yellow (to pale- 

(Martius) Mull. Arg.]. 

green at maturity), without lenticel, glabrous to


: ~ "- I ...- '- _. _ A . - 

ii 

FIG. 85. Molopanthera paniculata var. paniculata. (A-C from Williams & Assis 6861, GH; D from Orlandi 2237, F). 

A. Habit of inflorescences. B. Curved flower bud. C. Flower with exserted and connected anthers. D. Mature capsule.

Systematic Treatment of the Genera Studied 

sparsely short-pubescent below the disk; disk gla- 1972 (fl), Spada 31 (RB[2]); Reserva Forestal Linhares, near 

brous, not exceeding the calyx; disk septicidal dehis- rd. 143, talhAo 602, 29 May 1972 (fl), Lino 75 (RB). BAHIA: 

cence absent. Seeds 0.8-3 x 1-2.2 mm, with concen- 

Mun. Sta. Cruz de Cabralia, Pau-Brasil Ecological Station, 

ca. 16 km W of Porto 

tric 

Seguro, 27 Jul 1978 (fl), Mori et al. 

wing deeply fringed. 

10328 (K, NY); Bahia, Encruzilhada, rd. to Itamb6, Mata 

Cip6, 27 May 1968 (fl), Belem 3677 (CEPEC, NY); 

Turczaninow (1848) described Molopanthera Jacobina, 1 1?16'S, 40?27'W, 25 Aug 1980 (fr), Orlandi 223 

paniculata citing two syntypes, Blanchet 2557 and (CEPEC, F, HRB, MG, RB); Mun. Jacobina, Serra do Brite, 

Blanchet 3282. Curiously, one year later, Miquel 11009'S, 40?01'W, 500-600 m, 21 Aug 1984 (fl), Bautista 

(1849) described Coffeafloribunda based on Blanchet & Orlandi 985 (CEPEC, HRB, MG); Dto. Itaitu, Mun. 

3282, the syntype of M. paniculata, present at U, Jacobina, km 8 on rd. Jacobina-Capim Grosso, 27 Oct 1995 

which is here selected as the holotype of (fr), Jardim et al. 711 (CEPEC); Mun. Anderai, km 10 on 

C.floribunda. 

rd. 

Because of the difficulties in receiving material on 

Andarai-LengSis (BA 142), Fda. Lagoa Encantada, 22 

May 1989 (fl), Mattos Silva et al. 2842 (CEPEC); Eunapolis, 

loan from KW and LE, Blanchet 255 7 preserved at K 

Itabela, Itamaraju, Prado, 8 Jul 1970 (fl), Santos 920 

is here selected as the lectotype of M. paniculata. (CEPEC); Itamaraju, Fazenda Boa Vista, 27 Jul 1971 (fl), 

The specific epithet is sometimes misspelled as Monteiro 23498 (RB); Barra da Estiva, 13?38'S, 41?07'W, 

panniculata (Schumann 1889, 1891), but it was origi- 21 Jun 1978 (fl), Vaillant 21 (CEPEC, HRB, MG, RB). 

nally published as paniculata and should be main- MINAS GERAIS: Mun. Serro, 3 km from Serro, 12 May 1945 

tained as such. 

(fl), Williams & Assis 6861 (F, GH, K, NY, RB); Itaobim, 4 

Apr 1979 (fl), Rizzini & Mattos 1126 (RB); Mun. Aracuai, 

16?53'S, 41056'W, 6 Jul 1981 (fl-fr), Salim J. Filho 148 

(HRB, MG); Mun. Jequitinhonha, 16?33'S, 41?06'W, 8 Jul 

1981 

Key to the varieties of (fl), Salim J. Filho 159 (HRB, MG). Rio DE JANEIRO: 

Molopanthera paniculata 

Rio de Janeiro, Lagoinha, Silvestre, 16 Jun 1959 (fl-fr), 

1. Leaves glabrous beneath, glabrous-shiny above; Duarte 4842 (RB110068) (F[2], RB). 

stipules glabrous outside; lower nodes of the 

inflorescences glabrous ................. 1a. var. paniculata Local name. Brazil: saco de mono (Bahia, Mori 

1. Leaves densely golden-pubescent beneath, coarse- et al. 10328). 

pubescent above; stipules pubescent outside; 

lower nodes of the inflorescences short- 

This variety is easily distinguishable from var. 

pubescent ......................................... Ib. var. burchellii 

burchellii) in having by the characters given in the 

key; it occurs mainly in the provinces of Bahia and 

northernmost Minas Gerais. Individuals of this varila. 

Molopanthera paniculata Turczaninow var. ety have been reported to reach 30 m in height and 80 

paniculata. Figs. 3T, 4N, 85A-D, 86 cm dbh (Mori et al. 10328). 


disturbed seasonal forests, remnant in pastures, of 

Brazil, mainly in the provinces of Bahia, Espirito 

Santo, and northernmost Minas Gerais; commonly in 

clay-siliceous soils. Flowering specimens were col- 

lected in May, July, and August. Fruiting specimens 

were collected in August and October. 

Pollination ecology. The flowers of this variety 

(and probably the other variety) were reported to be 

cream-white, fragrant, and visited (and most likely 

pollinated) by small bees (label data of Bautista & 

Orlandi 985). 

Specimens examined. BRAZIL. ESPIRITO SANTO: 

Reserva Forestal Linhares, near rd. 161, talhao 604, 16 Jul 

1973 (fl), Spada 293 (RB); Reserva Forestal Linhares, near 

rd. 143, talhao 403, 10 Jul 1973 (fl), Spada 289 (RB); 

Reserva Forestal Linhares, near quadra 1, talhao 605, 2 Jul 

195 

lb. Molopanthera paniculata Turczaninow var. 

burchellii (J. D. Hooker) K. Schumann in Martius, 

Fl. Bras. 6(6): 200. 1889. Molopanthera burchellii 

J. D. Hooker in Bentham, G. & J. D. Hooker, Gen. 

P1. 2: 42. 1873. Type. Brazil. Minas Gerais: Lagoa 

Santa, 1863-65 (fr), Warming s.n. (B*; lectotype, 

F, here selected; isolectotypes, C-n.v., GH). 

Figs. 1 B, 85, 86 


disturbed forests of southeastern Brazil, mainly in the 

provinces of Minas Gerais and Rio de Janeiro. Flow- 

ering specimens were collected in February, May, and 

June. Fruiting specimens were collected in May, July, 

and August. 

Specimens examined. BRAZIL. MINAS GERAIS: 

Caratinga Biological Station, 19?40'S, 41?50'W, 24 Aug


FIG. 86. Distribution of Afolopanthera paniculata var. 

paniculata (circles) and M. paniculata var. burchelli (tri- 

angles). 

1983 (fr), Strier 1007 (NY); Mun. Tombos, Fazenda da 

Cachoeira, 29 Jul 1935 (fr), Barreto 1789 (JBBH16686) (A, 

F); Mun. Santa Luzia, Lagoa Santa, 3 May 1934 (fl-fr), 

Barreto 8305 (JBBH26371) (F); Mun. Santa Luzia, Lagoa 

Santa, 8 May 1937 (fl), Magalhaes s.n. (IAN); Mun. Belo 

Horizonte, Bento Pires (Eng. Nogueira), Feb 1945 (fl), Williams 

5066 (F, GH); Grno Mogol, valley of Riberio creek, 

ca. 900 m, 24 May 1987 (fl), Pirani & Mello Silva SPF47733 

(NY). Rio DE JANEIRO: Morro de S. Joao, Botafogo, Jul 1914 

(fl), Kulhmann 1221 (RB); Horto Florestal, 9 Jun 1927 (fl), 

Personnel ofthe Herbario Floresta s.n. (RB); Depto. Veiase, 

between Arapinga and Caman, 21 Apr 1935 (fl), Kuhlmann 

82 (RB149083) (RB). 

Local name. Brazil: pesquim (Minas Gerais, 

Barreto 1789). 

This variety is distinct from the typical variety in 

having blades densely golden-pubescent below and 

coarse sparsely pubescent above, stipules externally 

pubescent, and lower nodes of the central rachis of 

inflorescences short-pubescent, correlated with its 

geographical distribution. The minute bent corollas 

and small bilobed capsules are morphologically consistent 

throughout the range of the species. Future 

collections throughout the range of this species may 

prove that the two varieties here recognized are not 

distinguishable. 

Schumann annotated Warming s.n. (the lectotype 

here selected) as "Molopanthera panniculata Turcz. 

var. scabrida K. Schum. et Burchell II - det. 

Schumann." 

DOLICHODELPHYS 

9. Dolichodelphys K. Schumann & K. Krause in E. 

Ule, Verh. Bot. Vereins. Prov. Brandenburg 50: 

102, fig. 1 (103). 1908; Standley, Publ. Field 

Columbian Mus., Bot. Ser. 8: 159. 1930; 

Steyermark in Lasser & Steyermark, Fl. Venez. 

9(1): 262-265, fig. 42. 1974; Delprete in Harling 

& Andersson, Fl. Ecuador (in press). Type species. 

Dolichodelphys chlorocrater K. Schumann 

& K. Krause. 

Shrubs to treelets, much branched shrubs to 

single-stemmed trees; bark grayish. Stipules 

interpetiolar, free at base, often connate to the petiole, 

deltoid to narrowly triangular, persistent. Leaves 

petiolate, lanceolate to obovate, acuminate to obtuse, 

mucronate or acuminate at apex; petioles thickened 

at base; domatia absent. Inflorescences terminal, solitary, 

pyramidal, openly paniculate; lateral branches 

opposite to subopposite, decussate. Flowers protandrous, 

extremely variable in size within the same inflorescence, 

first flowers to 3 cm long, gradually 

smaller toward the end of flowering season. Calyx 

cupular, lobes ovate to lanceolate, persistent. Corolla 

campanulate (in earlier, big flowers) to urceolate (in 

laster, smaller flowers), membranous, cream-white to 

yellowish green; tube cylindrical to narrowly funnelform; 

lobes 5(-6); aestivation narrowly imbricate. 

Stamens 5(-6), included, equal or subequal; anthers 

cylindrical, narrowly oblong, green, dorsifixed near 

the base, base sagittate, dehiscing by longitudinal slits. 

Pollen tricolporate, exine openly reticulate. Style included; 

style branches membranous, reflexed. Ovary 

2-celled, placentation axile, turbinate to narrowly 

obconical, glabrous, with many ovules in each locule 

horizontally inserted. Capsules woody, long, narrowly 

cylindrical, pluricostate; dehiscing septicidally, 

the septa detaching and expanding. Seeds ovoid, 

rhomboid to trapezoid, not winged, exotesta reticulate.


Dolichodelphys is a rare monotypic genus endemic 

to northwestern South America. Its name is derived 

from the Greek Dolicho- (= long) and delphys (= 

ovary), referring to the slender elongated capsules 

typical of this genus. Dolichodelphys is readily distinguishable 

by its inflorescences with the first internal 

flowers campanulate and 1-3 cm long, and gradually 

smaller on outer branchlets of inflorescence 

(down to 3-5 mm long), its yellowish green to creamwhite 

thinly membranous flowers, and its long-slender 

multicostate capsules (see Fig. 87A-G). This 

threatened species is mainly encountered in the Cordilleras 

of Colombia, and few rare collections are 

available from other northwestern South American 

countries. 

Schumann and Krause (1908c) described this genus 

from plants with only immature capsules and 

flower buds. They assigned it to the "Cinchonoideae- 

Gardenieae group" and related to Retiniphyllum, 

Synisoon [= Retiniphyllum], and Tocoyena, because 

of its two-parted ovaries and linear calyx lobes. 

Steyermark (1974) did not cite this genus in Flora de 

Venezuela, but treated the specimens of 

Dolichodelphys as Rustia longifolia. Robbrecht (1988, 

1993) maintained Dolichodelphys in the Gardenieae, 

but in his survey of the tribe (Robbrecht & Puff, 1986) 

lar, acuminate, glabrous outside and inside, 13-30 x 

8-15 mm, older stipules sometimes splitting into two 

parts, giving the impression of being bilobed. Leaves 

31-62 x 10-30 cm, L/W 2:1 to 3:1, lanceolate to oblanceolate 

to obovate, acute at base, acuminate to obtuse 

and mucronate at apex, sometimes tapering to an 

acuminate apex up to 3 cm long; dark green above, 

pale green below, semi-coriaceous; drying brown to 

olive-green, semi-coriaceous to stiff-chartaceous; glabrous 


glabrous (hirtellous in Fernandez & Oyos 182), secondary 

veins 17-25 each side, tertiary veins starting 

subparallel and reticulate in the middle; petioles 10- 

15 mm long, 3-5 mm thick, terete to costate, sometimes 

adaxially flattened. Inflorescences openly paniculate 

with opposite decussate branches with terminal 

cymules of 10-25 flowers; inflorescence (15-)25-65 

cm long, basal branches 15-28 cm long, apical 

branches 4-7 cm long, lateral branches 3-4 pairs, 

basal portion of axis not branched (4-)15-25 cm long; 

rachis and branches laterally compressed, 4-5 costate, 

glabrous (rarely short hirtellous); bracts subtending 

basal branches foliose, to 7 x 3 cm, upper bracts 10- 

20 x 3-6 mm, lanceolate to linear; bracteoles subtending 

flowers 1-2 mm x 1-2 mm, ovate to linear. Flowers 

pedicellate, pedicels 1-11 mm long, glabrous (but 

it was not mentioned. Its terminal, paniculate inflo- hirtellous in Fernandez & Oyos 182); hypanthium 

rescences, seed coat morphology, bilocular ovary, and obconical, 1-5 x 1-2 mm, glabrous. Calyx cupular, 

T-shaped placenta resemble those of Raritebe 0.5-2 x 2-6 mm, glabrous; lobes ovate to narrowly 

(Isertieae s.l.). Nevertheless, Dolichodelphys is ten- lanceolate (margins entire to fringed), 1-6 x 1-4 mm, 

tatively included in the Rondeletieae s.l. (Delprete, ciliolate. Corolla size and shape extremely variable, 

1996d), although future investigations may result in first largest flowers opening, 1.5-3 cm long, campanuits 

placement elsewhere. 

late; later flowers gradually smaller, (1-)6-8 mm long, 

tubular to urceolate, the smallest ones lacking sexual 

structures; tube cylindrical to narrowly funnelform, 

1-20 mm x 1-6 mm, glabrous outside and inside, 

1. Dolichodelphys chlorocrater K. Schumann & K. whitish cream to yellowish green to pale-green; lobes 

Krause, Verh. Bot. Vereins. Prov. Brandenburg 5(-6), 1-4 x 1-4 mm, triangular to ovate, sometimes 

50: 102. 1908. Type. Peru. Loreto: Cerro de 

mucronate, glabrous outside and inside, margins en- 

Escaler, 1200 m, Jan 1903 (fl), Ule 6774 (B*, tire to fringed, cream-white to yellow (rarely greenphoto-B 

at F; lectotype, K, selected by Delprete, ish). Stamens 5(-6), included, attached V2 to '/io from 

1999b; isolectotypes, frag-F, G[2], frag-IAN). the base of the tube; filaments 1-7 mm long, flattened, 

Figs. 3S, 12A,B, 19A,B, 84, 87A-G 

glabrous; anthers cylindrical, narrowly oblong, green, 

Rustia longifolia Standley, Publ. Field Columbian Mus., 4-7 x 1-2 mm, base sagittate, dehiscing by longitu- 

Bot. Ser. 8: 159. 1930. Type. Colombia. Caqueta: dinal slits. Pollen exine reticulate, with denser reticu- 

Between Santa Marta and Marsella, Aug 1926 (fr), lation at the 

Woronow & Jusepczuk 6426 

poles. Style included, 5-23 mm long, 

(holotype, LE-n.v.; 

glabrous; style branches ovate to obovate, membraisotypes, 

F, LE-n.v.). 

nous, reversed, entire to fringed, ca. 3 mm long, gla- 

Shrubs to treelets to 8(-10) m tall, to 10 cm dbh, 

much branched, rarely a single-stemmed treelet; bark 

grayish-whitish. Leafy branchlets glabrous to golden 

pubescent, slender, tetragonous to costate, grayish; 

lenticels absent. Stipules deltoid to narrowly triangubrous, 

stigmatic surface smooth. Capsules narrowly 

cylindrical, pluri-costate, apex truncate, 14-23 x 3.5- 

5 mm, dark brown, glabrous (hirtellous in Hernandez 

& Hoyos 182) throughout, without lenticels; disk truncate, 

not exceeding the calyx. Seeds ovoid, rhomboid to 

trapezoid, 0.1-0.3 long, not winged, exotesta reticulate.


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secondary rain forests and cloud forests, to second- 

ary moist forests, of medium elevations, 400-1400 m, 

on the Andes of Venezuela, Colombia, Ecuador, and 

Peru. Flowering specimens were collected in January, 

March, June, August, October, November, and De- 

cember. Specimens in intermediate late flowering- 

early fruiting stage have been collected in July, and 

September. Fruiting specimens were collected in Feb- 

ruary, March, and August. 

Reproductive biology. The inflorescences of this 

species are laxly paniculate, and its flowering pattern 

is unique. The first flowers to open are large and campanulate 

(1.5-3 cm long). As the blooming continues 

on outer lateral branchlets, the flowers decrease in size 

(down to 3-5 mm), gradually changing shape from 

campanulate to short-tubular to urceolate (Fig. 87B- 

E). Also, as the flowers become smaller, they gradually 

lose their reproductive parts, first only with functional 

anthers (and non-functional ovary), down to the 

minute flowers, and the most reduced without any 

sexual organs and entirely sterile. 

Probably the flowering strategy of this species 

relies on the memory of its pollinators (Larry Gilbert, 

pers. comm.): the first few large perfect flowers have 

the main function to attract pollinators, the following 

flowers gradually diminish in corolla size (but repro- 

ductive organs are still functional), economizing en- 

ergy devoted to attraction. The pollinators, remem- 

bering the locality of the individuals, continue to visit 

the gradually less visible, but still functional flowers. 

The pollinators of this species have never been ob- 

served, but considering the size and shape of the flow- 

ers, they are probably butterflies. 

Specimens examined. COLOMBIA. ANTIOQUIA: Valley 

of Rio Anori, Planta Providencia, 26 km S of Saragoza, 

07?13'N, 75?03'W, 400-700 m, 10 Feb 1975 (fr), Denslow 

2680 (COL, US); Planta Providencia, 30 Sep 1984 (fl-fr), 

Zarucchi 3338 (MO, NY); Planta Providencia, 27 Jun 1976 

(fl), Shepherd 459 (COL, MO[2]); Hwy. Medellin-Bogota, 

Rio Samani-Rio Claro, San Luis, 600 m, 7 Jul 1981 (fl-fr), 

Herndndez & Hoyos 182 (COL); Mun. San Luis, quebrada 

La Cristalina, 06?N, 74?45'W, 770 m, 25 Aug 1987 (fr), 

Ramirez & Cdrdenas 179 (MO), 1017 (COL, MO), 1501 

(COL), 1620 (COL), 1879 (MO). CAQUETA: Florencia, Cerro 

de la Sardina, 500 m, 30 Mar 1940 (fl), Cuatrecasas 8926 

(COL, US). META: Parque Nacional La Macarena, macizo 

Renjifo, 1300-1900 m, 6-20 Jan 1951 (fl), Idrobo & Schultes 

1011 (COL, US); Parque Nacional La Macarena, cafto 20, 

850 m, 2 Mar 1962 (fr), Idrobo 4874 (COL[2]); Parque 

Nacional La Macarena, Central mtns., N ridge, 1400 m, 27 

Dec 1949 (fl), Philipson & Idrobo 1955 (COL[2], US[2]); 

Parque Nacional La Macarena, Rio Guapaya, 450 m, 26 Nov 

1949 (fl), Philipson et al. 1576 (COL, US). PUTUMAYO: 

Umbria, 00?54N, 76?10'W, 325 m, Oct 1930 (fl), Klug 1804 

(A, K, MO, NY, US); Rio Rumiyaco, 800 m, 9 Aug 1964 

(fl), Soejarto et al. 1287 (COL, GH, US[2]). 

VENEZUELA. CAABOBo: On steep rocky slopes above 

Rio San Gean, above La Toma, E of Los Tanques, S of 

Borburata, 1000 m, 6 Dec 1967 (fr), Steyermark 100495 

(G[2], VEN). WrIHour LOCALrTY: "Venezuela," Linden 1604 

(BR, photo-BR at VEN). 

ECUADOR. MORONA-SANTIAGO: Along new rd. 

M6ndez-Morona, km 25-30, forest remnant, 800 m, 21 Aug 

1989 (fl), van der Werff& Gudinio 11326 (QCNE). 

PERU. LORETO: Prov. Alto Amazonas, N slopes of Cerro 

Campanquiz, Pongo de Manseriche, right bank of Rio 

Marafion, 300-550 m, 19-21 Oct 1962 (fl), Wurdack 2348 

(US). SAN MARTIN: Prov. Lamas, Rio Curiyacu, affluent of 

Rio Cumbasa, 8 km above San Antonio, 450 m, 6 Nov 1937 

(fl), Belshaw 3593 (BH, GH, K, LL, NY, UC, US). 

Standley (1930b) placed Rustia longifolia Standl. 

close to R. splendens (= R. thibaudioides). Steyermark 

(1974) maintained Standley's (1930b) point of view 

(both authors without seeing mature flowering material), 

keying it and describing it under Rustia. Simpson 

(1976) and Andersson (1992) erroneously reported R. 

longifolia to be synonymous with Tresanthera 

condamineoides. The type of R. longifolia (Woronow 

& Jusepczuk 6426, F) has leaves and a portion of the 

inflorescence that exactly matches those of D. 

chlorocrater, with which is here treated as synonymous. 

Ferndndez & Oyos 182 (Prov. Antioquia, Colombia) 

differs from the rest of the collections of this species 

in having young vegetative parts, rachis, and capsules 

golden-hirtellous. I do not consider this character 

alone sufficient so as to assign any formal taxonomic 

recognition to this collection. 

PARACHIMARRHIS 

10. Parachimarrhis Ducke, Arch. Jard. Bot. Rio de 

Janeiro 3: 253. 1922. Type species. P.breviloba 

Ducke. 

Trees; bole irregular, without buttresses; bark 

smooth; wood brownish yellow, hard. Stipules 

interpetiolar, above leaf attachment, free at base, ad- 

nate to the petioles, caducous, leaving a scar encir- 

cling the stem above the petiole attachments. Leaves 

elliptic to ovate to obovate, usually acuminate at apex; 

microscopically glandular-punctate on the abaxial 

side; petioles slightly thickened at base; domatia ab- 

sent (exceptionally a few short hairs). Inflorescences 

terminal corymbiform panicles, with opposite to


subopposite lateral branches terminating in cymules. 

Flowers protandrous, sessile to short-pedicellate, very 

fragrant; hypanthium obconical. Calyx reduced, with 

acute lobes, persistent; in some of the flowers one 

calyx lobe expanded into a small, white calycophyll. 

Corolla short-tubular, shortly lobed, with reflexed 

lobes, white; tube cylindrical, with a ring of pubescent 

hairs inside on the tube; lobes 5, aestivation narrowly 

imbricate, superficially resembling valvate aestivation. 

Stamens 5, alternate to the petals, exserted; 

filaments attached on the upper part of the tube, slender, 

basally flattened, with a tuft of pilose hairs at base; 

anthers elliptic, dorsifixed, dehiscing by lateral slits. 

Pollen tricolporate, exine sparsely, irregularly reticulate. 

Style exserted; style branches rounded to ovate, 

reflexed at maturity. Ovary 2-celled, placentation 

horizontal on the septum, obconical, with many ovules 

in each locule; immature fruits green and semi-fleshy 

when fresh. Capsules woody, bi-globular; dehiscing 

loculicidally, disk septicidal dehiscence absent. Seeds 

hemi-elliptic, dorsiventrally flattened, laterally attached 

in two rows each locule, margin entire, with 

lateral wing. 

Ducke (1922) treated Parachimarrhis as closely 

related to Chimarrhis (hence the name), stating that 

it differed by having deeply laciniate calyx, a shortly 

lobed corolla, and capsules with few seeds per locule. 

In Chimarrhis the inflorescences are axillary (vs. 

terminal in Parachimarrhis), the flowers are 

protogynous (vs. protandrous), the capsules are mostly 

obconical to turbinate (vs. biglobular), with peltate 

placentation (vs. horizontal) and septicidal (vs. loculicidal) 

dehiscence, minute seeds with deeply fringed, 

concentric wing irregular in outline (vs. small and with 

entire, lateral wing hemielliptic in outline), and testa 

reticulate (vs. minutely foveolate; cf. Fig. 12E). 

Because of the terminal inflorescences, protandrous 

flowers with exserted anthers, hemi-elliptic 

seeds (Figs. 12G, 88A-G), seed inserted horizontal 

on the central placenta, testa sculpturing (Fig. 12G), 

and the biglobular capsules (Fig. 87F), Parachimarrhis 

is related to Simira (Fig. 12E,F). From Simira it 

differs in having much smaller capsules and seeds, and 

yellowish wood (most species of Simira have wood 

with a reddish tinge when exposed to air). 

1. Parachimarrhis breviloba Ducke, Arch. Jard. Bot. 

Rio de Janeiro 3: 254. 1922. Type. Brazil. Para: 

Rio Tapajoz, close to Maranhaosinho waterfalls, 

Maria Luisa, 6 Dec 1919 (fl), Ducke s.n. (RB 

15687) (lectotype, RB, here selected; isotypes, B*, 

F, US; photo-B at F, MO, NY). 

Figs. 12G, 88A-G, 89 

Trees to 20 m tall, to 40-50 cm dbh, without buttresses; 

bark smooth, pale brown; wood brownish 

yellow, hard. Leafy branchlets 2.5-5 mm thick, terete 

to compressed, glabrous to puberulent, 

cinnamoneous; older branches glabrate, terete, brown; 

lenticels sparse, whitish, linear. Stipules free at base, 

contorted in bud, narrowly triangular, acuminate, appressed 

puberulent to gray-sericeous outside, glabrous 

with basal colleters inside, 15-27 x 2-4 mm, readily 

caducous, deep-green, leaving a white scar encircling 

the stem above petiole attachments, 0.5-1 mm wide. 

Leaves (10-)13-30 x 4-10 cm, L/W = 2.5:1 to 3:1, 

elliptic to oblong-obovate, acute-decurrent at base, 

acute and acuminate at apex, the narrowly triangular 

acumen 1.5-2.5 cm long; dark green above, pale green 

below, foliaceous to thinly coriaceous; drying olivegreen 

to pale-brown, stiff-chartaceous to coriaceous; 

glabrous above and below; primary, secondary and 

tertiary veins glabrous to very sparsely puberulent 

below; secondary veins 10-18 each side; tertiary veins 

subparallel; petioles 10-26 mm long, 1-2 mm thick, 

adaxially concave, glabrous; domatia absent or a few 

puberulent hair. Inflorescences corymbiform 

panicles, with opposite decussate branches, short-pedunculate, 

5.5-11 cm long; basal branches 2-8 cm 

long, lateral branches 2-5 pairs; rachis basally terete 

to subterete, decussately compressed distally, without 

lenticels, rachis and branches densely puberulent; 

flowers in terminal few-flowered cymules. Flowers 

sessile to short-pedicellate; pedicels to 1.5 mm long, 

puberulent; hypanthium obovoid, faintly bilobed, 0.5- 

0.9 x 0.6-0.7 mm, puberulent. Calyx minutely 

cupular, short-lobed, 1-1.5 x 1.5-2 mm, glabrous; 

lobes narrowly triangular, 0.6-1.7 x 0.3-0.6 mm; several 

flowers of the same inflorescences with one calyx 

lobe expanded into small calycophyll. 

Calycophylls 7-13 x 3-5 mm, blades elliptic to ovate, 

with the adaxial side covered with a thick mat of glandular 

hairs (0.2-0.4 mm long). Corolla tubular-campanulate, 

shortly lobed, the lobes reflexed at anthesis, 

3-4 mm long, glabrous throughout, white to 

cream-white; tube short cylindrical, 2-3 x 0.5-0.6 

mm; lobes 5, 0.5-0.7 x 0.4-0.6 mm, oblong-linear, 

round at apex. Stamens 5, exserted well above the 

corolla, equal in length, attached 1-1.5 mm from the 

base of the tube; filaments 4-5 mm long, slender-terete, 

basally flattened and short-pubescent; anthers 

narrowly elliptic, 1-1.5 x 0.3-0.4 mm, dorsifixed near 

the middle. Pollen exine irregularly reticulate. Style 

exserted, elongating only after anthers dehiscence, 3-


,'- ; 

, 

. . .o,'... 

~" 

FIG. 88. Parachimarrhis breviloba. (A-E from Fundaci6n Biol6gica Puerto Rastrojo 5052, NY, and Ducke (RB15567), 

RB, lectotype; F, G from McDaniel & Rimachi 20503, NY). A. Habit of inflorescence, with two mature leaves. B. Flower 

with exserted anthers. C. Detail of expanded calyx lobe. D. Stamen. E. Apical portion of style. F. Mature capsule. G. Seed.


0 200 no so. 10no 

oI 

m O 

a10~~~ 7~FG0 D o to ma f 0 o 

5 mm long, terete, glabrous; style branches clavate, 

reflexed, 0.5-1 mm long, stigmatic surface minutely 

papillose. Capsules sessile to short-pedicellate, 

strongly bilobed the two sides globose, rounded at 

base and at apex, 3-4.5 x 4.5-6 mm, without lenticels, 

rust-brown, glabrous throughout; disk not exceeding 

the calyx; disk septicidal dehiscence absent. Seeds 

hemi-elliptic in outline,-2.2-4.5 x 1.5-2.5 mm, with 

lateral wing with entire margin; seed embryo about 

one half the size of the seed outline. 


secondary rain forests of lowland, seasonally inun- 

dated, Amazonian rain forest of Colombia, Ecuador, 

Peru, and Brazil. Flowering specimens were collected 

in December. One specimen at late flowering-early 

fruiting stage was collected in March. Fruiting speci- 

mens were collected in January, March, April, July, 

and December. 

Specimens examined. COLOMBIA. AMAZONAS: 

Parque Nacional Cahuinari, Rio Bernardo, 01?28'S, 

70?45'W, 7 Mar 1991 (fl-fr), Fundaci6n Biol6gica Puerto 

Rastrepo 5052 (NY). 

PERU. LORETO: Rio Mazan, above La Libertad, ca. 35 

km above Mazan, 150 m, 10 Jul 1976 (fr), Gentry & Revilla 

16645 (NY); Maynas, Dtto. Iquitos, Rio Mom6n, ca. 2 km 

FIG. 89. Distribution of Parachimarrhis breviloba. 

above Rio Nanay, 13 Mar 1976 (fr), McDaniel & Rimachi 

20503 (NY); Maynas, Dtto. Puchana, Rio Amazonas, 

Caserio de Santa Maria de Ojeal, Quebrada de Sinchicuy, 

90 m, 13 Apr 1993 (fr), Rimachi 10501 (NY). 

BRAZIL. PAA: Rio Tapajoz, vic. of Francez, 19 Dec 

1919 (fr), Ducke s.n. (RB 15400)(B* [photo-B at F, MO, 

NY], F, RB, US, paratypes). RorND6N: Rio Machado, curso 

inferior, Jan 1981 (fr), Goulding 1048 (MG). 

Local names and uses. Peru: Huacamayo micuna 

(Rimachi 10501), quillobordon (McDaniel & Rimachi 

20503). Ducke (1922) reported that this species has 

good wood, but no report about its use has been found. 

Molopanthera breviloba is easily distinguishable 

by its small (3-4.5 x 4.5-6 mm), bilobed capsules, 

and minute (2.2-4.5 x 1.5-2.5 mm) seeds that are 

laterally attached and hemi-elliptic in outline. It is also 

typical in having flowers with small (7-13 x 3-5 mm) 

calycophylls covered with a thick mat of glandular 

hairs. The function of this matt of hairs is unknown. 

Molopanthera breviloba is a little-known species 

represented by mid-canopy trees, known from a few 

collections from Amazonian forests that have been 

little collected. In the last decades a few more collec- 

tions have shown its wide distribution throughout the 

Amazon basin.

Literature Cited 203 

ACKNOWLEDGMENTS 

My personal gratitude goes to Billie Turner (TEX), 

who was my doctoral supervisor during my Austin 

years (1991-1996); not only has he assisted me with 

his supervision, but he also helped sponsor my travel 

expenses to Costa Rica in 1991 and Ecuador in 1992, 

and the direct expenses for SEM studies of seeds and 

pollen in 1994. My extreme appreciation goes to 

Grady Webster (DAV), the first botanical teacher who 

guided my first steps through the cloud forests of 

Ecuador, and who was the first to inspire me to study 

Rubiaceae (sitting on a log, at sunset, in the outdoors 

near Austin). Larry Gilbert (TEX) showed me the 

beauty and diversity of tropical forests in Corcovado 

National Park, Costa Rica; from him I learned the 

attitude of wonder and curiosity that is present inside 

of me. The contribution by Paul Fryxell (TEX) toward 

the writing of this monograph has been among the 

most valuable and influential, helping me to solve 

problems ranging from taxonomic history to typification, 

from nomenclatural problems to species concepts, 

and carefully proofreading my entire doctoral 

dissertation. My most sincere thanks go to Joseph 

Kirkbride (USDA), who read every single word of the 

final version of the manuscript, contributing many 

valuable suggestions and helping to solve nomenclatural 

problems. My particular thanks go to Charlotte 

Taylor (MO) and John Dwyer (MO), who, in fall 1990, 

suggested that I study Rustia or Chimarrhis; following 

their suggestions the original study terminated in 

the ongoing monographic treatment of the entire Condamineeae 

(sensu Hooker). I am grateful to Charlotte 

Taylor for reviewing the manuscript and for helpful 

comments. I am also grateful to Rupert Bareby (NY), 

who improved the Latin diagnoses of the new taxa 

described here and elsewhere, and for helping me 

solve many nomenclatural and taxonomical problems. 

U, UB, UC, US, USCH, and W (acronyms according 

to Holmgren et al., 1990). 

I owe final thanks to the many colleagues and 

friends, some of them no longer with us, who contributed 

to my research on the Condamineeae: Linda 

Albert de Escobar, Lennart Andersson, Patricia 

Apreza, Alberto Areces, William Burger, John Clark, 

Hermes Cuadros, Todd Barkman, Brian Boom, 

Birgitta Bremer, Joan Busquets, Daryon Cardenas, 

Arthur Cronquist, Miles Davis, Ted Delevoryas, 

Bianca Dickerson, Imre Eifert, Javier Francisco- 

Ortega, Alina Freire-Fierro, Javier Fuertes, Fabio 

Garcia Cossio, Al Gentry, Steve Ginzbarg, Doug 

Goldman, Mario Gomes, Vera Gomes-Klein, Favio 

Gonzales, Barry Hammel, Jim Henrickson, Lauritz 

Holm-Nielsen, Otto Huber, Robert Jansen, Jaime 

Jaramillo, Dennis Kearns, Michael Kiehn, David 

Lorence, Jim Luteyn, Tom Mabry, Mark Mayfield, 

June McCaskill, Tim McDowell, John Mendenhall, 

Philip Morat, Scott Mori, Michael Nee, David Neill, 

Guy Nesom, Benjamin 0llgaard, Jose Panero, Cristina 

Panfet-Valdez, Henrik Pedersen, Claes Persson, Luis 

Poveda, Alan Prather, Christian Puff, Don Pullen, 

Montserrat Rios, Elmar Robbrecht, Katya 

Romoleroux, Johan Rova, Jan Saunders de Palacios, 

Beryl Simpson, Neil Snow, Bertil Stahl, Stella Sylva, 

Armen Takhtajahn, Ivan Valdespino, Renato 

Valencia, Angela Verduga, Justin Williams, and Tom 

Zanoni. 

During my collecting trips in the tropical forests 

of South America I was hosted by several native tribes, 

among them the Achuare, Embera, Kofan, Kogi, 

Secoya, Sharanahua, Shuar, Tikuna, Tukano, and 

Waorani. I regard the members of these tribes as my 

ultimate masters, and their ancestral wisdom should 

be the inspiration for the morals and ethics of modem 

societies. 

Jack Neff (TEX) identified all the insects that visited 

the flowers of the taxa presented in this study. Jim LITERATURE CITED 

Luteyn (NY) was always patient in answering all the 

editorial questions that came up as the manuscript was 

reaching its final form. The illustrations were all 

drawn by myself during the last three years in Austin 

and my first year in New York. The SEM and field 

photographs are all taken by myself, unless otherwise 

indicated in figure captions. 

I am grateful to the directors and staff of the following 

institutions for making their specimens available 

(a total of nearly 13,000), hosting me, and helping 

in the field: A, AAU, B, BH, BR, C, CAS, CAY, 

CEPEC, CHOCO, CLEMS, CM, COL, CR, DAV, 

DS, F, FI, FLAS, FSU, G, GA, GB, GH, GOET, HAC, 

HAJB, HRB, IAN, INPA, JBGP, JEPS, K, LL, M, 

MA, MG, MO, NY, P, QCA, QCNE, RB, S, SP, TEX, 

Adams, R. J. & J. K. Morton. 1972. An improved technique 

for examining pollen under the scanning electron 

microscope. Pollen & Spores 14: 203-212. 

Adanson, M. 1763. Rubiaceae [as "Aparines"]. Families 

des Plantes 140-147. 

Aiello, A. 1979. A reexamination ofPortlandia (Rubiaceae) 

and associated taxa. J. Arnold Arb. 60: 38-123. 

Andersson, L. 1992. A provisional checklist of Neotropical 

Rubiaceae. Scripta Bot. Belg. 1: 1-200. 

- . 1994. Rubiaceae-Cinchoneae. In: G. Harling & L. 

Andersson (eds.), Flora of Ecuador 50: 3-82. 

& C. Persson. 1991. Circumscription of the tribe 

Cinchoneae (Rubiaceae)-A cladistic approach. PI. Syst. 

Evol. 178: 65-94. 

Aublet, J. B. C. F. 1775. Histoire des plantes de la Guiane 

Francoise. P. F. Didot, Paris. 

Baillon, H. E. 1879. Les ailes seminales de certainnes 

Rubiacees. Adansonia 12: 296-302.


- . 1880. Rubiac6es-Dipsacacees. Histoire naturelle des 

plantes 7: 257-503. Leipzig (L. Hachette et Cie), Paris. 

Barbosa, M. R. de V. & A. L. Peixoto. 1989. As esp6cies 

de Simira (Rubiaceae, Rondeletieae) da Amaz6nia 

brasileira. Acta Amazon. 19: 27-46. 

Benoist, M. R. 1920. Bathysa difformis. Bull. Mus. Hist. 

Nat. Paris 26: 185-186. 

- . 1933. Pseudochimarrhis difformis. Bois de la 

Guyane. Arch. Bot. 5 (Mem. 1): 264-265. 

Bentham, G. 1844. The botany of the voyage H. M. S. Sulphur, 

under the command of Captain Edward Belcher,... 

during the years 1836-1842. Smith, Elder & Co., London. 

Blackmore, S. & S. H. Barnes. 1986. Harmomeghatic 

mechanisms in pollen grains. Linn. Soc. Symposium Ser. 

12: 137-149. 

Boom, B. M. 1984. A revision of Isertia (Isertieae: Rubiaceae). 

Brittonia 36: 425-454. 

& M. T. V. do A. Campos. 1991. A preliminary 

account of the Rubiaceae of a central Amazonian terra 

firme forest. Bol. Mus. Paraense Emilio Goeldi, n.s., Bot. 

7(2): 223-247. 

Borhidi, A. 1982 [1983]. Studies in Rondeletieae (Rubiaceae) 

III: The genera Rogieria and Arachnotryx. Acta 

Bot. Acad. Sci. Hung. 28: 65-71. 

--- . 1987 [1989]. Studies in Rondeletieae (Rubiaceae) 

X. New combinations of Central American taxa. Acta 

Bot. Hung. 33: 301-303. 

--- . 1991. Phytogeography and vegetation ecology of 

Cuba. English translation by A. Borhidi, J. Podani & I. 

K. Kecskes. Szegedi Nyomda, Szeged, Hungary. 

& M. Fernandez Zequeira. 1981 a [1982]. Studies 

in Rondeletieae (Rubiaceae) I. A new genus: Roigella. 

Acta Bot. Hung. 27: 309-312. 

--- & .- 1981b [1982]. Studies in Rondeletieae 

(Rubiaceae) II. A new genus: Suberanthus. Acta Bot. 

Hung. 27: 313-316. 

- & M. Jarai-Koml6di. 1983 [1986]. Studies in Rondeletieae 

(Rubiaceae), IV. A new genus: Javorkaea. Acta 

Bot. Hung. 29: 13-27. 

, & M. Moncada. 1980 [1981]. Acunaeanthus, 

a new genus of Rubiaceae. Acta Bot. Acad. Sci. 

Hung. 26: 277-287. 

Bremekamp, C. E. B. 1934. Notes on the Rubiaceae of 

Surinam. Recueil Trav. Bot. Neerl. 31: 248-308. 

. 1940. On Urophyllum Wall. (Rubiaceae) and its 

nearest allies. Recueil Trav. Bot. Neerl. 37: 171-197. 

-- . 1954a. The identity of Simira tinctoria Aubl. Acta 

Bot. Neerl. 3: 150-153. 

1954b. Les sousfamilles et les tribus des Rubiaceae. 

Rapports et communications parvenus avant le congress 

aux section 2, 4, 5 et 6, Huitiem Congres International 

de Botanique, Paris, 1954, pp. 113-114. 

-- . 1966. Remarks on the position, the delimitation, 

and the subdivision of the Rubiaceae. Acta Bot. Neerl. 

15: 1-33. 

Bremer, B. 1992. Phylogeny of the Rubiaceae (Chiococceae) 

based on molecular and morphological data-Useful approaches 

for classification and comparative ecology. Ann. 

Missouri Bot. Gard. 79: 380-387. 

. 1996. Phylogenetic studies within Rubiaceae and 

relationships to other families based on molecular data. 

Opera Bot. Belg. 7: 33-50. 

& O. Eriksson. 1992. Evolution of fruit characters 

and dispersal modes in the tropical family Rubiaceae. 

Biol. J. Linn. Soc. 47: 79-95. 

& R. K. Jansen. 1991. Comparative restriction site 

mapping of chloroplast DNA implies new phylogenetic 

relationships within the Rubiaceae. Amer. J. Bot. 78: 

198-213. 

& L. Struwe. 1992. Phylogeny of the Rubiaceae 

and the Loganiaceae: Congruence or conflict between 

morphological and molecular data? Amer. J. Bot. 79: 

1171-1184. 

, K. Andreasen & D. Olsson. 1995. Subfamilial and 

tribal relationships in the Rubiaceae based on rbcL sequence 

data. Ann. Missouri Bot. Gard. 82: 383-397. 

Brummitt R. K. & C. E. Powell (eds.). 1992. Authors of 

Plant Names. Royal Botanic Gardens, Kew. 

Burger, W. & C. M. Taylor. 1993. Rubiaceae. In: W. Burger 

(ed.), Flora Costaricensis. Fieldiana, Bot. n.s. 33: 1-333. 

Candolle, A. P. de. 1830. Rubiaceae. Prodromus Systematis 

Naturalis Regni Vegetabilis 4: 341-622. Treuttel & 

Wiirtz, Paris. 

Carrasco, L., A. Jimenez & D. Visquez. 1976. Specific 

inhibition of translocation by tubulosine in Eukaryotic 

polysomes. Europ. J. Biochem. 64: 1-5. 

Casaretto, G. 1843. Chrysoxylon. Novarum stirpium 

Brasiliensium decades. Decas 7: 59. 

Chamisso, L. A. & D. F. L. von Schlechtendal. 1829. De 

Plantis in Expeditionis speculatoria Romanzoffiana 

observatis. Linnaea 4: 129-202. 

Classen-Bockhoff, R. 1996. A survey on flower-like inflorescences 

in the Rubiaceae. Opera Bot. Belg. 7: 329-367. 

Condamine, C. M. de la. 1738. Sur l'arbre du quinquina. 

Mem. Acad. Roy. Sci. Paris 1738: 226-243, tabs. 5-6. 

--- . 1745. Relation abr6eg d'un voyage fait dans 

l'interieur de l'Amerique m6ridionale. Paris. 

Correll D. S. & D. H. Correll. 1982. Flora of the Bahama 

Archipelago. J. Cramer, Vaduz. 

Darwin, S. P. 1976a. The subfamilial, tribal and subtribal 

nomenclature of the Rubiaceae. Taxon 25: 595-610. 

. 1976b. The genus Lindenia (Rubiaceae). J. Arnold 

Arb. 57:426-449. 

Davis, P. H. & V. H. Heywood. 1963. Principles of angiosperm 

taxonomy. Van Nostrand, New York. 

Dean, W. 1995. With broadax and firebrand: The destruction 

of the Brazilian Atlantic forest. University of California 

Press, Berkeley. 

Delprete, P. G. 1995a. Proposed circumscription of the tribes 

Chiococceae, Condamineeae, and Catesbaeeae using 

morphological characters. Second International Rubiaceae 

Conference (13-15 September 1995, Meise, Belgium). 

Programme and abstracts. Scripta Bot. Belg. 11: 80. 

. 1995b. Three new species of Rustia from Panama 

and Ecuador. Novon 5: 133-139. 

. 1996a. Studies on pendulous-flowered species of 

Rustia (Rubiaceae, Condamineeae), including a new 

species, Rustia schunkeana. Brittonia 48: 29-34. 

- . 1996b. Notes on calycophyllous Rubiaceae, part I. 

Morphological comparison of the genera Chimarrhis, 

Bathysa, and Calycophyllum, with new combinations and 

anew species, Chimarrhis gentryana. Brittonia 48: 35-44. 

- . 1996c. Systematics, typification, distribution, and 

reproductive biology of Pinckneya bracteata (W. 

Bartram) Raf. (Rubiaceae). PI. Syst. Evol. 201: 243-261. 

- . 1996d. Evaluation of the tribes Chiococceae, Condamineeae, 

and Catesbaeeae (Rubiaceae) based on morphological 

characters. Opera Bot. Belg. 7: 165-192. 

. 1996e. Notes on the taxonomic position of the 

monotypic Brazilian genus Kerianthera (Rubiaceae). 


. 1996f. Revision of the tribe Condamineeae (sensu 

Hooker, 1873) (Rubiaceae). Ph.D. dissertation, University 

of Texas at Austin.

Literature Cited205 

. 1997a. Revision and typification of Brazilian Au- Gray, A. 1859. Notes upon some Rubiaceae, collected in 

gusta (Rubiaceae, Rondeletieae), with ecological obser- South Sea Exploring Expedition under Captain Wilkes. 

vation on riverine vegetation of Cerrado and Atlantic Proc. Amer. Acad. Arts 4: 306-324. 

Forest. Brittonia 49: 487-497. 

Grisebach, A. H. R. 1861. Rubiaceae. In: Flora of the Brit- 

- . 1997b. Notes on calycophyllous Rubiaceae. Part ish West Indian Islands 316-351. Lovell Reeve & Co., 

II. Morphological comparison of the genera Bathysa and London. 

Schizocalyx. Brittonia 49: 480-486. 

- . 1862 [1860-1862]. Plantae wrightianae, e Cuba 

. 1998a. A new species of Rustia (Rubiaceae) from orientali. Cambridge, Boston. 

Bilsa Biological Station, Esmeraldas Province, Ecuador. .1866. Phyllomelia. Catalogus plantarum cubensium 

Novon 8: 231-234. 

exibens collectionem wrightianam aliasque minores ex 

. 1998b. Notes on calycophyllous Rubiaceae. Part insula Cuba missas, 139-140. G. Engelmann, Lipsia. 

III. Systematic position of the monotypic Mexican ge- Hall6, F. 1961. Contribution a l'6tude biologique et taxononus 

Cosmocalyx and notes on the calycophyll develop- mique des Mussaendeae (Rubiacees) d'Afrique tropicale. 

ment. Brittonia 50: 309-317. 

Adansonia, ser. 2, 1: 266-298. 

- . 1998c. Notes on calycophyllous Rubiaceae. Part .1967. Etude biologique et morphologique de la 

IV. The monotypic Brazilian genus Blandibractea tribu des Gardeni6es (Rubiacees). Mem. O.R.S.T.O.M. 

Wernham is a Simira (Rubiaceae, Rondeletieae). 22: 1-146. 

Brittonia 50: 318-323. 

- , R. A. A. Oldeman & P. B. Tomlison. 1978. Tropi- 

. 1999a. Morphological and taxonomical compari- cal trees and forests. An architectural analysis. Springer, 

son of Ariadne, Mazaea, Phyllomelia, and Eosanthe New York. 

(Rubiaceae, Rondeletieae). Brittonia 51: in press. Hemsley, W. B. 1881. Rustia occidentalis. Biologia centrali- 

. 1999b. Rubiaceae-Condamineeae. In: G. Harling americana. Botany 2: 14. 

& L. Andersson (eds.), Flora of Ecuador (in press). Hesse, M. 1984. An exine architecture model for viscine 

& M. Nee. 1997. The enigmatic genus Wernhamia threads. Grana 23: 69-75. 

S. Moore is a synonym of Simira (Rubiaceae, Rondele- ----. 1986. Nature, form and function of pollen-connecttieae). 

Brittonia 49: 303-308. 

ing threads in angiosperms. Linn. Soc. Symp. Ser. 12: 

Ducke, A. 1922. Plantes Nouvelles ou peu communes de la 109-118. 

region amazonienne (II Partie). Arch. Jard. Bot. Rio Hiepko, P. 1987. The collections of the Botanical Museum 

Janeiro 3: 253-262. 

Berlin-Dahlem (B) and their history. Englera 7: 219-252. 

. 1925. Pseudochimarrhis. Plantes Nouvelles ou peu Hogeweg, P. & J. Koek-Noorman. 1975. Wood anatomicommunes 

de la region amazonienne (III Partie). Arch. cal classification using interactive character weighing. 

Jard. Bot. Rio Janeiro 4: 177. 

Acta Bot. Neerl. 24: 269-283. 

. 1935. Dendrosipanea. Plantes Nouvelles ou peu Holmgren, P. K., N. L. Holmgren & L. C. Barnett. 1990. 

communes de la region Amazonienne (VIII Serie). Arq. Index herbariorum. Part I: The herbaria of the world. 

Inst. Biol. Veg. 2: 69-71. 

Ed. 8. Regnum Veg. 120. International Association for 

. 1945. Chimarrhis. New forest trees and climbers Plant Taxonomy, New York Botanical Garden, Bronx. 

of the Brazilian Amazon. Bol. Techn. Inst. Agron. Norte, Hooker, J. D. 1868. Limnosipanea spruceana. Hook. Icon. 

Para 4: 26-29. 

PI. 11: 38. PI. 1050. 

Duke, J. A. & R. Vasques Martinez. 1994. Amazonian --- . 1871a. Phitopis multiflora. Hook. Icon. PI. 11: 75. 

ethnobotanical dictionary. PCR Press, Boca Raton, FL. PI. 1093. 

Dwyer, J. D. 1980. Family 179. Rubiaceae. In: Woodson --. 187 lb. Phyllacanthus grisebachianus. Hook. Icon. 

E. et al. (eds.), Flora of Panama, part IX. Ann. Missouri PI. 11: 76-77. PI. 1095. 

Bot. Gard. 67: 1-522. 

.1873. Ordo LXXXIV. Rubiaceae. In: G. Bentham 

Endlicher, S. 1838. Rubiaceae. Genera plantarum, part 7: & J. D. Hooker (eds.). Genera Plantarum ad exemplaria 

520-566. Fr. Beck, Wien. 

imprimis in herbariis kewensibus servata definita, 2: 7- 

Esau, K. 1965. Plant anatomy. Ed. 2. John Wiley, New York. 151. Lovell Reeve & Co., London. 

Estrella, E. 1991. La Naturaleza Americana y las Expedi- Howard, R. A. & G. R. Proctor. 1958. Acrosynanthus 

ci6nes Botinicas Espafiolas del Siglo XVIII. In: I. C. jamaicense. J. Arnold Arb. 39: 101-102. 

Cevallos (ed.), Flora Huayaquilensis, La Espedici6n Hutchinson, J. 1910. Pogonopus febrifugus. Kew. Bull. 

Botanicas de Juan Tafalla a la Real Audiencia de Quito, 1910:200-201. 

1799-1808. Abya-Yala Editing, Quito, Ecuador. ----. 1926. The families of flowering plants. I. Dicotyle- 

Fagerlind, F. 1937. Embryologische, zytologische und dons. Arranged according to a new system based on their 

bestaubungs-experimentelle Studien in der Familie Ru- probable phylogeny. Macmillan, London. 

biaceae nebst Bemerkungen fiber einige Polyploiditats- ~--- & J. M. Dalziel. 1931. Rubiaceae. Flora of West 

probleme. Acta Horti Berg. 11: 195-470. 

Tropical Africa 2(1): 65-137. Whitefriars Press, London. 

Garcia Kirkbride, M. C. 1979. Review of the neotropical Jacquin, N.J. 1763. Selectarum stirpium americanum historia, 

Isertieae (Rubiaceae). Brittonia 31: 313-332. 

in qua ad linnaeanum systema determinatae descripti- 

. 1981. Duas novas tribos de Rubiaceae neotropicais. aeque situntur plantae illiae, quas in insula Martinica, 

Revista Brasil. Bot. 4: 119-123. 

Jamaica, Domingo. Ex officina Krausiana, Wien. 

.1982. A preliminary Phylogeny for the neotropical - . 1797. Macrocnemum speciosus. Plantarum rariorum 

Rubiaceae. PI. Syst. Evol. 141: 115-121. 

horti caesarei schoenbrunnensis 1: 19. C. F. Wappler, Wien. 

Germano Filho, P. 1993. Estudo taxon6mico do genero Jansen, M. E. 1979. A revision of the genus Kajewskiella 

Bathysa (Rubiaceae, Rondeletieae) no Brasil. Master's the- (Rubiaceae). Blumea 25: 283-294. 

sis, Universidade Federal do Rio de Janeiro, Rio de Janeiro. Jussieu, A. L. de. 1789. Rubiaceae. Genera Plantarum se- 

Goudot, J. 1909. Rustia mosenii. Bull. Soc. Bot. France cundum ordines naturales disposita, juxta methodum in 

56, mem. 3: 337. 

horto regio parisiensis exaratum, 196-210. Herissant & 

- . 1828. Cinchona muzonensis. Phil. Mag. n.s. 3: 132. Barrois, Paris.


--- . 1807. Septieme memoire sur les caracteres generaux 

des families, tires des graines, et confirm6 ou rectifies 

par les observations de Gaertner. Ann. Mus. Natl. Hist. 

Nat. 10: 307-332. 

- . 1820. Sur la famille des plantes Rubiacees. Mem. 

Mus. Hist. Nat. 6: 365-410. 

Karsten, H. 1858-1869. Florae Columbiae terrarumque 

adiacentium specimina selecta in peregrinatione duodecim 

annorum observata delineavit et descripsit H. Karsten. 

2 volumes. Ferdinandi Duemmleri successores, Berlin. 

. 1887. Bentham-Hooker's "Genera Plantarum" un 

Florae Columbiae specimina selecta. Bot. Jahrb. Syst. 

8: 354-360. 

Kiehn, M. 1995. Chromosome survey of the Rubiaceae. 

Ann. Missouri Bot. Gard. 82: 398-408. 

Kirkbride, J. H. 1985a. A revision of the genus Capirona. 

Acta Amazon. 15: 47-60. 

Amat. Bonpland et Alex. de Humboldt. [sumptibus 

librariae graeco-latini-germanicae] Paris [1815-1825]. 

Lanjou, J. & F. A. Stafleu. 1954. Index herbariorum. Part 

II. Collectors, A-D. Regnum Veg. 2: 1-174. 

Leppik, E. E. 1956. The form and function of numeral patterns 

in flowers. Amer. J. Bot. 43: 445-455. 

Lersten, N. R. 1974a. Colleter morphology in Pavetta, 

Neorosea and Tricalysia (Rubiaceae) and its relationships 

to the bacterial leaf nodule symbiosis. Bot. J. Linn. 

Soc. 69: 125-136. 

-. 1974b. Morphology and distribution ofcolleters and 

crystalsin relation to taxonomy and bacterial leaf nodule 

symbiosis of Psychotria (Rubiaceae). Amer. J. Bot. 

61: 973-981. 

--- . 1975. Colleter types in Rubiaceae, especially in 

relation to the bacterial leaf nodule symbiosis. Bot. J. 

Linn. Soc. 71: 311-319. 

- . 1985b. Manipulus rubiacearum IV. Kerianthera 

(Rubiaceae), a new genus from Amazonian Brazil. 

Brittonia 37: 109-116. 

.- 1997a. Manipulus rubiacearum VII. BioLlania, 

Edici6n Esp. 6: 393-406. 

. 1997b. Manipulus rubiacearum VI. Brittonia 49: 

354-379. 

Klotzsch, J. F. 1846. Rustia. In: F. G. Hayne, Getreue 

Darstellung und Beschreibung der Arzneykunde 

gebrauchlichen Gewachse 14: tabs. 14, 15 [555]. 

- . 1853. Einige neue Gattungen der Rubiaceen. 

Monatsber. Koenigl. Preuss. Acad. Wiss. 1853: 494-502. 

Koek-Noorman, J. 1969a. Contribution to the wood 

anatomy of the South American Rubiaceae. I. Acta Bot. 

Neerl. 18: 108-123. 

---- . 1969b. Contribution to the wood anatomy of the 

South American Rubiaceae. II. Acta Bot. Neerl. 18: 

377-395. 

~--- . 1970. Contribution to the wood anatomy of the 

Cinchoneae, Coptosapelteae and Naucleeae (Rubiaceae). 

Acta Bot. Neerl. 19: 154-164. 

.1972. The wood anatomy of the Gardenieae, Ixoreae 

and Mussaendeae (Rubiaceae). Acta Bot. Neerl. 21: 

301-320. 

. 1976. Juvenile characters in the wood of certain 

Rubiaceae with special reference to Rubia fruticosa. 

IAWA-Bull. 3: 38-42. 

-- . 1977. Systematische Holtzanatomie einer Rubiaceen. 

Ber. Deutsch. Bot. Ges. 90: 183-190. 

---- . 1980. Wood anatomy and classification of 

Henriquezia Spruce, Platycarpum Hum. et Bonpl. and 

Gleasonia Standl. Acta Bot. Neerl. 29: 117-126. 

& P. Hogeweg. 1974. The wood anatomy of 

Vanguerieae, Cinchoneae, Condamineae and Rondeletieae 

(Rubiaceae). Acta Bot. Neerl. 23: 627-653. 

& C. Puff. 1983. The wood anatomy of Rubiaceae 

tribes Anthopermeae and Paederieae. PI. Syst. Evol. 147: 

17-45. 

Krause K. 1915. Rubiaceae. In: A. Engler & K. Prantl (eds.). 

Die Natiirlichen Pflanzenfamilien. Nachtrage [IV] zu den 

Teilen II-IV: 290-301. Engelmann, Leipzig. 

Krug, J. C. & I. Urban. Mazaea. 1897a. Ueber einigen 

Rubiaceen-Gattungen. Ber. Deutsche Bot. Ges. 15: 265- 

266, t. 9, fig. 10-21. 

& - . 1897b. Neomazaea. Ueber einigen 

Rubiaceen-Gattungen. Ber. Deutsche Bot. Ges. 15: 542. 

Kunth, C. S. 1818, 1820. Macrocnemum. In: F. H. A. 

Humboldt, A. J. de Bonpland & C. S. Kunth, Nova genera 

et species plantarum, quas in peregrinatione orbis 

novis collegerunt, descripserunt, partim adumbraverunt 

Linden, J. J. & J. E. Planchon. 1854. Pinckneya ionantha. 

Bot. Zeit. 365. 

Linnaeus, C. 1753. Species plantarum. Laurentii Salvii, 

Stockholm. 

Liogier, A. L. 1963. Rubiaceae. Flora de Cuba 5: 13-146. 

Industria Graficas Diario-Dia, Palencia de Castlla, Spain. 

. 1995. Rubiaceae. La Flora de la Espafiola. 7: 

207-439. 

Lorence, D. H. 1991. New species and combinations in 

Mexican and Central American Rondeletia (Rubiaceae). 

Novon 1: 135-157. 

- & J. D. Dwyer. 1988. A revision of Deppea (Rubiaceae). 

Allertonia 4: 389-436. 

Ma, W. W. 1991. Bioactive constituents of Pogonopus 

speciosus (Rubiaceae), Persea major (Lauraceae), and 

Endlicheria dysodantha (Lauraceae). Ph.D. dissertation. 

Dissertation Abstract International, Vol. 53/05-B. 

,J. E. Anderson, A. T. McKenzie, S. R. Byrn, J. 

L. McLaughlin, and M. S. Hudson. 1990. Tubulosine: 

An antitumor constituent ofPogonopus speciosus. J. Nat. 

Prod. (Lloydia) 53: 1009-1014. 

Maddison W. P. & D. R. Maddison. 1992. MacClade: 

Analysis ofphylogeny and character evolution. Version 

3.0. Sinauer Associates, Sunderland, Massachusetts. 

Marques-Souza, A. C., M. L. Absy, I. P. de A. Miranda 

& H. E. C. Kfichmeister. 1993. Caracteristica de flores, 

nectar y visitantes de Kerianthera preclara (Rubiaceae). 

Rev. Biol. Trop. 41: 483-489. 

Martius, C. F. P. de. 1837 [-1841]. Herbarium Florae 

Brasiliensi. Plantae brasilienses exsiccatae, quas 

denominatas, partim diagnosi aut observationibus 

instructas botanophilis offert Dr. C. Fr. Ph. de Martius. 

[Publisher unknown] Miinchen. 

McDowell, T. 1996. Exostema (Rubiaceae): Taxonomic 

history, nomenclature, position and subgeneric classification. 


Merrill E. D. & L. M. Perry. 1947. Kajewskiella, a new 

rubiaceous genus from the Solomon Islands. J. Arnold 

Arb. 28: 331-332. 

Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the dicotyledons. 

Clarendon Press, Oxford. 

Michaux, F. A. 1817. Pinckneya. The North American sylva 

1: 180-181, pl. 49. Thomas Dobson, Philadelphia. 

Miquel, F. A. W. 1849. Coffeafloribunda. Linnaea 22: 804. 

Moore, S. 1922. Wernhamia. In: Herzog, T., Die von Dr. 

Th. Herzog auf seiner zweiten Reise durch Bolivien in 

den Jahren 1910 und 1911 gesammelten Pflanzen. 

Medel. Rijks. Herb. Leiden 46: 22-23. 

Muller, J. 1979. Form and function in angiosperm pollen 

grains. Ann. Missouri Bot. Gard. 66: 569-632.

Literature Cited 207 

Orsted, A. S. 1852. Macrocnemum exsertus. Vidensk. Robbrecht, E. 1988. Tropical woody Rubiaceae. Charac- 

Meddel. Naturhist. Foren. Kjobenhavn 2-4: 45. (reimpr. teristic features and progressions. Contributions to a new 

in Centralamerikas Rubiaceer. K0benhavn. 1853). subfamilial classification. Opera Bot. Belg. 1: 1-271. 

. 1863. Pogonopus exsertus. L'Amerique centrale 17, - . 1993 [1994]. Supplement to the 1988 outline of the 

Tab. 13. K0benhavn. 

classification of the Rubiaceae. Index to genera. In: E. 

Pacini, E. 1986. An approach to armomegathy. In: M. Cresti Robbrecht (ed.), Advances in Rubiaceae macro- 

& R. Dallai (eds.), Biology of reproduction and cell systematics. Opera Bot. Belg. 6: 173-196. 

motility in plants and animals. Siena University, Siena. & D. M. Bridson. 1993 [1994]. Nomenclatural notes 

. 1990. Harmomegathic characters of Pteridophyta on three Rubiaceeae genera. In: E. Robbrecht (ed.), 

spores and Spermatophyta pollen. PI. Syst. Evol. Suppl. Advances in Rubiaceae macrosystematics. Opera Bot. 

5: 53-69. 

Belg. 6: 197-200. 

Parrotta, J., J. K. Francis & R. R. de Almeida. 1995. & C. Puff. 1986. A survey of the Gardenieae and 

Trees of the Tapaj6s-A photographic field guide. Gen. related tribes (Rubiaceae). Bot. Jahrb. Syst. 108: 63-138. 

Tech. Rep. IITF - 1. U.S.D.A., Forest Service. International ---, --- & E. Smets. 1996. Second International 

Institute of Tropical Forestry, Rio Piedras, Porto Rico. Rubiaceae Conference. Proceedings. Opera Bot. Belg. 

Peixoto, A.L. 1981. Novas especies de Simira Aublet (Ru- 7: 1-432. 

biaceae) do norte do Espirito Santo. Bol. Mus. Bot. Rogers, G. K. 1987. The genera of the Cinchonoideae 

Munic. 44: 1-7. 

(Rubiaceae) in the southeastern United States. J. Arnold 

. 1982. As especies de Simira (Rubiaceae, Rondele- Arb. 68: 137-183. 

tieae) do Norte do Estado do Espirito Santo. Arq. Univ. Roig y Mesa, J. T. 1928. Diccionario botanico de nombres 

Fed. Rural Rio de Janeiro 5: 115-128. 

vulgares cubanos. [Boletin Estaci6n Experimental 

Planchon, J. E. 1849. Rogiera amoena. Fl. Serres Jard. Eur. Agron6mica, Santiago de las Vegas 54] Rambla, Bouza 

5: t. 442 (a-c). 

y Ca., La Habana. 

Pohl, J. E. 1828-1829. Augusta. Plantarum Brasiliae icones Rova, J. H. E., L. Andersson, P. G. Delprete & V. A. 

et descriptiones hactenus ineditae 2: 1-8, tabs. 101-105. Albert. 1997. Macrophylogeny of the Rubiaceae from 

[Publisher unknown] Wien. 

trnL-F sequence data. Abstract of the AIBS meetings, 

Poiret, J. L. M. 1811. Chimarrhis. In: J. B. A. P. M. de Canada. Amer. J. Bot. 84 (supplement): 227. 

Lamark, Encyclopedie methodique. Botanique. Suppl. Ruiz L6pez, H. 1778. Descripcion del Arbol de la Quina, 

2: 231. H. Agasse, Paris. 

hecha en el 1737 por Mr. de La Condamine de la Real 

Poiteau, P. A. 1804. Stevensia, nouveau genre de la famille Academia de Sciencias de Paris, entre cuyas memorias 

des Rubiacees. Ann. Mus. Natl. Hist. Nat. 4: 235-236, pertenecientes al afio 1738 se halla en la pagina 226. 

pi. 60. 

Traducida del Frances al Castellano, y aumentada con 

Prance, G. T. 1976. Inventario forestal de um hectare de algunas notas relativas a los arboles de Quina, nuevamata 

de terra firme km 30 de Estrada Manaus- mente descubiertos en Santa Fe de Bogota, Capital del 

Itacoatiara. Acta Amazon. 6: 9-35. 

Nuevo Reyno de Granada en la America Meridional. 

Presi, C. B. 1845. Bathysa. Abh. Konigl. B6hm. Ges. Wiss. [Publisher unknown] Madrid. 

3: 514- 515. (reimpr. Bot. Bemerk. 84. 1844 [1845]). 

& J. A. Pav6n y Jiminez. 1799. Rubiaceae. Flora 

Puff, C., A. Igersheim & U. Rohrhofer. 1993. peruviana et chilensis, sive descriptiones et icones plantarum 

Pseudomussaenda and Schizomussaenda (Rubiaceae): peruvianarum et chilensium, secundum systema linneanum 

Close allies of Mussaenda. Bull. Nat. Plantentuin. Belg. digestae, cum characteribus plurimum generum evulga- 

62: 35-68. 

torum reformatis. Vol. 2. Gabrielis de Sancha, Madrid. 

Radford, A. E., W. C. Dickison, J. R. Massey & C. R. Rusby, H. H. 1893. Malanea grandis. On the collections of 

Bell. 1974. Vascular plant systematics. Harper & Row, Mr. Miguel Bang in Bolivia. Mem. Torrey Bot. Club 3: 

New York. 

43-44. 

Raffauf, R. B. 1970. A handbook of alkaloids and alkaloid- . 1895. Elaeagia grandis. On the collections of Mr. 

containing plants. John Wiley, New York. 

Miguel Bang in Bolivia-Part II. Mem. Torrey Bot. Club 

Rafinesque, C. S. 1820. Tableau analytique des ordres 4: 208. 

naturels, families naturelles et genres, de la classe . 1896. Condaminea. On the collections of Mr. 

Endogynie, sous classe Corisantherie. Ann. Gen. Sci. Miguel Bang in Bolivia-Part III. Mem. Torrey Bot. 

Phys. 6: 76-89. 

Club 6: 45-46. 

Record, S. J. & R. W. Hess. 1943. Timbers of the New - . 1927. Flexanthera. Mulford exploration of the 

World. Yale University Press, New Haven. 

Amazon Valley. Mem. New York Bot. Gard. 7: 367-370. 

Reinders, E. 1935. Fiber-tracheids, libriform wood fi- Saldanha da Gama, J. de. 1872. Configuraqao e Estudio 

bers and systematics in wood anatomy. Trop. Woods Botanico dos Vegetaes Seculares da Provincia do Rio 

44: 30-36. 

de Janeiro e de Outros Pontos do Brasil. Laemmert, Rio 

Richard, A. 1830. Memoire sur la famille des Rubiacees. de Janeiro. 

Imprimerie de J. Tatsu. [First copies apparently distrib- Sandwith, N. Y. 1949. Contributions to the flora of tropical 

uted in July 1829]. Paris. (Reimpr. Mem. Soc. Hist. Nat. America: XLIX. Notes on Rubiaceae. Kew Bull. 1949: 

Paris, ser. 3, 5: 81-224. 1834). 

253-264. 

Richard, L. C. M. 1803. Pinckneya. In: A. Michaux, Flora Schumann, K. 1888a. Rubiaceae, trib. VII-IX. In: C. F. P. 

boreali-americana, vol. 1: 103-105, t. 13. Fratres von Martius, A. G. Eichler & I. Urban (eds.), Flora 

Levrault, Paris. 

brasiliensis enumeratio plantarum in Brasilia hactenus 

Ridsdale, C. E. 1982. A revision of Badusa (Rubiaceae, detectarum... 6(6): 1-124. Fleischer, Leipzig. 

Condamineeae, Portlandiinae). Blumea 28: 145-150. . 1888b. Uber einige verkannte oder wening gekannte 

Rizzini, C. T. 1947. Chimarrhis. Novitates rubiacearum Geschlechter der Rubiaceen Sfidamerikas. Bot. Jahrb. 

compositarumque. Rev. Brasil. Biol. 7: 275-277. 

Syst. 10: 302-363.


. 1889. Rubiaceae, trib. X-XIX. In: C. F. P. von 

Martius, A. G. Eichler & I. Urban (eds.), Flora brasili- 

ensis enumeratio plantarum in Brasilia hactenus 

detectarum... 6(6): 125-466. Fleischer, Leipzig. 

. 1891. Rubiaceae. In: A. Engler & K. Prantl (eds.). 

Die Natirlichen Pflanzenfamilien 4(4): 1-156. Engel- 

mann, Leipzig. 

& K. Krause. 1908a Chimarrhis. In: K. Krause, 

Rubiaceae andinae. Engl. Bot. Jahrb. 40: 312-313. 

- & -- . 1908b. Acrobotrys. In: I. Urban, Plantae 

novae andinae imprimis Weberbauerianae. III. Bot. 

Jahrb. 40: 316-318. 

& - . 1908c. Dolichodelphys. In: E. Ule, Verh. 

Bot. Vereins. Prov. Brandenburg 50: 102-104. 

Simpson, D. R. 1976. Studies in neotropical Rubiaceae. I. 

Rustia. Phytologia 33: 4-8. 

Skvarla, J. J., P. H. Raven, W. F. Chissoe & M. Sharp. 

1978. An ultrastructural study of viscin threads in 

Onagraceae pollen. Pollen & Spores 20: 5-143. 

Soejarto, D. D., P. G. Delprete, J. C. Regalado & D. A. 

Madulid. 1996. The true provenance and identity of 

Badusa philippica (Rubiaceae). 

Solereder, H. 1890. Rustia pauciflora. Studien uber die 

Tribus der Gaertnereen Bentham et Hooker. Ber. 

Deutsch. Bot. Ges. 8: 99-100. 

. 1893. Ein Beitrag zur anatomischen Charakteristik 

und zur Systematik der Rubiaceen. Bull. Herb. Boissier 

1: 167-183. 

. 1908. Systematic anatomy of the dicotyledons. 

Clarendon Press, Oxford. 

Spjut, R. W. 1994. A systematic treatment of fruit types. 

Mem. New York Bot. Gard. 70: 1-182. 

Standley, P. C. 1916. Rustiaferruginea. Systematic inves- 

tigations of tropical American plants. Contr. U.S. Natl. 

Herb. 18: 142. 

- . 1918, 1921, 1934a, 1934b. Rubiaceae. N. Amer. 

Fl. 32: 1-300. 

. 1926. Chimarrhis microcarpa. In: N. L. Britton, 

West Indian plants. Bull. Torrey. Bot. Club 53: 471. 

. 1929. Condaminea breviflora. Studies on American 

plants-I. Publ. Field Columbian Mus., Bot. Ser. 4: 264. 

. 1930a. The Rubiaceae of Colombia. Publ. Field 

Columbian Mus., Bot. Ser. 7(1): 1-176. 

. 1930b. Studies on American plants-IV, Publ. Field 

Columbian Mus., Bot. Ser. 8: 131-236. 

. 1931a. The Rubiaceae of Ecuador. Publ. Field 


. 1931b. The Rubiaceae of Bolivia. Publ. Field 


. 1931c. The Rubiaceae of Venezuela. Publ. Field 

Columbian Mus., Bot. Ser. 7: 341-485. 

-- . 193 Id. Phitopis sterculioides. Studies on American 

plants-V. Publ. Field Columbian Mus., Bot. Ser. 8: 341. 

-- . 1932. Holtonia, a new genus of trees of the family 

Rubiaceae. Trop. Woods 30: 37-38. 

.1936. Rubiaceae. In: J. F. Macbride, Flora of Peru. 

Field Mus. Nat. Hist., Bot. Ser. 13: 1-263. 

- . 1940. Rubiaceae. Studies on American plants. Publ. 

Field Mus. Nat. Hist., Bot. Ser. 22: 48-58, 107-125, 

171-218, 278-294, 382-390. 

. 1947. Stomandra. Studies on Central American 

plants-VII. Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 

247-248. 

& J. A. Steyermark. 1953. Chimarrhis 

venezuelensis. Botanical exploration in Venezuela-III. 

Fieldiana, Bot. 28: 571-573. 

& L. O. Williams. 1975. Rubiaceae. In: Flora of 

Guatemala. Fieldiana, Bot. 24: 1-274. 

Stellfeld, C. 1948. Os novos generos e as novas species de 

Freire AllemAo. Pongetti, Rio de Janeiro. 

Stearn, W. T. 1957. Achille Richard's "M6moire sur la 

famille des Rubiac6es." Taxon 6: 186-188. 

Steyermark, J. A. 1963. Dioicodendron, a new genus of 

Rubiaceae. Bol. Soc. Venez. Ci. Nat. 25: 23-28. 

. 1964a. A new Tresanthera from Venezuela. Bol. 

Soc. Venez. Ci. Nat. 25: 248-249. 

. 1964b. Cuatrecasasiodendron. Novedades en las 

Rubiaceas Colombianas de Cuatrecasas. Acta Biol. 

Venez. 4: 29-35. 

-. 1964c. Rubiaceae. In: B. Maguire, J. J. Wurdak & 

Collaborators, Botany of the Guayana Highlands, Part 

V. Mem. New York Bot. Gard. 10: 186-278. 

.1965. Rubiaceae. In: B. Maguire, J. J. Wurdak & 


VI. Mem. New York Bot. Gard. 12: 178-285. 

---. 1967. Rubiaceae. In: B. Maguire, J. J. Wurdak & 


VII. Mem. New York Bot. Gard. 17: 230-436. 

. 1971. Pogonopus speciosus. Rubiaceas nuevas de 

los Andes y Cordilleras de la costa de Venezuela. Acta 

Bot. Venez. 6: 108-110. 

. 1972. Rubiaceae. In: B. Maguire, J. J. Wurdak & 


IX. Mem. New York Bot. Gard. 23: 227-832. 

-- . 1974. Rubiaceae. In: T. Lasser & J. A. Steyermark, 

Flora de Venezuela 9: 1-2070. Instituto BotAnico, Caracas. 

--- . 1975. Revision of the genus Joosia (Rubiaceae). 

Brittonia 27: 251-262. 

---- . 1986. Holstianthus, a new genus of Rubiaceae from 

the Guyana Highland. Ann. Misssouri Bot. Gard. 73: 495. 

Swofford, D. L. 1993. PAUP: Phylogenetic analysis using 

parsimony. Version 3.1.1. Computer program distributed 

by the Illinois Natural History Survey, Champaign, Illinois. 

Tange, C. 1995. The identity of Siderobombyx and a new 

species ofXanthophytum (Rubiaceae). Nordic J. Bot. 15: 

575-581. 

Taylor, C. M. 1995. First International Conference on the 

Rubiaceae: Introduction. Ann. Missouri Bot. Gard. 82: 

355-356. 

& B. E. Hammel. 1993. Notes on Elaeagia 

myriantha, Comb. Nov. (Rubiaceae). Novon 3:490-491. 

Troll, W. 1950. Uber den Infloreszenzbegriff und seine 

Anwendung aufdie krautige Region bliihender Pflanzen. 

Abh. Akad. Wiss. Lit. Mainz, Mat.-Nat. KI. Nr. 13. 

Tryon, A. F. 1963. Hermann Karsten, his collections and 

the Flora columbiae. Taxon 12: 103-105. 

Turczaninow, N. S. 1848. Molopanthera. Decades Quarta 

et Quinta, generum adhuc non descriptorum. Bull. Soc. 

imp. Natural. Moscou 21: 580-581. 

Urban, I. 1899, 1903,1912,1913,1923. In: I. Urban(ed.), 

Symbolae antillanae seu fundamenta florae Indiae 

occidentalis. 9 Volumes, 1898-1928. Berlin, Leipzig, 

Paris, London. 

. 1922. Ariadne. Sertum antillanum XIV. Feddes 

Repert. Spec. Nov. Regni Veg. 18: 25-26. 

- . 1931. Rustia haitiensis. Plantae haitienses novae 

vel rariores a cl. Ekman 1924-1930 lectae. Stokholm. 

Almqvist & Wiksells Botryckeri. [reimpr. Ark. Bot. 24A 

(4): 36]. 

Verdcourt, B. 1958. Remarks on the classification of the 

Rubiaceae. Bull. Jard. Bot. Etat. 28: 209-281.

Numerical List of Taxa209 

Waha, M. 1984. Zur Ultrastruktur un Funktion pollenverbindender 

Faden bei Ericaceae und anderen Angiospermenfamilien. 

PI. Syst. Evol. 147: 189-203. 

Walpers, W. G. 1852. Henlea. Annales botanices 

systematicae 2: 788. F. Hofmeister, Leipzig. 

Weberling, F. 1977. Beitrige zur Morphologie der Rubiaceen- 

Infloreszenzen. Ber. Deutsche Bot. Ges. 90: 191-209. 

. 1992. Morphology of flowers and inflorescences. 

Cambridge Univ. Press. 

Weddell, H. A. 1849. Histoire Naturelle des Quinquinas, 

ou monographie du genre Cinchona suivie d'une description 

du genre Cascarilla et de quelques autres plantes 

de la meme tribu. Victor Masson, Paris. 

- . 1854. Howardia. Notice sur quelques Rubiacees 

de l'Amerique tropicale. Ann. Sci. Nat. 4: 66-71, pl. 10. 

Wernham, H. F. 1911. Pteridocalyx. A new genus of Rubiaceae. 

J. Bot. 49: 317-318. 

NUMERICAL LIST OF TAXA 

. 1912. Carmenocania. New Rubiaceae from tropical 

America-I. J. Bot. 50: 241-242. Tab. 520. 

- . 1913. Pteridocalyx minor. New Rubiaceae from 

tropical America-II. J. Bot. 51: 218. 

. 1916. Pseudomussaenda, a new genus of Rubiaceae. 

J. Bot. 54: 297-301. 

- . 1917. Blandibractea, Neobertiera. Tropical American 

Rubiaceae-VIII. J. Bot. 55: 170-171. 

Willaman, J. J. & B. G. Schubert. 1961. Alkaloid-bearing 

plants and their contained alkaloids. Techn. Bull. 

U.S.D.A. 1234: 1-287. 

Willdenow, C. L. von. 1801. Sickingia. Zhen Neue 

Gattungen von Gewichsen. Ges. Naturf. Freunde Berlin 

Neue Schriften 3: 445-448. 

Wodehouse, R. P. 1935. Pollen grains. McGraw-Hill, New 

York. 

1-1. Rustia costaricensis (Standley) Lorence 

5-3. P. tubulosus (A. Richard in A. P. de Candolle) K. 

1-2. R. occidentalis (Bentham) Hemsley 

Schumann 

1-3. R. dressleri Delprete 

1-4. R. alba Delprete 

6-1. Chimarrhis parviflora Standley 

1-5. R. bilsana Delprete 

6-2. C. latifolia Standley 

1-6. R. rubra Standley ex D. Simpson 

6-3. C. cubensis Steyermark 

1-7. R. viridiflora Delprete 

6-4. C. jamaicensis (Urban) Steyermark 

1-8. R. schunkeana Delprete 

6-5. C. cymosa Jacquin 

1-9. R. thibaudioides (Karsten) Delprete 

6-6. C. glabriflora Ducke. 

1-10. R. venezuelensis Standley & Steyermark 

6-7. C. hookeri K. Schumann 

1-11. R. formosa Klotzsch 

6-8. C. microcarpa Standley 

1-12. R. 6-9. C. 

simpsonii Delprete 

speciosa (Steyermark) Delprete 

1-13. R. angustifolia K. Schumann 

6-10. C. turbinata A. P. de Candolle 

1-14. R. gracilis K. Schumann 

6-11. C. barbata (Ducke) Bremekamp 

6-12. C. duckeana Delprete 

2-1. Tresanthera condamineoides Karsten 

6-13. C. brevipes Steyermark 

2-la. T. condamineoides var. condamineoides 

6-14. C. gentryana Delprete 

2-lb. T. condamineoides var. thyrsiflora (Steyermark) Delprete 7-1. Dioicodendron dioicum (K. Schumann & K. Krause) 

3-1. Condaminea corymbosa (Ruiz & Pav6n) A. P. de Candolle 

Steyermark 

3-2. C. elegans Delprete 

8-1. Molopanthera paniculata Turczaninow 

8-la. M. paniculata var. paniculata 

4-1. Picardaea cubensis (Grisebach) Britton ex Urban 8-lb. M. paniculata var. burchellii (J. D. Hooker) K. 

Schumann 

5-1. Pogonopus exsertus (Orsted) Orsted 

5-2. P. speciosus (Jacquin) K. Schumann 

9-1. Dolichodelphys chlorocrater K. Schumann & K. Krause 

5-2a. P. speciosus var. speciosus 

5-2b. P. speciosus var. sandwithianus (Steyermark) Steyermark 10-1. Parachimarrhis breviloba Ducke


Abbiatti, D. & L. Claps, 288 (5-3). 

Acevedo-Rodriguez, P. et al., 6774, 6900 (1-2). 

Acosta-Solis, M., 12635 (3-1). 

Acufia, J., 8917, s.n. (6-3). 

Adair, s.n. (6-12). 

Adams, C. D., 9289 (6-4). 

Aguilera, V., 17 (5-1). 

Alain, Bro. & J. Acufia, 7673 (4-1). 

Albert de Escobar, L., 1373 (3-1). 

Alexander, R. C., s.n. (6-4). 

Allard, H. A., 22129 (3-1). 

Allart, A., 236 (5-2a). 

Allen, C., 622 (5-2b). 

Allen, P. H., 860 (1-2); 2017 (5-1); 3217 (1-9); 5613 (6-2); 

5621 (5-1); 5762 (6-2); 5910 (3-1); 6302 (6-2). 

Almeda, F. et al., 4264 (3-1). 

Alvarado, A., 233 (1-6); 292 (6-7). 

Alverson, W. S. et al., 28 (10-1); 214 (5-1). 

Ancuash, E., 504 (6-7); 604 (3-1); 1256, 1372 (6-6); 1509 

(3-1). 

Anderson, s.n. (6-5). 

Anderson, W. R., 9927 (1-11). 

Andersson, L. et al., 2108 (7-1). 

Andre, E., 1955 (3-1). 

Antonio, T. M., 1935 (1-2). 

Antonio, T. M. & W. J. Hahn, 4240 (1-2). 

Apparicio, P. A., 919, 59161 (1-11). 

Argeli, C. & C. A. de Oliveira, 691 (1-11). 

Aristeguieta, L., 5414, 6635 (5-2a). 

Asplund, E., 7844, 8571, 12428, 18347 (3-1); 18686 (1-6). 

Assuncao, 369 (6-11). 

Aymard, G. et al., 5233 (7-1). 

Badcock, W. J., 107, 739 (5-3). 

Badillo, V. M., 335 (5-2a). 

BAFOG Service Forestiere, 166M, 1187, 1262, 7608 (6-10). 

Bailey, L. H. & E. Z. Bailey, 272 (6-5); 865 (5-2a). 

Baker, M. A. & D. Neil, 5556 (3-1). 

Baker, M. A. et al., 6000 (6-7). 

Balee, W. L. & B. G. Ribeiro, 431, 2826 (6-10). 

Bang, M., 610 (3-1); 1344(5-3); 1429(3-1); 1716, s.n. (5-3). 

Barclay, A. S. et al., 3424 (7-1); 3463 (3-1). 

Barfod, A., 41389 (3-1). 

Barreto, M., 1789 (JBBH16686), 8305 (JBBH26371) (8-1b). 

Barrett, G. E., s.n. (5-2a). 

Barrier, S., 4993, 5060 (6-10). 

Barroso, G. et al., 66 (RB 179015) (5-3). 

Bartlett, H. H., 20361, 20389 (5-3). 

Bautista, H. P. & R. P. Orlandi, 985 (8-la). 

Bayley, E., s.n. (5-2a). 

Beard, P., 1134 (6-5). 

Beck, H. T. et al., 1055 (6-14). 

Belanger, 195, 196 (6-5). 

Belem, P., 3677 (8-la). 

Belshaw, C. M., 3422 (3-1); 3514 (1-8); 3593 (9-1). 

Bena, P., 1066 (6-5); 1187, 1262 (6-10). 

Benavides, O., 5651 (3-1). 

Benitez de R., C. E., 1083 (5-2a); 1513 (3-1). 

Bennett, B. C. & P. G6mez Andrade, 3638 (6-6). 

Bennett, B. C. et al., 4028 (6-6). 

Benoist, R., 1190 (6-10). 

Bensman, R., 331 (6-6). 

Berg, 1227 (1-7). 

LIST OF EXSICCATAE 

Berlin, B., 1812(3-1). 

Berardi, A. L., 563, 2209 (3-1); 5829 (2-lb); 5943 (6-9). 

Berry, P., 1847 (5-2a). 

Besse, L. et al., 602 (5-3). 

Billiet, F. & B. Jadin, 5010 (6-5). 

Bisby, F. et al., P18095 (6-10). 

Black, G. A., F8-47-955 (6-10). 

Black, G. A. et al. F8-50-9831 (6-10). 

Blanchet, J., 2557, 3282, s.n. (8-la). 

Blanco, C., 822 (6-8). 

Blum, K. E., 1260 (5-1). 

Boeke, J. D., 1501 (3-1). 

Boeke, J. D. & S. Utzschneider, 2925 (5-1). 

Bonpland, M. A., 622 (3-1). 

Boom, B., 10698 (6-8). 

Boom, B. & S. Mori, 2037, 2041, 2043, 2050, 2067, 2069, 

2078, 2080, 2180, 2213, 2217, 2226, 2230, 2263, 2445 

(6-8). 

Boom, B. & D. Beardsley, 8443 (3-1). 

Boom, B. et al., 8756 (6-10). 

Borsini, O., s.n. (5-3). 

Brade, A. C., 10639 (1-11). 

Brandbyge, J. & E. Asanza C., 30431 (3-1); 31615 (6-7). 

Bravo & G6mez, 306 (6-7). 

Bredemeyer, F., s.n. (5-2a). 

Breedlove, D. E., 22535, 28572 (5-1). 

Breedlove, D. E. & R. F. Thore, 30715 (5-1). 

Breteler, F. J., 3612 (3-1). 

Bridges, s.n. (5-3). 

Bristan, N., 1330, 1346, 1425, 1519 (5-1). 

Britton, N. L., 610 (6-4). 

Broadway, W. E., 3076 (2-la); 6464, 10011 (6-8); s.n. (2-la). 

Bruijn, J. de, 1646, 1647, 1651 (6-8). 

Buchtien, 0., 761 (5-3); 1393 (3-1); 1397 (5-3); 8186, s.n. 

(3-1). 

Burchell, W. J., 3181 (1-11). 

Burger, W. & G. Matta, 4593 (3-1). 

Burger, W. et al., 12314 (1-1). 

Burkart, A., 17038 (5-2a). 

Busey, P., 464 (1-1). 

Cabrera, A. L., 4159, 14523 (5-3). 

Cabrera, A. L. et al., 22348, 30389 (5-3). 

Cabrera R., I., 3264 (1-2); 3701 (1-9). 

Calder6n, S., 1225 (5-1). 

Callejas, R. & O. Escobar, 3234 (3-1). 

Callejas, R. et al., 2512 (3-1); 4160(10-1); 5038 (1-2); 5824 

(3-1). 

Camp, W. H., A-C25, E-25, E-1016, E-1540, E-2384 (3-1). 

Campos, M. D. et al., 48 (1-11). 

Campos, M. T. V. do A. et al., 468 (INPA 185019) (6-11). 

Cardenas, D., 2593, 4261, 5895 (5-3). 

Cavalcante, P. & M. Silva, 1598 (6-10). 

Cazalet, P. C. D. & T. D. Pennington, 7789 (6-6). 

Castillo, A., 1966 (5-2a). 

Ceballos, A. et al., 36 (5-3). 

Cer6n, C., 1600 (3-1). 

Cer6n, C. E., 3661 (6-7). 

Cer6n, C. E. & J. Ayala, 9656 (6-7). 

Cer6n, C. E. et al., 4353 (3-1); 8458, 8462 (6-7). 

Chagas, A. S. L. et al. 2217 (6-10). 

Chagas, J., 486 (MG25131) (6-12). 

Chavez, F., 265 (6-7).

List of Exsiccatae211 

Chavez-Alfaro, R., 359 (5-3). 

Churchill, H. W. & G. de Nevers, 4446 (5-1). 

Cid, C. A. et al., 4896, 4939, 5022 (5-3). 

Claes, F., s.n. (1-9). 

Clark, J. L., 2979 (1-5). 

Clark, J. L. et al., 139 (1-5). 

Claussen, P., 669, s.n. (1-11). 

Clement, Bro. & P. Roca, 9941 (6-3). 

Constance, L. & O. Tovar, 2272 (3-1). 

Cook, O. F. & G. B. Gilbert, 913 (3-1); 1389, 1564 (5-3). 

Cooper, G. P., 38 (6-5). 

Cooper, G. P. & G. M. Slater, 3, 120 (6-1); 260 (6-2); 401 

(6-1). 

Cordeiro, 676 (7-13). 

Core, E. L. 566, 1307 (3-1); 1388 (7-1); 1416 (3-1). 

Cornejo, X., 582 (6-6). 

Correa, M. N. et al., 4577 (5-3). 

Cosson, E., 18 (6-5). 

Cowan, R. S., 1439 (2-la). 

Croat, T. B., 8959 (5-1); 9828 (1-2); 11244 (6-1); 12400, 

12465, 12511, 12652, 12895 (5-1); 21099 (3-1); 22191 

(1-1); 26547 (3-1); 27508, 27551, 27709 (1-1); 27755, 

36431 (6-1); 57632, 69930 (3-1). 

Cruz, R., WB-00482 (5-1). 

Cuadros V., H., 950 (1-2); 1243 (3-1); 1677 (5-2b). 

Cuadros V. J. H. et al., 3884 (5-1). 

Cuatrecasas, J., 2325 (3-1); 7370, 7819 (1-9); 8665 (7-1); 

8926 (9-1); 8947 (6-6); 11059 (3-1); 11076 (1-9); 13074 

(3-1); 14317 (1-2); 15192 (7-1); 15905, 17580, 19736 

(1-2); 22458, 22516 (7-1). 

Cuatrecasas, J. et al., 28672 (7-1). 

Cuezzo, A. R., 10.262 (5-3). 

Daly, D. C. et al., 4312 (6-11); 6496 (3-1). 

Daniel, Bro., 314 (3-1); 2787 (5-1); 5533 (1-9). 

Dannouse, I., 6946 (6-8). 

D'Arcy, W. & K. Sytsma, 14523 (5-1); 14584 (3-1). 

Davidse, G., 4174 (5-2a). 

Davidse, G. & G. Herrera C., 26263 (3-1). 

Davidson, C., 4805 (3-1). 

Davidson, M. E., 792 (3-1). 

Davis, E. W., 184 (1-9). 

Dayton, W. A. & W. R. Barbour, 3007 (6-1). 

Duarte, A. P. et al., 60854 (1-11). 

Delprete, P., 6196 (6-6); 6319,6329,6330 (1-2), 6526 (6-7), 

6527, 6528 (6-6). 

Delprete, P. & P. Apreza, 6359 (5-2b). 

Delprete, P. & G. Fagua, 6378, 6381, 6382 (1-9). 

Delprete, P. & L. Gilbert, 5267 (1-2); 5271 (6-2). 

Delprete, P. & J. Luteyn, 6205, 6207 (3-1). 

Delprete, P. & A. Verduga, 6412 (3-1); 6414, 6419, 6420 

(1-4); 6421, 6422 (6-7); 6423, 6424 (6-6); 6436 (3-1). 

Delprete, P. & G. L. Webster, 6104 (3-1). 

Delprete, P. et al., 6049, 6218 (3-1). 

Del Risco, E., 27343, 27395, 27458 (6-3). 

De Marco, N. E. et al., 10262 (5-3). 

Denslow, J., 2680 (9-1). 

De Paula, 3233 (1-11). 

Devia, W., 316, 399, 558, 559, 559A (3-1). 

Dezzeo, N., 73 (6-13). 

Diaz, V., 32 (5-1). 

Diaz Pedrahita, S., 3446, 3817 (7-1). 

Dick, 290 (6-7). 

D'Orbigny, A., 620 (5-3). 

Dodge, C. W. et al., 16977 (5-1). 

Dodson, C. H. & P. M. Dodson, 12001 (3-1). 

Dodson, C. H. & A. Gentry, 13726 (3-1). 

Dombey, J., 555 (3-1). 

Dorr, L. J. et al., 5719 (3-1). 

Dressier, R. L., 4207 (1-2); 4703, 4749 (1-3); 4912 (3-1). 

Drew, W. B., 546, E-607 (3-1). 

Dryander, E., 1976 (3-1); 2583 (1-2). 

Duarte, A. P., 989 (RB61684), 3037 (RB71834) (1-11); 4842 

(RB 110068) (8-lb). 

Duarte, A. P. et al., s.n. (RB60854) (1-11). 

Ducke, A., 38 (RB20719) (6-11); 860 (6-11); 1235 (MG19119) 

(6-10); 1618 (6-6); s.n. (MG15480), s.n. (MG15757) (6-10); 

s.n. (RB15400) (10-1); s.n (RB15685) (6-10); s.n. 

(RB15687) (10-1); s.n. (RB17355), s.n. (RB17356) (6-10); 

s.n. (RB17358) (6-11); s.n. (RB21691), s.n. (RB21692) (6- 

10); s.n. (RB22821) (1-11); s.n. (RB22845) (6-11, 6-12 

[mixed collection]); s.n. (RB22846) (6-12); s.n. (RB22847) 

(6-11); s.n. (RB35055) (6-11). 

Ducke, A. & J. C. Kuhlmann, s.n. (RB4620), s.n. (RB4688) 

(1-1 1). 

Dudley, T. R., 11488 (3-1). 

Duke, J. A., 4465, 5361, 6127 (5-1); 9653, 10567, 10605 

(1-2); 10621 (3-1); 14597 (5-1). 

Duke, J. A. & J. Idrobo, 11580 (1-2). 

Dunlap, V. P., 33 (6-1). 

Duss, R. P., 345, 1452, 1880, 1883, 2548, 3424 (6-5). 

Dwyer, J., 2231, 2290 (1-2); 3095 (5-1); 6178 (3-1); 8372, 

15265 (5-1). 

Dwyer, J. & B. MacBryde, 9626 (3-1). 

Dwyer, J. et al., 523 (3-1). 

E. L. C., s.n. (5-1). 

Edwall, G., 23938 (1-11). 

Eggers, F., 5812 (2-la); 13392 (5-2a). 

Elias et al., 583 (5-1). 

Elias, Bro., 450 (5-2a). 

Ekman, E. L., 3642, 4641, 5772, 7859, 7919, 9384 (4-1); 

9417, 9692 (6-3); 10408 (1-15); 10389 (6-4). 

Espina, J. et al., 2503 (1-2). 

Espinal S., T., 623, 633 (5-1). 

Eyerdam, W. J., 180 (4-1). 

Eyerdam, W. J. & A. A. Beetle, 22876 (5-3). 

Falcao, M. & D. Coelho, 209 (6-12). 

Fanshawe, D. B., F1249 (3985) (6-8). 

Famey, C. et al., 1237 (1-11). 

Fassett, N. C., 25427 (1-9); 25756 (3-1). 

Fendler, A., 289 (1-2); 584 (5-2a); 2366 (2-la). 

Fenmndez, A., 256 (1-2). 

Ferreira, V. F. et al., 285 (1-14). 

Ferreyra, R., 1112, 3612, 4208, 4542, 6823 (3-1). 

Fiebrig, K., 2705 (5-3). 

Field, A., 424 (6-9). 

Fishlock, W. C., 10, H338-16 (6-5). 

Flores-Paitan, S., 3B, 61 (6-7). 

Folsom, J. P., 6241 (5-1). 

Folsom, J. P. & R. Lantz, 1931 (5-1). 

Folsom, J. P. & R. Robinson, 2365 (6-1). 

Folsom, J. P. et al., 6581 (5-1). 

Fonnegra, R. & F. J. Roldan, 2278 (3-1). 

Fonnegra, R. et al., 2514 (3-1). 

Forero, E., 7689 (1-11). 

Forero, E. et al., 4418 (1-2); 6093 (3-1). 

Forest Department of British Guiana, 3985 (F1249), 5885 

(WB479), 6991 (F3427), 7892 (RB68) (6-8). 

Forestry Bureau Suriname, 1151, 1680, 2040, 2415, 2464, 

2585, 2907, 4952 (6-10).


Fors, A., 142 (6-3). 

Fosberg, F. R., 19440 (1-9); 19527 (7-1); 19880,20388 (3-1); 

21857 (7-1); 27750 (3-1); 28653 (5-3); 28672 (3-1). 

Fosberg, F. R. & L. R. Holdridge, 19409 (3-1). 

Foster, R. B., 1411 (1-2); 2877 (6-1); 4133 (1-2). 

Foster, R. B. & E. Vivar, 13333 (3-2). 

Frazod, A., s.n. (RB2316) (1-14). 

Freire, E. & J. Cerda, 461 (1-6). 

Freire-Fierro, A. et al., 2333 (3-1). 

Freire-Fierro, A. & J. Luteyn, 2159 (3-1). 

Freire-Fierro, A. & P. Yanez, 2697 (1-6). 

Freitas, C. A. A. et al., 83 (5-3). 

French Guiana Forest Service, 7608 (6-10). 

Friedrichshal, E. R. von, s.n. (1-2). 

Froes de Lemos, R., 20777 (6-10); 31049, 32097 (6-10); 

s.n. (RB22596) (6-12). 

Fuchs, H. P. & L. Zanella, 21703 (1-2). 

Fuentes, A. F., 471 (5-3). 

Fuertes, P., 649 (4-1). 

Funck, N., 372, 463 (5-2a). 

Funck, N. & L. J. Schlim, 1637 (3-1). 

Fundaci6n Biol6gica Puerto Rastrepo, 5052 (10-1). 

Garcia, 147 (6-9). 

Garcia-Barriga, H., 8197 (3-1); 18763 (7-1). 

Garcia-Barriga, H. & G. Stout, 18907 (3-1). 

Garwood, N. C., 719, 802, 1015, 1102, 1129 (3-1); 2531 

(6-1). 

Garwood, N. C. et al., 10 (1-2); 70 (3-1). 

Gaudichaud, C., 1830 (1-14). 

Gay, M. C., 919 (3-1). 

Gehrt, A., s.n. (1-11). 

Gentry, A., 1785, 5609, 6720 (5-1); 70741 (6-7). 

Gentry, A. & H. Cuadros, 55525 (5-2b). 

Gentry, A. & J. Aronson, 25307 (6-14). 

Gentry, A. & C. Diaz, 61118 (3-1); 58655 (5-3). 

Gentry, A. & S. Estensoro, 70562 (3-1); 70733 (6-7). 

Gentry, A. & M. Fallen, 17201 (3-1). 

Gentry, A. & E. Forero, 7195 (3-1); 7377 (1-2). 

Gentry, A. & N. Jaramillo, 28093 (6-6); 41321 (3-1). 

Gentry, A. & A. Juncosa, 40718 (3-1). 

Gentry, A. & W. Morawetz, 63183 (6-10). 

Gentry, A. & S. Mori, 13874 (5-1). 

Gentry, A. & G. Morillo, 10339 (5-2a). 

Gentry, A. & OTS class, 48572 (6-2). 

Gentry, A. & J. Revilla, 16645 (10-1). 

Gentry, A. & G. Shupp, 26475 (3-1). 

Gentry, A. et al., 27780, 31219,31384, 36636 (6-7); 37904 

(3-1); 40418A (1-2); 40125 (5-3); 40160 (3-1); 42942 

(6-6); 43018, 43027, 43135, 45661,46255 (6-7); 53411 

(1-2); 55625, 60913, 65564, 65621, 72152 (6-7). 

Gillis, W. T., 7535, 11168 (5-1). 

Giordano, L. C. et al., 1005 (1-11). 

Glaziou, A., Ibis, 148 (1-11); 1098 (1-12); 2607 (1-13); 

6564 (1-12); 12036 (1-14); 12038 (1-13); 13974, 13975, 

14892 (8-1); 14893 (1-14); 14937 (1-13); 18305 (5-2a); 

18306(1-13); 18309, 18905 (5-2a); 18906(1-13); 19437 

(1-14); 19438 (8-1); 21458a, s.n. (1-11). 

Gomes, M., 323 (1-13). 

G6mez-Laurito, J., 10694 (3-1); 11698 (1-1). 

G6mez P., L. D., 3307 (1-2). 

G6mez P., L. D. et al., 22770, 22944 (6-1), 23844 (3-1); 

24504 (5-1). 

Gosling, C., s.n. (5-3). 

Gottsberger, G. K., G213-13888 (3-1). 

Goudot, J., s.n. (1-9, 3-1). 

Goulding, M., 1048 (10-1). 

Grandez, G. et al., 1768 (6-13). 

Grant, M. L., 10620 (3-1); 10891 (5-2b). 

Grant, V., 630 (5-1). 

Granville, J. J. de, B-2671, B-2799, B-5167 (6-8); B-5377 

(6-10). 

Granville, J. J. de et al., 9041 (6-8). 

Grayum, M. et al., 4689 (3-1). 

Grenand, 1238, 1500(6-10). 

Grubb, P. J. et al., 1485 (1-8). 

Gudiflo, E., 1707 (6-6). 

Gudifio, E. & N. Andi, 2220 (6-7). 

Gudifio, E. & G. Grefa, 2336 (6-7). 

Gudifto, E. & C. Gualinga, 1640 (6-7). 

Guti6rrez, A., 32 (6-7). 

Haber, W. & E. Bello, 1735, 1736, 6837, 7152 (6-1). 

Hahn, M., 116 (5-1); 429 (6-5). 

Halton, J. & L. Besse, 101 (3-1). 

Hammel, B., 9753 (6-1); 12446 (6-1). 

Hammel, B. et al., 16858 (6-1); 17540 (3-1); 17726 (5-1); 

17719 (6-2). 

Harling, G., 3763 (1-6); 6138, 11016 (3-1); 11076 (1-6). 

Harling, G. & L. Andersson, 11860, 13755, 14065 (3-1); 

16423 (1-6); 16979 (3-1); 17003 (1-6); 23151,23944 (3-1). 

Harling, G. et al., 9916, 9769, 10079 (3-1). 

Harmon, E. W. & J. A. Fuentes, 4965 (5-1). 

Harris, W., 5810, 5828, 8783, 9449 (6-4). 

Harris, W. & N. L. Britton, 10576 (6-4). 

Hart, J., 109 (1-2). 

Hartshorn, G. S., 1211, 1500 (6-1); 1687 (6-7). 

Hartshorn, G. S. & J. Quijano, 2925 (6-6). 

Hartshorn, G. S. et al., 2906,2925,2932 (6-6); 62451 (5-3). 

Hatschbach, J. H., 23784 (5-3); 38796,42945,43851 (1-11). 

Haught, 0., 2236, 3705 (5-2b); 5427 (1-2). 

Hauman II, 5520 (5-3). 

Haymard, G. et al., 5233 (7-1). 

Hayes, S., 164 (5-1). 

H6guert, J. S. d', 642 (5-2a). 

Herbario Florestal Personnel, s.n. (8-lb). 

Heringer, E. P., 7065, 7681, 9156, 10468 (1-11). 

Heringer, E. P. et al., 1439, 1784, 3157, 3188, 3232, 4448, 

5082,6157(1-11). 

Hernmndez, J. J. & S. E. Hoyos, 182 (9-1). 

Herrera, G., 1976 (6-1); 3519 (3-1). 

Herrera, H., 1663 (1-3). 

Hill, S. R., 24770 (6-5). 

Hinds, 354, s.n. (1-2, 3-1). 

Hioram, Bro. & C. Maurel, 6066 (6-3). 

Hodge, W. H., 9 (6-7); 695 (6-5); 6048 (3-1). 

Hoene, F. C., s.n. (1-11). 

Hoffman, B. et al., 1532 (6-8). 

Holdridge, L. R., 6723 (6-1). 

Holm-Nielsen, L., 16311 (3-1). 

Holm-Nielsen, L. & J. Jaramillo, 28968 (3-1). 

Holm-Nielsen, L. et al., 3776, 20567, 20613 (3-1); 25235, 

26039, 26057 (1-2). 

Holton, I. F., 410 (7-1); 440 (3-1). 

Hoogte, L. v. d. & C. Roersch, 2791 (5-3). 

Hoover, W. S. & S. Wormsley, 1420 (3-1). 

Hoover, W. S. et al., 3047 (3-1). 

Horowitz, S., 663 (5-3). 

Howard, R. A., 11143 (6-5); 16044 (5-1). 

Howard, R. A. & L. I. Nevling, 16910 (5-1). 

Howard, R. A. & E. S. Howard, 18846, 19487 (6-5). 

Howard, R. A. et al., 19930 (6-5).

List of Exsiccatae213 

Howell, J. T., 10184A (1-2). 

Huashikat, V., 2136 (6-14). 

Huertas, G. & L. A. Camargo, 6282 (7-1). 

Humbert, H., 26973 (3-1). 

Humboldt, A., 567 (3-1). 

Hunziker, A. T., 2037 (5-3). 

Hurtado, F., 2365 (6-7). 

Hurtado, F. & A. Alvarado, 2240, 2261 (6-7). 

Hurtado L., R., 5, 111 (3-1). 

Hutchison, P. C., 1153 (3-1). 

Hutter, C., 1215 (7-1). 

Idrobo, J. M., 287 (3-1); 4874 (9-1). 

Idrobo, J. M. & R. E. Schultes, 570 (1-9); 1011 (9-1). 

Ijjasz-Madriz, E., 422 (3-1). 

INPA/WWF 1109.25 (6-10). 

Irvine, D., 634 (6-6). 

Irwin, H. S. & T. R. Soderstrom, 5369 (1-11). 

Irwin, H. S. et al., 8881, 14160, 25206(1-1 1); 48440, 50615, 

55443 (6-10). 

Isem, J., 2382 (3-1). 

Jack, J. G., 8173, 8430 (5-1). 

Jacobs, B., 2315 (6-1). 

Jahn, A., 1339 (6-9). 

Jangoux, J. & R. P. Bahia, 86 (6-10). 

Jansen-Jacobs, M. J. et al., 1899 (6-10). 

Jaramillo, J., 14167, 14290 (1-8); 14990 (1-2). 

Jaramillo, J. & F. Coello, 4322 (1-8). 

Jaramillo, J. et al., 13199 (3-1); 31361 (6-6). 

Jardim, A. & A. Cadden, 2153 (5-3). 

Jardim, A. & Mamami, 1423 (5-3). 

Jardim, J. G. et al., 711 (8-la). 

Jativa, C. & C. Epling, 771 (1-2); 870 (3-1); 2001,2201 (1-2). 

Jelski, de, 376 (7-1). 

Jimenez, Q. & R. Zuniga, 760 (6-2). 

Jim6nez, Q. et al., 751 (5-1). 

Jim6nez L., 0., 541 (5-1). 

Jim6nez M., A., 2451 (3-1); 2898, 3173 (1-2); 3418 (6-1); 

3728 (3-1). 

Johnson, H., 57, 1022 (5-1). 

Johnston, I. M., 1745 (1-2). 

Jorgensen, P. M. & S. Laegaard, 56452 (3-1). 

Jorgensen-Hansen, P., 35009 (5-3). 

Juncosa, A., 1589 (1-2); 1660, 2435 (3-1); 2543 (1-2). 

Juzepczuk, S., 6869 (3-1). 

Kalbreyer, W., 927 (5-2a); 1246 (1-2). 

Kanehira, R., 107 (3-1). 

Karsten, H., s.n. (1-9); s.n. (2-la). 

Kayap, R., 929, 2012 (3-1). 

Keman, H. S., 19, 89 (3-1); 167 (5-2b). 

Kiesling, R., 7109 (5-3). 

Killip, E. P., 3284 (5-1); 9735 (3-1); 11707 (1-2); 11199 

(3-1); 34986 (1-2). 

Killip, E. P. & J. Cuatrecasas, 38680, 38970, 39150 (1-2). 

Killip, E. P. & H. Garcia-Barriga, 32209, 33128, 33209, 

33338 (1-2); 33635 (3-1). 

Killip, E. P. & A. C. Smith, 22449, 23506 (3-1); 23811 (5-3); 

24130 (3-1). 

King, S. R. & M. Ramirez-Rengifo, 372 (3-1). 

Kirizawa, M. & S. A. C. Chiea, 2350 (1-1 1). 

Kirkbride, J. H., 2637, 2667 (5-2b); 5431 (1-11). 

Klawe, W. L., 1492 (1-2). 

Klug, G., 1804 (9-1); 3672 (3-1). 

Knapp, S., 1335 (1-2); 2713 (5-1). 

Knapp, S. & J. Mallet, 3035 (3-1); 3070 (1-2); 6764 (2-la). 

Knapp, S. et al., 4586 (6-1). 

Krapovickas, A. & A. Schinini, 31024,31193,31475,39017 

(5-3). 

Krapovickas, A. et al., 19143 (5-3). 

Krukoff, B. A., 1018 (6-10); 8782 (6-14); 10032 (3-1); 

10054, 10167 (5-3). 

Kuhlmann, M., 3769 (1-11). 

Kuhlmann, M. & S. Jimbo, 310, 329 (6-10). 

Kuhlmann, J. G., 82 (RB14083) (8-lb); 503 (RB139367) 

(1-11); 1221 (8-lb); 2353 (RB15366) (5-3); s.n. 

(RB16318)(1-11); s.n. (RB17357)(6-12); s.n. (RB17384) 

(6-11); s.n. (RB70222) (1-1 1). 

Kuntze, O., s.n. (5-3). 

Lamarck, W., s.n. (6-5). 

Lanjouw, J. & J. C. Lindeman, 2810 (6-10). 

Lansberge, J. G. van, 214 (5-2a). 

Lao Magin, R., 44, 51, 5107 (6-7). 

Lasser, T., 2264 (6-9). 

Lee, K., 49 (6-5). 

Legname, P. R. & A. R. Cuezzo, 5967, 6129, 7964, 9047 

(5-3). 

Lehmann, F. C., 473 (6-4); 5651 (3-1); 7459 (5-1); 7718 

(7-1); K-132 (1-2); s.n. (1-9). 

Leite, I., 2 (6-12). 

Lemos, C., 28678 (1-11). 

Lent, R. W., 359 (1-2). 

Le6n, Bro.,455 (5-1); 11011, 12813 (6-3). 

Le6n, Bro. & Matos, 19644 (4-1). 

Le6n, Bro. & M. Victorin, 17235 (4-1). 

Le6n, Bro. et al., 9941 (6-3). 

Le6n, J., 1484 (3-1); 2143 (6-1). 

Lescure, 2060 (6-6). 

Lewis, M., 88148, 88825, 88914 (3-1). 

Lewis, W. H. et al., 1 (5-1); 692 (3-1); 3392 (5-1); 13543 

(6-14). 

Liesner, R. & A. Gonzales, 10940 (1-9). 

Lillo, M., 10785, 22336 (5-3). 

Lima, J. & B. Zimmerman, 519 (6-10). 

Linden, J., 874 (3-1); 1604 (9-1); s.n. (5-2b). 

Lino, A. M., 75 (8-la). 

Liogier, A., 14353 (4-1). 

Liogier, A. & J. Acunia, 7673 (4-1). 

Liogier, A. & P. Liogier, 26174 (4-1). 

Little, E. L. Jr., 7186 (1-9); 8205 (7-1); 8490, 8757, 15872 

(3-1). 

Little, E. L. Jr. et al., 298 (3-1), 763 (6-6). 

L0jtnant, B. & U. Molau, 13513 (6-7). 

L6pez-Palacios, S., 1939, 1940 (5-2a). 

Lorentz & Hieronymus, 384 (5-3). 

Lourenco, s.n. (RB1042) (1-11). 

Lourteig, A., 714 (5-3). 

Lozano C., G. & J. Diaz, 3734 (7-1). 

Lozano C., G. et al., 2832 (5-2b); 4395, 4885 (7-1). 

Lugo, M., 111(3-1); 115(1-6); 141 (3-1). 

Lugo S., H., 1871 (3-1); 3674, 3946 (1-7); 4427,6034 (3-1). 

Luschnath, B., s.n. (1-11). 

M. A. V., 42 (6-7). 

Maas, P. J. M. & L. Y. T. Westra, 3912 (6-8). 

MacBride, J. F., 3662 (3-1); 5343 (5-3). 

MacBride, J. F. & J. Dwyer, 1408 (6-6). 

Macedo, A., 2402 (3-1). 

Machado, A. A., 11-1984 (5-2a). 

Macias & Burbano, 7788 (6-7).


Maciel, U. N. et al., 776 (6-10). 

Madison, M. T. et al., 4562 (3-1); 4808 (1-4); 5051 (3-1). 

Madsen, J., 85961 (7-1). 

MagalhAes, M., s.n. (8-lb). 

Maguire, B. & L. Politi, 27971 (6-13). 

Mamani, F., 436 (5-3). 

Manara, B., s.n. (5-2a). 

Marcano Berti, L., 241, 1426 (6-8). 

Marin, 2109 (3-1). 

Marin, E., 740 (6-13). 

Maroto, R. & E. Salsa, 123 (5-2a). 

Marques, M. C. et al., 285 (1-11). 

Marquete, R. et al., 832, 1084, 1371 (1-11). 

Martinelli, A. et al., 4133 (1-11). 

Martinelli, G., 6835 (6-10). 

Martinelli, G. et al., 13320 (RB28042) (1-11). 

Martinet, M., 1412, s.n. (3-1). 

Martius, K. F., 57 [in. obs.] (1-13); 111 [in. obs.] (1-11); 

2607, 2608 (1-9); 2964, 2965, 2966, 2967 (1-13); 3139, 

3201 (1-9). 

Mathias, M. E. & D. Taylor, 5932 (3-1). 

Mathews, A., 3030 (3-1). 

Mattheus, 1954 (3-1). 

Mattos, A. et al., 322 (1-11). 

Mattos Silva, L. A. et al., 2847 (8-la). 

Matuda, E., 273, 560, 16811, 18748 (5-1). 

McDaniel, S. & M. Rimachi Y., 20503 (10-1). 

McDowell, T. et al., 4411, 4472 (6-8). 

McPherson, G., 9146 (6-1). 

Mendonca, R.C. et al., 1292 (1-11). 

Meneces, E. & W. Terceros, 9 (6-6). 

Metcalf, R. D., 30676 (3-1). 

Mexia, Y., 6351 (3-1); 8035 (5-3). 

Meyer, T., 903, 4810, 4811, 8477 (5-3). 

Meyer, W. & S. Llamosa, 3627 (2-la). 

Meyer, W. et al., 2039 (2-la). 

Michel, R. de, 41 (5-3). 

Molina, F., s.n. (3-1). 

Molina R., A. et al., 1868, 2085 (1-2); 18005 (3-1). 

Montalvo, E. A., 3849 (5-1). 

Monteiro, M. T., 23498 (8-la). 

Moore, A., 10648 (6-4). 

Morales, J. F. et al. 1124 (6-2). 

Morawetz, W. & B. Walln6fer, 11-161085 (3-1). 

Moreno, L. M., 193 (5-2b). 

Moreno, P. P., 23868, 26317 (6-1). 

Mori, S., 421 (5-1). 

Mori, S. & J. Cardoso, 17128 (6-10). 

Mori, S. & C. Gracie, 18943 (6-8). 

Mori, S. & J. Kallunki, 3086 (1-1). 

Mori, S. & J. Pipoly, 15562 (6-8). 

Mori, S. et al., 10328 (8-la); 15016, 15109 (6-8); 15786, 

16248, 18691 (6-10); 20910 (6-8); 21567, 23098, 23645 

(6-10). 

Morillo, G. et al., 3298 (2-la). 

Moritz, K., 840 (5-2a). 

Mos6n, H., 3408 (1-11). 

Mulford Biological Exploration, 1016 (5-2a). 

Museu Goeldi Personnel, 9425, 9713 (6-10). 

Mutis, J. C., 2 (3-1); 3 (1-9); 1004 (3-1); 2194 (7-1); 4651 

(3-1). 

Nascimento, J. R. M. et al., s.n. (INPA/WWF 1109.302) 

(6-12). 

Nee, M., 8846, 9125 (5-1); 41570 (5-3). 

Nee, M. & G. Coimbra S., 35199 (5-3). 

Nee, M. & M. Saldias P., 36320 (5-3). 

Neill, D., 9817, 9840, 9848 (6-7). 

Neill, D. & A. Alvarado, 9040 (6-7). 

Neill, D. & W. Palacios, 7104 (6-7); 9718 (1-8). 

Neill, D. & J. Zaruma, 365, 7021 (6-7). 

Neill, D. et al., 6105 (3-1); 6204 (6-7); 6220 (6-6); 6515 (6-6); 

7394, 9145 (3-1). 

Nufiez, P., 6104 (3-2). 

Nufiez, P. & C. Mufioz, 5122 (3-1). 

Nufiez, P. et al., 10098 (5-3). 

Occhioni, P., 6655 (1-14); s.n. (RB15455) (5-1). 

Oldeman, R. A. A., B-674 (6-10). 

Oldeman, R. A. A. & Arevalo, 16 (6-7). 

Oliveira, A. de, s.n. (INPA 60564) (6-12). 

Oliveira, E. de, 3836, 3854, 4036, 4798 (6-10). 

Oliveira, R. P., 5 (3-1). 

011gaard, B. et al., 34627, 34667, 35520, 90375 (3-1). 

Orlandi, R., 223 (8-la). 

Orsted, A. S., 11536, 11537, s.n. (5-1). 

Otto, E., 901 (5-1). 

Ownbey, M., 2682 (1-6); 2686 (3-1); 2735 (1-8). 

Pab6n E., M., 960 (1-9). 

Pabst, G., 4667 (1-11). 

Palacios, W., 1994 (6-6); 2828 (6-7); 6616 (1-8). 

Palacios, W. & J. Clark, 12633 (7-1). 

Palacios, W. & D. Rubio, 7290 (7-1). 

Palacios, W. et al., 240, 365 (6-7); 3380 (6-6); 9663 (1-4). 

Parodi, L. R., 9220 (5-3). 

Patris, s.n. (6-10). 

Paula, J. E. de, 3233 (1- 1). 

Paulino Filho, H. F. & M. G. Silva, 93-96 (5-3). 

Pearce, R. W., s.n. (5-3). 

Pennell, F. W., 3503 (3-1). 

Peredo, Y., 525 (5-3). 

Pereira, B. A. S., 270 (1-11). 

Pereira, B. A. S. & D. Alvarenga, 2503 (1-11). 

Pereira, B. A. S. & V. V. Mecenas, 2442 (1-11). 

Pereira, E., 7363 (1-11). 

Pereira, E. & A. P. Duarte, 4484 (1-11). 

Pereira, E. et al., 3762 (RB266353) (1-11). 

Perez, P., s.n. (3-1). 

Perez-Arbelaez, E., 697 (3-1). 

Perez-Arbelaez, E. & J. Cuatrecasas, 5733 (3-1). 

Perrottet, 283 (6-5). 

Persson, C. et al., 43 (1-7). 

Pflanz, K., 937, 2053 (5-3). 

Philipson, W. R. & J. M. Idrobo, 1921 (1-9); 1955 (9-1). 

Philipson, W. R. et al., 1576 (9-1). 

Picarda, L., 1129(4-1). 

Pierotti, S. A., 76 (5-3). 

Pinto et al., 370 (1-9). 

Pipoly, J. J. et al., 12525 (6-7). 

Pirani, J. R. & R. Mello Silva, SPF 47733 (8-lb). 

Pires, J. M., 6826, 6922, 6969, 6978, 7107, 48902 (6-10). 

Pires, J. M. & R. P. Belem, 13006 (6-10). 

Pires, J. M. & N. T. Silva, 1742, 10879, 10880, 11895, 

11897, 11898 (6-10). 

Pires, J. M. et. al., 4997 (6-10); 9270, 9539 (1-11); 50615, 

50853, 51224, 51420 (6-10). 

Pittier, H., 58,236 (5-2a); 2396,2476 (5-1); 4166 (1-2); 4817 

(5-1); 5251 (3-1); 5507 (5-1); 7468 (5-2a); 9799, 10299, 

11672 (5-2a); 13854,14138,15275 (6-9); 15582 (5-2a). 

Pl1e, A., s.n. (6-5). 

Plowman, T. & W. Davis, 4762, 5164 (5-3).

List of Exsiccatae 215 

Plowman, T. & H. Kennedy, 5649 (5-3). 

Pohl, J. E., 809, 809d, 810, 2681, 3442, 3444, s.n. (1-11). 

Ponthieu, de, s.n. (6-5). 

Popenoe, W., 802 (5-1). 

Prance, G. T. et al., 4452 (6-8); 12462 (6-6). 

Prieto, F., ChuP-20 (1-7). 

Proctor, G. R., 19742, 19818, 21467, 22591 (6-4). 

Proctor Cooper III, G., 38, 38A (6-5). 

Pulle, A., 379 (6-8). 

Purdie, W., s.n. (1-9). 

Quevedo, R., 316 (5-3). 

Rabelo, B. V. & J. Cardoso, 2887 (6-10). 

Ragonesi & Coras, 37794, 37799 (5-3). 

Ramage, G. A., s.n. (6-5). 

Ramirez, J. G. & D. Cardenas Lopez, 179, 1017, 1501, 1620, 

1879 (9-1). 

Ramos, J. & G. Mota, 368 (5-3). 

Ramos, L. S. & C. Rosario S., 15 (6-10). 

Renteria, E. et al., 658 (3-1). 

Revilla, J. & C. H. Froehner, 2449 (6-6). 

Reynel, C. & J. Le6n, 4044 (3-1). 

Ribamar, J. & J. Ramos, 164 (6-10). 

Ribeiro, J. E. L. S. & P. A. C. L. Assuncao, 1776 (INPA 

185097) (6-12). 

Richard, L. C. M., s.n. (6-5). 

Riedel, L., 66,68 (1-13); 187 (1-11); 367 (1-14); 369 (5-3); 

424 (1-14); 480 (1-11); 1236 (1-11); 1229 (5-3); 1462 

(1-11); s.n. (1-11). 

Riedel, L. & B. Luschnath, 1062 (1-13). 

Riera, B. & D. Sabatier, 1844 (6-10). 

Rimachi Y., M., 2973 (6-6); 10178 (3-1); 10501 (10-1). 

Rizzini, C. T. & A. Mattos, 1126 (8-la). 

Robinson, W. & M. W. Lyon, s.n. (5-2a). 

Rodrigues, R. R. & N. Figueredo, 15745 (1-11). 

Rodrigues, W., 8573, 9954, 9976 (6-12). 

Rodrigues, W. & D. Coelho, 7600 (INPA 16903) (6-12). 

Rodriguez, H., 266 (5-2a). 

Rodriguez, W., 8098, 8099, 8692 (6-10); 9954 (6-12). 

Roig, F. A., 6419 (6-3). 

Rojas, T., 1145, 1546 (5-3). 

Romero-Castafieda, R., 132, 609 (5-2b); 2309 (3-1); 11198 

(5-2b); s.n. (1-2). 

Rosa, N.A., 1096 (6-10). 

Rosa, P., 145 (RB131708) (1-11). 

Rosario S., C. et al., 852, 858 (6-11); 1092 (6-10). 

Rose, J. N. & G. Rose, 21691 (5-2a); 23426 (3-1); s.n. (5-2a). 

Rudas, A. et al., 3016 (6-6). 

Ruiz L., H. & J. J. A. Pav6n, s.n (3-1). 

Ruiz Teran, L. E., 488 (6-9); 1204 (3-1). 

Ruiz Teran, L. E. & M. L6pez Figueras, 658 (3-1). 

Rusby, H. H., 106, 711, 1898 (3-1); 2090 (5-3). 

Rusby, H. H. & R. W. Squires, 126 (6-8). 

Sabatier, D., 606 (6-10). 

Sabatier, D. & M. F. Prevost, 2116, 3356, 3731 (6-10). 

Saer, d'H., J., 642 (5-2a). 

Sagastegui A., A., 5872 (3-1). 

Saint George Expedition, 538, 624 (1-2). 

Saint Hilaire, A., 348 (1-11). 

Saint John, H., 20813, 20871 (3-1). 

Salim, J. Filho, 148, 159 (8-1a). 

Sampaio, A., 612 (1-11). 

Sanchez Vega, I. & M. Dillon 9006, 9052 (3-1). 

Sandemer, 6055 (3-1). 

Sandino, J. C., 3294, 3355, 3412, 4842 (6-1). 

Sandwith, N. Y., 1757 (2-1a). 

Sanoja, R. E., s.n. (3-1). 

Santillan, P., 90436 (5-3). 

Santos, T. S., 920 (8-la). 

Saundeman, C., 6055 (3-1). 

Schlim, L., 755 (5-2a); s.n. (5-2b). 

Schnee, L., 1174 (5-2a). 

Schreiter, 220, 3536, 11168 (5-3). 

Schultes, R. E., 24356 (1-9). 

Schultes, R. E. & I. Cabrera, 18929 (6-6). 

Schultz, A. G., 978 (5-3). 

Schulz, J. P., 7289, 8581 (6-10). 

Schunke, J. M., 331 (6-6). 

Schunke Vigo, J., 1527 (6-6); 2072 (3-1); 2901 (6-6); 3205, 

3952 (1-8); 4364,4815 (3-1); 4877,5683 (1-8); 5807,5845 

(3-1); 6162 (1-6); 6214 (5-3); 6696 (1-8); 7267,7659,9494, 

10079(3-1); 10270(1-8); 10362,10512(3-1); 10653(6-6). 

Schwacke, C. A. W., 1441 (5-1). 

Scolnick, R., 1678 (5-3). 

Scolnick, R. & R. Luti, 497 (3-1). 

Seemann, B., 478 (5-1); 1593, s.n. (3-1), s.n. (5-1). 

Servicio Forestal do Brasil, Personnel, 1043 (1-11). 

Shafer, J. A., 4406 (6-3); 7850 (4-1). 

Shattuck, O. E., 688 (5-1). 

Shepherd, J. D., 459 (9-1). 

Sieber, F. W., s.n. (6-5). 

Silva, A. S. L. da, et al., 1553, 2217 (6-10). 

Silva, M., 2794, 5073 (6-10). 

Silva, M. F., 576 (6-10). 

Silva, M. G., 5517 (6-10); 6125 (5-3). 

Silva, M. G. & C. Rosario S., 2420, 2475, 3732 (6-10). 

Silva, N. T., 51, 628, 905, 999, 1246, 2561, 2568, 3421, 

3924 (6-10). 

Silverstone-Sopkin, P. et al., 3001 (7-1). 

Skutch, A. F., 3950 (5-1); 3981 (3-I); 4777 (1-1). 

Slane, V., 140 (6-5). 

Sleumer, H., s.n. (5-3). 

Smith, A., 1890 (6-1). 

Smith, A. C., 3433 (6-8). 

Smith, C. E. et al., 3282 (5-1). 

Smith, D. N., 1311 (6-6); 1415 (5-3); 2075 (1-6); 4019 (6-6). 

Smith, D. N. et al., 1415 (5-3); 1423 (3-1); 5605 (5-3); 6325 

(3-1). 

Smith, E. E., 396 (6-3). 

Smith, H. H., 100 (5-2b). 

Smith, H. H. & G. W. Smith, 889, 1245 (6-5). 

Smith, L. B. & E. L. McWilliams, 15364 (1-11). 

Smith, R. F., VI-280 (5-2a). 

Sobel, G. L. & J. Strudwick, 2430 (3-1). 

Sodiro, P. L., s.n. (1-6). 

Soejarto, D. D., 1086 (7-1); 3443 (5-1). 

Soejarto, D. D. et al., 1287 (9-1). 

Solanilla, O., 16 (5-1). 

Solomon, J. C., 3097 (5-3); 3153 (3-1); 7321 (5-3); 7322 

(3-1); 7359, 17641 (5-3); 18445 (3-1). 

Solomon, J. C. & J. Kuijt, 11585 (3-1). 

Sothers, C. et. al., 568 (INPA185102), 625 (INPA185105), 

750 (INPA185110) (6-11). 

Soukup, J., 597, 2281 (3-1). 

Spada, J., 31, 289, 293 (8-lb). 

Sparre, B., 14380 (3-1); 18300, 18340 (1-2). 

Spegazzini, C., 13814 (5-3). 

Spruce, R., 4579 (3-1); 4930 (6-7). 

Stahel, G., 261, 261a (6-10). 

Standley, P. C., 21672, 29972, 62055, 79549 (5-1).


Standley, P. C. & E. Padilla V., 2885 (5-1). 

StAhl, B. et al., 1822, 1866 (1-7); 2800 (1-6). 

Steere, W. C., 8016 (3-1); 8021 (1-6); 8060, 8337 (3-1); 

8339 (1-6). 

Steere, W. C. & W. H. Camp, 8228 (1-6); 8255 (3-1). 

Stehle, H., 987, 2298, 3015, 3652, 5068, 6626, 7178 (6-5). 

Stein, B. A. & D. D'Alessandro, 2776 (3-1). 

Stein, B. A. & C. Todzia, 2142, 2358 (3-1); 2372 (5-3). 

Steinbach, J., 9292 (3-1). 

Stem, W. L., & D. Wasshausen, 2582 (6-5). 

Stem, W. L., et al., 724 (5-1). 

Stevens, F. L., 662 (5-1). 

Stevens, W. D., 24597 (6-1). 

Stevens, W. D. et al., 24859 (6-1). 

Stewart, A., 315 (1-2). 

Steyermark, J. A., 54450 (7-1); 54732, 54865 (3-1); 54921 

(5-2a); 55220, 55223, 55408 (7-1); 56208, 56306 (3-1); 

56333 (1-10); 56659 (3-1); 57662 (5-2a); 61136, 62185 

(6-8); 65741 (5-2a); 87104 (6-9); 89514 (6-8); 89885, 

89961 (6-9); 91807 (5-2a); 100495 (9-1). 

Steyermark, J. A. & P. Berry, 112141 (5-2a). 

Steyermark, J. A. & C. B. Carias, 105353 (5-2a). 

Steyermark, J. A. & G. Davidse, 116278, 116885 (6-9). 

Steyermark, J. A. & V. C. Espinoza, 112706,108790 (5-2a). 

Steyermark, J. A. & M. Rabe, 96163 (2-la). 

Steyermark, J. A. & C. Steyermark, 95211 (6-9). 

Steyermark, J. A. et al., 111625 (6-8); 114008 (5-2a); 

121534 (2-la). 

Stork, H. E. et al., 10494 (5-3). 

Strier, K. B., 1007 (8- b) 

Sucre, D., 88 (5-3). 

Sucre, D. & E. P. Heringer, 624 (1-11). 

Svenson, H. K., 331 (1-2). 

Sytsma, K. & W. G. D'Arcy, 3391 (5-1); 3480 (3-1). 

Tafalla, J. J. & J. A. Manzanilla, s.n. (3-1). 

Tate, H. H., 488 (3-1); 543 (5-3). 

Tello, 87, 129,313 (6-6); 489, 862, 1284 (6-7); 1295, 1434, 

1995 (6-6). 

Tessmann, G., 4512 (3-1); 4668 (6-7). 

Thiel, J., 878 (6-10). 

Thomas, J. et al., 1437 (1-9). 

Thompson, S. A. & J. E. Rawlins, 816 (3-1). 

Thompson, S. A. et al., 9859 (4-1). 

Tonduz, A., 3947, 4671 (3-1). 

Toro, R. A., 421 (3-1); 540 (5-1). 

Tovar, O., 1421, 4629 (5-3). 

Triana, J., 1782 (3262.1) (3-1); 1784 (3262.3) (1-2); 1785 

(3262.4) (1-9); 1869 (7-1); s.n. (3262) (1-9). 

Trigoso, J. R., 1025 (3-1). 

Troll, C., 1514 (5-3); 1745 (3-1). 

Trujillo, B., 1413, 3416 (5-2a). 

Trujillo, B. & A. Fernmndez, 293 (5-2a). 

Tunqui, S., 205, 945 (6-7). 

Turpe, A. M. et al., 4690 (5-3). 

Tuxill, J. et al., 63 (6-5). 

Tyson, E. L., 2629 (5-1). 

Uferide, T., 13 (6-9). 

Ule, E., 6774 (9-1); 9848, 9848b (5-3). 

Uribe Uribe, L., 912, 1441, 1481 (5-1); 5468 (7-1); 6385 (3-1). 

Utley, J. & K. Utley, 5337 (6-1). 

Vaca, A. A., 18(5-3). 

Vaillant, P., 21 (8-la). 

Van der Werff, H. & E. Gudifio, 10874 (3-1); 11098 (7-1); 

11326 (9-1). 

Vargas C., C., 1719, s.n. (5-3). 

Vargas, H. & P. Grefa, 908 (1-6). 

Vargas, H. et al. 563 (6-7). 

Vargas V., V., 2575 (3-1). 

Vasquez, C. B., 14-CBV (6-6). 

Vasquez, R., 10631(6-6). 

Vasquez, R. & N. Arevalo, 8969 (6-14). 

Vasquez, R. & G. Criollo, 1855 (6-7). 

Vasquez, R. & N. Jaramillo, 2431 (6-6). 

Vasquez, R. et al., 5404 (6-13). 

Vauthier, 15, 205, s.n. (1-11). 

Ventenat, L., s.n. (1-11). 

Ventura, E. & E. L6pez, 424, 2856 (5-1). 

Venturi, S., 5207, 7648 (5-3). 

Vivas, J. M., 2 (5-2a). 

Vogel, S., 54 (1-9). 

Wagner, M., 3495, 3496 (5-1). 

Wagner, R. J., 711, 897 (5-1). 

Walker, F., 1277 (5-2b). 

Walln6fer, B., 26-19588 (3-1); 113-29788 (1-6). 

Warming, E., s.n. (8-1b). 

Warner, R. N., 316 (1-2). 

Warscewicz, J., 10, 11 (1-9). 

Weberbauer, A., 7940 (5-3). 

Webster, G. L., 13478 (6-5). 

Webster, G. L. & N. McCarten, 28816 (3-1). 

Webster, G. L. et al., 8341 (6-4); 9205, 9403 (6-5); 9805 

(2-la). 

Weddell, H. A., 2637, 2638 (1-11); 4190 (5-3); s.n. (5-3). 

Wedel, H. von, 2114 (1-2). 

Werling, L. & S. Leth-Nissen, 226 (3-1). 

West, J., 8261 (5-3). 

White, J. W. & R. H. Warner, 82 (1-2). 

Widgren, J., 491, 1036, s.n., s.n. (1-11). 

Wiggins, I. L., 10880 (8-1); 10947 (3-1). 

Wilbur, R. L., 37205 (6-1). 

Wilbur, R. L. et al., 8137 (6-5). 

Williams, LI., 369, 409 (6-7); 7794 (3-1); 10299, 10415, 

10646, 12408 (5-2a). 

Williams, L1. & A. H. G. Alston, 42 (5-2a). 

Williams, L. O., 5066 (8-Ib). 

Williams, L. O. & V. Assis, 6861 (8-la). 

Williams, L. O. et al., 28805 (3-1). 

Williams, R. O., 11881(6-8). 

Williams, R. S., 322 (3-1). 

Willink, 174(5-3). 

Wilson, F. G. & S. J. Browne, WB479 (5885) (6-8). 

Woods, C. W., 497 (5-2a). 

Woronow, G. J. N. & S. V. Jusepczuk, 6426 (9-1). 

Woytkowski, F., 6663 (3-1); 7380 (6-6). 

Wright, C., 1262, 1622 (6-3); 2661 (4-1). 

Wullschlagel, 1344bis (6-4). 

Wurdack, J. J., 1383 (7-1); 1831 (3-1); 2348 (9-1). 

Young, K., 942 (3-1). 

Young, K. & M. Eisenberg, 233 (5-3); 255, 342 (3-1). 

Young, K. & G. Sullivan, 610, 800, 864 (3-1). 

Young, K. et al., 493 (3-1). 

Yuncker, T. G., 4529 (5-1). 

Zabala, S., 165 (5-3). 

Zak, V., 3979 (6-7). 

Zak, V. & S. Espinoza, 5130 (6-6). 

Zak, V. & J. Jaramillo, 3577 (3-1). 

Zamora, N. et al. 1901 (6-2). 

Zanderij, I., 261 (6-10).

Nomenclatural List 217 

Zanoni, T. & J. Pimentel, 35991 (4-1). 

Zanoni, T. et al., 25634, 33125, 40662 (4-1). 

Zarucchi, J. L., 3265, 3338 (9-1). 

Zarucchi, J. L. et al., 5398 (3-1); 5465 (5-1); 6104 (3-1); 

7044 (5-1). 

NOMENCLATURAL LIST 

Zaruma, J., 802 (3-1). 

Zaruma, J. & A. Arguello, 451 (6-6). 

Zehntner, M., 38 (5-2a). 

Zuloaga, F. O. & N. B. Deginani, 331 (5-3). 

Zuluaga R., S., 1184 (5-1). 

Due to the complexity of the taxonomic problems and large number of transfers and rearrangements in the 

present treatment, a list of accepted taxa (boldface), orthographic variants, and synonyms, as recognized by 

the author, is given. Synonyms, orthographic variants, and unpublished names are followed by the corresponding 

accepted taxon. Authority abbreviations follow Brummitt & Powell, 1992. 

Allenathus Standl. 

A. erythrocarpus Standl. 

Acunaeanthus Borhidi, Jarai-Koml. & Moncada 

A. tinifolius (Griseb.) Borhidi 

Arachnotryx Planch. =Rondeletia 

Ariadne Standl. =Mazaea 

A. ekmanii Urb. =Mazaea shaferi 

A. shaferi (Standl.) Urb. =Mazaea shaferi 

Augusta Pohl (nomen cons.) 

Augusta subgen. Lindenia (Benth.) J.H.Kirkbr. 

Augusta subgen. Augusta (Pohl) J.H.Kirkbr. 

A. attenuata Pohl =Augusta longifolia var. parvifolia 

A. austro-caledonica (Brogn.) J. H. Kirkbr. 

A. glaucescens Pohl =Augusta longifolia var. longifolia 

A. lanceolata Pohl =Augusta longifolia var. longifolia 

A. longifolia (Spreng.) Rehder 

A. loagifolia (Spreng.) Rehder var. longifolia 

A. longifolia (Spreng.) Rehder var. parvifolia (Pohl) Delprete 

A. oblongifolia Pohl =Augusta longifolia var. longifolia 

A. parvifolia Pohl =Augusta longifolia var. parvifolia 

A. rivalis (Benth.) J.H.Kirkbr. 

A. vitiensis (Seem.) J.H.Kirkbr. 

Badusa A.Gray 

B. corymbifera (Forst.f.) A.Gray 

Bartramia W. Bartram =Pinckneya 

B. bracteata W.Bartram = Pinckneya bracteata 

(W.Bartram) Raf. 

Bartramia L. =Triumfetta Hedw. (Tiliaceae) 

Bartramia Hedw. (Bryophyta) 

Bartramia Salisb. =Penstemon Schmidel 

Bathysa Presl 

B. bathysoides (Steyerm.) Delprete 

B. bracteosa (Wedd.) Delprete 

B. difformis Benoist =Chimarrhis turbinata 

B. perijaensis (Steyerm.) Delprete 

B. pittieri (Standl.) Steyerm. 

Bignonia W. Bartram =Pinckneya 

Bignonia L. (Bignoniaceae) 

B. bracteata W.Bartram = Pinckneya bracteata 


Blandibractea Werham =Simira 

B. brasiliensis Wernham =Simira glaziovii 

Bonifacia Manso ex Steud. =Augusta 

B. riparia Manso ex Steud. =Augusta longifolia var. longifolia 

Buena muzonensis (Goudot) Wedd. =Ladenbergia 

muzonensis 

Calycophyllum DC. 

C. tubulosum Seeman =Pogonopus exsertus 

C. tubulosum (A.Rich. in DC.) DC. =Pogonopus tubulosus 

Capirona erythroxylon Standl. =Simira ecuatoriensis 

Carmenocania Wernham =Pogoaopus 

C. porphyrantha Wernham =Pogonopus speciosus var. 

speciosus 

Cascarilla muzonensis (Goudot) Wedd. =Ladenbergia 

muzonensis 

Chimarrhis Jacq. 

Chimarrhis Jacq. subgen. Chimarrhis 

Chimarrhis Jacq. subgen. Pseudochimarrhis (Ducke) 

Delprete 

C. barbata (Ducke) Bremek. 

C. bathysoides Steyerm. =Bathysa bathysoides 

C. brevipes Steyerm. 

C. clausicorollata J.H.Kirkbr. =Simira tinctoria 

C. cubensis Steyerm. 

C. cymosa Jacq. 

C. cymosa Jacq. subsp. genuina Urb. =Chimarrhis cubensis 

C. cymosa Jacq. subsp. jamaicensis Urb. =Chimarrhis 

jamaicensis 

C. cymosa Jacq. var. jamaicensis (Urb.) Standl. =Chimarrhis 

jamaicensis 

C. cymosa Jacq. subsp. microcarpa Urb. =Chimarrhis cubensis 

C. cymosa Jacq. var. microcarpa (Urb.) Standl. =Chimarrhis 

cubensis 

C. cyrrhosa Stend. (orth. var.) =C. cymosa 

C. decurrens Steyerm. =Allenanthus erythrocarpus 

C. dioica K.Schum. & K.Krause =Dioicodendron dioicum 

C. duckeana Delprete 

C. duckei Rizzini =Chimarrhis turbinata 

C. ferruginea (Standl.) Standl. =Bathysa sp. 

C. gentryana Delprete 

C. glabriflora Ducke 

C. goudotii Baill. =Simira goudotii 

C. hookeri K. Schum. 

C. jamaicensis (Urb.) Steyerm. 

C. latifolia Standl. 

C. longifolia (Willd.) Baill. =Simira longifolia 

C. longistipulata Bremek. =Chimarrhis microcarpa 

C. microcarpa Standl. 

C. microcarpa Standl. var. microcarpa =Chimarrhis 

microcarpa 

C. microcarpa Standl. var. speciosa Steyerm. =Chimarrhis 

speciosa 

C. paraensis Baill. =Simira paraensis


C. parviflora Standl. 

C. perijaensis Steyerm. =Bathysa perijaensis 

C. pisoniaeformis Baill. =Simira pisoniiformis 

C. pittieri Standl. =Bathysa pittieri 

C. speciosa (Steyerm.) Delprete 

C. turbinata DC. 

C. venezuelensis Standl. & Steyerm. =Dioicodendron dioicum 

C. williamsii Standl. =Chimarrhis hookeri 

Chrysoxylon Casaretto =Plathymenia (Mimosaceae) 

Chrysoxylon Wedd. =Pogonopus 

C. febrifugum Wedd. =Pogonopus tubulosus 

C. speciosum (Jacq.) Kuntze =Pogonopus speciosus var. 

speciosus 

C. tubulosum (A.Rich. in DC.) Kuntze =Pogonopus tubulosus 

Cinchona caroliniana Poir. =Pinckneya bracteata 


C. corymbifera Cav. =Badusa corymbifera 

C. muzonensis Goudot =Ladenbergia muzonensis 

C. philippica Cav. =Badusa corymbifera 

Coffea floribunda Miq., nom. superfl. =Molopanthera 

paniculata var. paniculata 

C. floribunda Martius = Ixora densiflora 

Condaminea DC. 

C. angustifolia Rusby =Condaminea corymbosa 

C. breviflora Standl. =Condaminea corymbosa 

C. corymbosa (R. & P.) DC. 

C. corymbosa (R. & P.) DC. var. pubescens Spruce ex 

K.Schum. =Condaminea corymbosa 

C. corymbosa (R. & P.) DC. var. subsessilis Regel, in herb. 

=Condaminea corymbosa 

C. elegans Delprete 

C. glabrata Bartl. ex DC. =Not Condaminea! 

C. macrophylla Poepp. & Endl. =Capirona decorticans 

C. microcarpa (R. & P.) DC. =Bathysa sp. 

C. petiolata Dwyer =Rondeletia panamensis 

C. tinctoria (Kunth in H.B.K.) DC. =Simira rubescens 

C. utilis Goudot =Elaeagia utilis 

C. venosa (R. & P.) DC. =Not Condaminea! 

Cuatrecasasiodendron Standl. & Steyerm. 

C. colombianum Standl. & Steyerm. 

C. spectabile Steyerm. 

Dioicodendron Steyerm. 

D. cuatrecasasii Steyerm. =Dioicodendron dioicum 

D. dioicum (K.Schum. & K.Krause) Steyerm. 

Dolichodelphys K.Schum. & K.Krause 

D. cholorocrater K.Schum. & K.Krause 

Elaeagia brasiliensis Standl. =Chimarrhis turbinata 

E. miryantha (Standl.) C.M.Taylor & Hammel 

Exandra Standl. =Simira Aubl. 

E. rhodoclada Standl. =Simira rhodoclada 

Exostema corymbifera (Forst.f.) Roem. & Schult. = Badusa 

corymbifera 

E. formosum Cham. & Schlecht. =Rustia formosa 

E. formosum var. laeve Cham. & Schlecht. =Rustia formosa 

E. formosum var. leprosum Cham. & Schlecht. =Rustia 

formosa 

E. leonis Standl. =Acunaeanthus tinifolius 

E. (Pseudostemma) occidentale Benth. =Rustia occidentalis 

E. philippica (Cav.) Roem. & Schult. =Badusa corymbifera 

Flexanthera Rusby =Simira Aubl. 

F. fragrans Rusby =Simira fragrans 

F. subcordata Rusby =Simira cordifolia 

Forsteronia panniculata Casar. (nom. nud.) =Molopanthera 

paniculata var. paniculata 

Henlea Griseb. (Malpighiaceae) 

Henlea H.Karst. =Rustia 

H. muzonensis (Goud.) Klotzsch & H.Karst. ex Walp. 

=Ladenbergia muzonensis 

H. rosea (Goudot) Klotzsch & H.Karst. ex Walp. 


H. splendens H.Karst. =Rustia thibaudioides 

H. thibaudioides H.Karst. =Rustia thibaudioides 

Holtonia Standl. =Elaeagia 

H. myriantha (Standl.) Standl. =Elaeagia miryantha 

Howardia Wedd. =Pogonopus 

H. caracasensis Wedd. =Pogonopus speciosus var. speciosus 

H. febrifuga (Wedd.) Wedd. =Pogonopus tubulosus 

H. grandiflora Wedd. =Pogonopus speciosus var. speciosus 

H. richardii Wedd. =Pogonopus tubulosus 

Ixora L. 

I. densiflora (Martius) Mull. Arg. 

Javorkaea Borhidi & Jarai-Koml. =Rondeletia 

Kajewskiella Merrill & Perry 

K. polyantha M.Jansen 

K. trichantha Merrill & Perry 

Kerianthera J.H.Kirkbr. 

K. preclara J.H.Kirkbr. 

Ladenbergia muzonensis (Goudot) Standl. 

Lindenia Benth. =Augusta subgen. Lindenia 

L. austro-caledonica =Augusta austro-caledonica 

L. rivalis Benth. =Augusta rivalis 

L. vitiensis Seem. =Augusta vitiensis 

Macrocnemum P. Br. 

M. corymbosum R. & P. =Condaminea corymbosa 

M. cubense Griseb. =Picardaea cubensis 

M. exsertus 6rst. =Pogonopus exsertus 

M. glabratum Bartl. =Not Condaminea! 

M. longifolium A. Rich. =Chimarrhis cymosa 

M. microcarpon R. & P. =Bathysa sp. 

M. speciosus Jacq. =Pogonopus speciosus var. speciosus 

M. tinctorium Kunth in H.B.K. =Simira rubescens 

M. tubulosum A.Rich. in DC. =Pogonopus tubulosum 

M. venosum R. & P. =Not Condaminea' 

Mazaea Krug & Urb. 

M. phialanthoides (Griseb.) Krug & Urb. 

M. pungens (Urb.) Alain = Rondeletia sp.? 

M. shaferi (Standl.) Delprete 

M. tinifolia (Griseb.) Alain =Acunaeanthus tinifolius 

Molopanthera Turcz. 

M. burchellii J. D.Hook. =Molopanthera paniculata var. 

burchellii 

M. colombiana Standl., in herb. =Dioicodendron dioicum 

M. paniculata Turcz. 

M. paniculata Turcz. var. paniculata 

M. paniculata Turcz. var. burchellii (J. D.Hook.) K.Schum. 

M. panniculata K.Schum. (hort. var.) =Molopanthera 

paniculata 

Mussaenda bracteata (W.Bartram) Raf. =Pinckneya 

bracteata (W.Bartram) Raf. 

M. speciosa (Jacq.) Poir. =Pogonopus speciosus var. speciosus 

Neobertiera = Pteridocalyx? 

N. gracilis Wernham = Pteridocalyx gracilis (Wernham) 

Delprete, ined.(?) 

Neomazaea Krug & Urb. =Mazaea 

N. phialanthoides (Griseb.) Krug & Urb. =Mazaea 

phialanthoides

Nomenclatural List 219 

N. shaferi Standl. =Mazaea shaferi 

N. pungens Urb. =Rondeletia sp.? 

N. tinifolia (Griseb.) Krug & Urb. =Acunaeanthus tinifolius 

R. brandegeeana Lorence 

R. phialanthoides Griseb. =Mazaea phialanthoides 

R. tinifolia Griseb. =Acunaeanthus tinifolius 

Rustia Klotzsch 

Otocalyx Brandegee =Rondeletia 

0. chiapensis Brandegee =Rondeletia brandegeeana 

Parachimarrhis Ducke 

P. breviloba Ducke 

Picardaea Urb. 

P. cubensis (Griseb.) Britt. ex Urb. 

P. haitiensis Urb. =Picardaea cubensis 

Pinckneya Rich. in A.Michx. 

P. acroma Freire & Allem?o =Simira rubra 

P. bracteata (W.Bartram) Harper =Pinckneya bracteata 


P. bracteata (W.Bartram) Raf. 

P. erubescens Glaz. =Simira glaziovii 

P. ionantha Planch. & Lind. =Pogonopus speciosus var. 

sandwithianus 

P. officinalis Lavalle (nom. nud.) =Pinckneya bracteata 


P. pubens L.C.M. Rich. in A.Mich. =Pinckneya bracteata 

(W. Bartram) Raf. 

P. rubescens Allemao =Simira rubra 

R. alba Delprete 

R. angustifolia K.Schum. 

R. bilsana Delprete 

R. costaricensis (Standl.) Lorence 

R. dressleri Delprete 

R. ferruginea Standl. =Bathysa sp. 

R. formosa (Cham. & Schltdl. ex DC.) Klotzsch 

R. haitiensis Urb. = not Rustia (?) 

R. isernii Standl. =Alibertia isernii 

R. longifolia Standl. =Dolichodelphys chlorocrater 

R. mosenii K.Schum. ex Glaziou =Rustia formosa 

R. occidentalis (Benth.) Hemsl. 

R. panamensis Dwyer =Rustia costaricensis 

R. pauciflora K.Schum. ex Soler. =Tresanthera pauciflora 

R. pohliana Klotzsch =Rustia formosa 

R. rosea (Klotzsch & H.Karst. ex Walp.) K.Schum 


R. rubra Standl. ex D. Simpson 

R. schunkeana Delprete 

R. secundiflora K.Schum. =Rustia thibaudioides 

R. sellowiana Klotzsch =Rustia formosa 

P. viridiflora Allemao & Saldanha =Simira viridiflora R. simpsonii Delprete 

Pinknea Persoon (orth. var.) =Pinckneya 

R. splendens (H.Karst.) Standl. =Rustia thibaudioides 

Pinknea pubescens Person (orth. var.) =Pinckneya R. thibaudioides (H.Karst.) Delprete 

bracteata (W.Bartram) Raf. 

R. venezuelensis Standl. & Steyerm. 

Pinkneya Raf. (orth. var.) =Pinckneya 

R. viridiflora Delprete 

Pogonopus A.Rich. in DC. 

R. warscewicziana Klotzsch, in herb. = Rustia thibaudioides 

P. amarus Hutch., in herb. =Pogonopus tubulosus 

P. bolivianus Britt., in herb. =Pogonopus tubulosus Schizocalyx =Bathysa 

P. erythroxylon (Standl.) J.H.Kirk. =Simira ecuatoriensis S. bracteosus Wedd. =Bathysa bracteosa 

P. febrifugus (Wedd.) J. D.Hook. =Pogonopus tubulosus S. hirsutus Standl. =Bathysa bracteosa 

P. febrifugus (Wedd.) J. D.Hook. var. macrosema Hutch. S. pohliana Berg (unpubl.?) =Faramea sp. 

=Pogonopus tubulosus 

Schreibersia Kuntze =Augusta 

P. exsertus (Orst.) Orst. 

S. longifolia (Spreng.) Kuntze =Augusta longifolia var. 

P. moritzii Klotch., in herb. =Pogonopus speciosus var. longifolia 

speciosus 

Semaphyllanthe L. Andersson =Calycophyllum 

P. ottonis Klotch. =Pogonopus speciosus var. speciosus Sickingia =Simira Aubl. 

P. speciosus (tubulosus) var. ottonis K. Schum., in herb. S. catappiifolia Standl. =Simira macrocrater 

=Pogonopus speciosus var. speciosus 

S. cordifolia J. D.Hook. =Simira cordifolia 

P. speciosus (Jacq.) K.Schum. 

S. ecuadorensis Standl. =Simira ecuadorensis 

P. speciosus (Jacq.) K.Schum. var. speciosus 

S. erythroxylon sensu Oliv. =Simira glaziovii 

P. speciosus (Jacq.) K.Schum. var. sandwithianus Steyerm. S. erythroxylon Willd. =Simira erythroxylon 

P. speciosus (Jacq.) K.Schum. subsp. sandwithianus Steyerm. S. fragrans (Rusby) Standl. =Simira fragrans 

= Pogonopus speciosus var. sandwithianus 

S. goudotii (Baill.) Standl. =Simira goudotii 

P. speciosus (Jacq.) K.Schum. subsp. speciosus var. S. longifolia Willd. =Simira longifolia 

sandwithianus Steyerm. =Pogonopus speciosus var. S. myriantha Standl. =Elaeagia myriantha 

sandwithianus Steyerm. 

S. paraensis (Baill.) K.Schum. =Simira paraensis 

P. speciosus (Jacq.) K.Schum. subsp. speciosus var. S. pisoniiformis (Baill.) K.Schum. =Simira pisoniiformis 

speciosus =Pogonopus speciosus var. speciosus S. tinctoria (Aubl.) Lem6e =Simira tinctoria 

P. speciosus (Jacq.) K. Schum. subsp. exsertus (Orst.) S. tinctoria (Kunth) K.Schum. =Simira rubescens 

Steyerm. =Pogonopus exsertus 

Simira Aubl. 

P. speciosus [tubulosus] (Jacq.) K.Schum. var. panamensis S. catappiifolia (Standl.) Steyerm. =Simira macrocrater 

K.Schum., in herb. =Pogonopus exsertus 

S. cordifolia (J. D.Hook.) Steyerm. 

P. tubulosus (A.Rich. in DC.) K.Schum. 

S. ecuadorensis (Standl.) Steyerm. 

Pseudochimarrhis Ducke =Chimarrhis subgen. S. erythroxylon (Willd.) Bremek. 

Pseudochimarrhis 

S. fragrans (Rusby) Steyerm. 

P. barbata Ducke =Chimarrhis barbata 

S. glaziovii (K.Schum.) Steyerm. 

P. difformis (Benoist) Benoist =Chimarrhis turbinata S. goudotii (Baill.) Steyerm. 

P. turbinata (DC.) Ducke =Chimarrhis turbinata S. longifolia (Willd.) Steyerm. 

S. macrocrater K.Schum. 

Rogiera Planch. =Rondeletia 

Roigella Borhidi & Z. M. Femrnndez =Rondeletia 

Rondeletia L. 

S. myriantha (Standl.) Steyerm. =Elaeagia myriantha 

S. paraensis (Baill.) Steyerm. 

S. pisoniiformis (Baill.) Steyerm.


S. rhodoclada (Standl.) Steyerm. 

S. rubescens (Benth.) Bremek. ex Steyerm. 

S. rubra (Mart.) Steyerm. 

S. tinctoria Aubl. 

S. viridiflora (Allemao & Saldanha) Steyerm. 

Simira Raf. =Ornithogalum L. (Liliaceae) 

Sprucea Benth. =Simira Aubl. 

S. rubescens Benth. =Simira rubescens 

Stomandra Standl. =Rustia 

S. costaricensis Standl. =Rustia costaricensis 

Suberanthus Borhidi & Z. M. Fernmndez =Rondeletia? 

S. pungens (Urb.) Borhidi & Z. M. Fernandez 

=Rondeletia sp.? 

agua fresca (6-8) 

amarelinha (6-12) 

amarilla (6-8) 

anime chiquita (3-1) 

aromuhe (6-7) 

bois chapelle (6-10) 

bois de riviere (6-5) 

bois des bams-jaunes (6-5) 

bois pagaie (6-10) 

bois riviere (6-5) 

bois riviere r6solu (6-5) 

bouganvillea (5-3) 

caapeba (1-11) 

cacahuito (3-1) 

cambur6n (2-lb) 

carapanauba (6-10) 

carapana6ba cinzeiro (6-10) 

carutillo (6-8) 

cascarilla (5-3) 

cascarilla masha (6-6; 1-8) 

cascarillita (1-8) 

caspi blanco (3-1) 

cera (6-3) 

chibigui (5-1) 

chollachaqui caspi blanco (6-7) 

chorcha de gallo (5-1) 

chullachasqui (3-1) 

citronelle (6-10) 

citronelle rouge (6-10) 

coralito (5-2a) 

doncello (1-9) 

flor de hoja (5-3) 

guamo (3-1) 

guapeba branca (1-11) 

INDEX OF LOCAL NAMES 

higueron (6-1) 

hilacho (6-3) 

huacamayo micuna (10-1) 

huacap6 masha (6-7) 

intacchi (6-7) 

itahuba (6-7) 

itauba (6-6, 6-7) 

itauba amarilla (6-6, 6-7) 

jagua amarilla (6-2) 

jatun mincha caspi (6-6) 

jilacho (6-3) 

kaire-e 

(6-10) 

koemaramaraballi 

(6-10) 

larangui (3-1) 

lengua de vaca (3-1) 

llema de huevo (6-1) 

madrofio (5-1) 

man wood (6-1) 

mecha caspi (6-6) 

mincha caspi (6-6) 

mukugd (6-7) 

mun baikuanin (3-1) 

naranjillo montafiero (5-2a) 

pablo manchana (6-6; 6-7) 

palo marfil (5-3) 

palo palillo (6-6) 

palo perro (6-7) 

pampa remo caspi (6-7) 

pampa remocaspi (6-6) 

papelillo caspi (6-7) 

paraka'y (6-10) 

paraku-'iran-'y (6-10) 

pau amarelo (6-12) 

pau de remo (6-10) 

Tresanthera H.Karst. 

T. condamineoides H.Karst. var. condamineoides 

T. condamineoides H.Karst. var. thyrsiflora (Steyerm.) 

Delprete 

T. pauciflora K.Schum. ex Soler. =Tresanthera 

condamineoides var. thyrsiflora 

T. thyrsiflora Steyerm. =Tresanthera condamineoides var. 

thyrsiflora 

Ucriana Spreng. =Augusta 

U. longifolia Spreng. =Augusta longifolia var. longifolia 

Wernhamia S. Moore =Simira Aubl. 

W. boliviensis S. Moore =Simira macrocrater K.Schumann 

pejiballito (6-1) 

penda (6-3) 

pesquim (8- b) 

purma-caspi (6-7) 

quillo-bordon (6-7) 

quillobordon (10-1) 

quina (5-1; 5-2a; 5-3) 

quina blanca (7-1) 

quina do Rio de Janeiro (1-11) 

quina morada (5-2b) 

quinoyer (6-5) 

ramal do pau rosa (6-10) 

river wood (6-5) 

resolu de montagne (6-5) 

resolu de riviere (6-5) 

roble amarillo (6-3) 

sacha cascarilla (1-8) 

sachajagua (6-7) 

sacha quina (5-3) 

saco de mono (8-la) 

samaati-palioudou (6-10) 

santonino (6-9) 

sienjahoedoe (6-10) 

sinjahoedoe (6-10) 

sobrasil (1-11) 

totumo amarillo (6-8) 

tuwara (6-7) 

walalu (6-10) 

waterwood (6-5) 

white calabash (6-8) 

wild fiddle wood (6-4) 

yacu-caspi(6-7) 

yema de huevo (6-1; 6-2) 

yerno prueba (6-6; 6-7)

Index of Scientific Names 221 

INDEX OF SCIENTIFIC NAMES 

Synonyms are in italics. Names in [square brackets] are nomen nuda or unpublished names. Page numbers in 

boldface indicate primary references. Page numbers with an asterisk (*) indicate pages with illustrations or maps. 

Acrobotrys, 8, 14, 22 

Acrosynanthus, 14, 16, 21 

Acunaeanthus, 8, 13, 14, 21 

Aleisanthia, 3, 9, 18 

Alibertia, 96 

isernii, 58, 96 

Allenanthus, 12* 

erythrocarpus, 141, 185 

Angiosperms, 7 

Anthospermeae, 53 

Antirheoideae, 8, 10 

Aparines, 4 

Aparinia, 5 

Arachnothryx, 9, 10, 13 

Ardisia, 80 

Ariadne, 13 

shaferi, 13 

Aristolochia, 121 

Aristolochiaceae, 121 

Augusta, 6, 9, 11, 12*, 13, 14, 16, 18 

subgen. Augusta, 22 

subgen. Lindenia, 16, 22 

longifolia, 16 

Augusta group, 13 

Badusa, 8, 12*, 18 

Bathysa, 6,9,12*, 13,14,21,58,96,105,116,138,186,187 

autralis, 58 

bathysoides, 141, 185 

bracteosa, 13, 20 

cuspidata, 58 

difformis, 140, 169, 175 

perijaensis, 186 

pittieri, 140, 187 

Bathysa group, 13 

Bikkia, 3, 6, 8, 12*, 29, 42, 52, 53, 54 

campanulata, 53 

palauensis, 53 

Blandibractea, 9, 17 

brasiliensis, 17 

Bradea, 17 

Buena muzonensis, 96 

Calycophylleae, 9, 138 

Calycophyllum, 9, 29, 53, 122, 138, 175, 185 

candidissimum, 140 

obovatum, 185 

tubulosum (A. Rich.) DC., 122, 123, 133, 136 

tubulosum Seeman, 123 

Capirona, 53, 137 

decorticans, 116 

erythroxylon, 136, 137 

Carmenocania, 121, 122 

porphyrantha, 121, 130, 131 

Carphalea, 6 

Carphalia, 6 

Cascarilla, 6 

muzonensis, 96 

sect. muzonia, 96 

Catesbaea, 3, 9, 11, 12* 

Catesbaeeae, 3, 5*, 9, 10, 11, 12*, 18, 42, 53 

Catesbaeeae-Chiococceae complex, 3, 16 

Cephalanthia, 5 

Ceratina, 54, 84 

Ceuthocarpus, 3, 8, 12*, 29 

Chalepophyllum, 6, 9, 14, 19, 22 

[Chimarhidia], 18 

[Chimarhis], 137 

[Chimarrheae], 7, 18 

[Chimarrhidoideae], 18 

Chimarrhis, 4,5,6,7, 8, 10, 11, 12*, 14, 16, 17, 18, 19,20, 

23, 24, 25, 26*, 27, 29, 30, 31*, 32, 33, 38, 39*, 

40*,48,50*, 51*, 52,53,54,55,56,137,138,139*, 

140, 141,150, 155, 158, 159, 169, 175, 177, 177*, 

183, 185, 186, 187, 188, 200 

subgen. Chimarrhis, 25, 138, 140, 141, 142 

subgen. Pseudochimarrhis, 25,137,138,140,141,169,175 

barbata, 23, 24, 27, 29, 32, 38, 48, 55, 56, 138, 139*, 

141,142, 159, 175, 176, 177, 177*, 178*, 181 

bathysoides, 141, 185 

brevipes, 24,29,38,40*, 137, 138, 139*, 141,142,177*, 

181,182*, 183, 185 

[corymbosa], 140, 152, 155 

clausicorollata, 185 

cubensis, 23, 24, 32, 38, 39*, 48, 50*, 55, 138, 139*, 

141,147,148*, 149*, 150, 152, 164 

cymosa, 23, 25, 26*, 27, 28, 32, 38, 39*, 48,50*, 53, 56, 

137,138,139*,140,141,147,149*,150,152,153*, 

154,155,166 

subsp. genuina, 140, 152, 155 

subsp.jamaicensis, 140, 141, 150 

subsp. microcarpa, 140, 141, 147, 164 

var.jamaicensis, 150 

var. microcarpa, 147, 164 

[cyrrhosa], 152 

decurrens, 141, 185 

dioica, 140, 141, 185, 187, 192 

duckeana,27,32,38,40*,51*, 138,142,177, 177*, 179, 

180*, 181 

duckei, 141, 169, 175, 179 

ferruginea, 140, 141, 186 

gentryana,24,29,32,38,40*, 137,138,139*, 142,177*, 

183,184*,185 

glabriflora, 23, 24, 26*, 27, 28, 31*, 38, 39*, 54, 55, 56, 

138, 139*, 141,155, 156*, 157*, 158, 159, 162*, 

163, 177* 

goudotii, 141, 186 

hookeri, 23, 24, 27, 38, 39*, 48, 51*, 53,54, 55, 56, 138, 

139*, 140, 141, 142, 157, 158, 159, 160*, 161*, 

162*, 163, 164, 177* 

jamaicensis, 23,24,32,48, 50*, 56, 137, 138, 139*, 141, 

149*,150,151*,152 

latifolia, 23, 24, 32, 38, 39*, 48, 50*, 54, 55, 56, 138, 

139*, 142, 144, 145*, 146* 

longifolia, 186 

longistipulata, 53, 141, 164, 166 

microcarpa, 23, 28, 32, 38, 48, 53, 56, 138, 139*, 140, 

141,144, 164, 165*, 166, 169, 169*, 183 

var. microcarpa, 141, 164 

var. speciosa, 141, 167 

paraensis, 186


Chimarrhis (continued) 

Cubanola, 3, 8, 12*, 29 

parviflora, 22, 28, 32, 28, 39, 48, 50*, 56, 139*, 141, 

142, 143*, 146*, 147, 169 

perijaensis, 186 

pisoniaeformis, 141, 186 

pittieri, 140, 141, 187 

[sessile], 187 

speciosa, 32, 38, 39*, 48, 51*, 141, 164, 167, 168*, 169, 

169*, 183 

Dendrosipanea, 9, 14, 22 

Deppea, 6 

Dilaria, 5 

Dioicodendron, 4, 8, 10, 11, 12*, 14, 18, 20, 23, 24, 25, 

26*, 27, 28, 29, 30, 31*, 32, 33, 38, 41*, 48, 51*, 

55, 140, 141,187, 188, 189*, 190*, 192 

cuatrecasasii, 188, 192 

turbinata, 23,25,26*, 27,29,32, 38,40*, 48,51*, 53, 55, 

56, 138, 139*, 140, 141, 142, 158, 159, 169, 170*, 

171,172*, 174*, 175, 177, 177*, 179, 181, 183 

venezuelensis, 141, 183, 187, 188, 192 

williamsii, 141, 159, 164 

dioicum,26*,31*,41*,48,51*, 140,183,186,187,188, 

189*, 190*, 191* 

Dolichodelphys, 4, 11, 12*, 14, 20, 23, 24, 25, 26*, 27, 

28, 29, 30, 32, 33, 38, 43*, 48, 52*, 55, 191*, 

196, 197, 198* 

Chiococca, 9, 12* 

chlorocrater, 26*, 43*, 48, 52*, 58, 85, 96, 196, 197, 

Chiococceae, 4, 5*, 9, 11, 12*, 13, 16 

198*, 199 

Chione, 12* 

Dolicholobium, 42 

Chrysoxylon Casaretto, 121 

Chrysoxylon Weddell, 121, 122 

Eizia, 9, 17 

febrifugum, 121, 122, 133, 136 

speciosum, 130 

tubulosum, 133 

Cigarilla, 8, 9 

Cinchona, 9, 11, 12*, 53, 105 


philippica 

Cinchona Series, 140 

Elaeagia, 6, 9, 14, 16, 17, 18, 19, 38, 53, 116, 117, 187 

brasiliensis, 169, 175 

cubensis, 19 

grandis, 187 

myriantha, 16 

Endogynie Rafinesque, 5 

Eosanthe, 16 

Ericaceae, 42 

Cinchonaceae, 6 

Erithalis, 9, 13* 

Cinchoneae, 6, 8, 9, 11, 17, 18, 28, 42, 53, 54, 117, 122, Eucondamineae, 3, 6, 18, 140 

138, 140, 175, 193 

Cinchoninae, 7 

Cinchonoideae, 7, 8, 9, 10, 11, 197 

Euglossa, 54, 64* 

Eupinckneya, 122 

Exandra, 16 

Clusia, 80 

rhodoclada 

Coffea, 9, 11 

Exostema, 8, 9, 12*, 53, 88 

floribunda Martius, 193 

australe, 58 

floribunda Miquel, 193, 195 

stipulata, 13 

Coffeae, 9, 11 

Coleoptera, 55, 81 

Condaminea, 4, 6, 8, 9, 10, 11,12*, 13, 14,18, 19,23,24,25, 

cuspidatum, 58 

formosum, 58, 85, 86, 88 

var. laeve, 85, 88, 89 

var. leprosum, 58, 85, 88 

occidentale, 58, 62, 65 

26*, 27, 28, 29, 30, 31*, 32, 33, 34, 36*, 44,47*, 48, souzanum, 58 

53,54,55,57,58,98,103,104*, 105,114,116,117 Exostemma, 58, 88 

angustifolia, 105 

breviflora, 105 

corymbosa, 23, 24, 26*, 31*, 33, 36*, 47*, 48, 53, 54, 

sect. Pseudostemma, 58, 62, 65 

formosum, 58 

occidentale, 25, 58, 62, 65 

55,98,99,103,104*,105,106*,107*,108*, 109*, 

110, 112*, 113, 114, 116,117,140 

var. pubescens, 105, 114 

elegans, 33, 36*, 48, 103, 105, 112*, 114, 115* 

glabrata, 105, 116, 159 

macrophylla, 105, 116 

microcarpa, 105, 116 

petiolata, 105, 116 

tinctoria, 105, 116 

Ferdinandusa, 53 

Ficus, 80 

Flexanthera, 8, 10, 16 

fragrans, 16 

subcordata, 16 

[Forsteronia], 192, 193 

[panniculata], 193 

utilis, 116 

venosa, 105, 117 

Gardenieae, 11, 38, 53, 197 

Geonoma, 80 

Condamineeae, 3, 4, 5*, 6, 7, 8, 9, 10, 11, 12*, 16, 17, 18, Gleasonioideae, 10 

19, 23, 25, 26*, 29, 31*, 38, 42, 52, 54, 122, 138, 

140, 193 

Glionnetia, 3, 9, 18 

Gloneria, 6, 9, 18 

Condamineinae, 3, 4, 6, 7, 10, 11, 18, 42, 140 

Condaminieae, 6, 18 

Couepia, 80 

Coutaportla, 3, 8, 12*, 29 

Coutarea, 3, 8, 9, 12*, 29, 53 

Cuatrecasasiodendron, 9, 12*, 14, 18, 20, 22, 41*, 42 

colombianum, 41* 

Greenea, 6, 9, 18 

Greenia, 6 

Grias, 80 

Guarea, 80 

Guettardeae, 53 

Guettardoideae, 7, 10 

Gyranthera caribensis, 101


[Hamelia albopurpurea], 84 

Hamelieae, 6, 17, 53 

Hedyoteae, 6 

Hedyotideae, 6, 8, 17, 18, 42, 53, 140 

Hedyotidinae, 6 

Henlea Grisebach, 57 

Henlea H. Karsten, 6, 7, 25, 57, 58, 84, 96, 97 


rosea, 58, 96 

splendens, 25, 58, 81, 84, 85 

thibaudioides, 57, 58, 81, 84 

Hillieae, 53 

Hillioideae, 10 

Hintonia, 3, 8, 12*, 29 

Hippotis, 13 

Hippotieae, 13 

Holstianthus, 9, 14, 22 

Holtonia, 16 

Howardia Klotzsch, 121 

Howardia Weddell, 121, 122, 136 

caracasensis, 122, 130 

febrifuga, 56, 121, 122, 133, 136 

grandiflora, 122, 130, 131 

richardi, 122, 133, 136 

Hymenocnemis, 25 

Iriartera fusca, 101 

Isertieae, 8, 9, 10, 18, 53, 197 

Isidorea, 3, 6, 8, 12*, 29 

Ixora, 186, 187, 193 

densiflora, 193 

podocarpa, 186, 187 

Ixoreae, 13, 53 

Ixoroideae, 3, 8, 9, 10, 11 

Javorkaea, 9, 10, 13 

Joosia, 9, 11, 12* 

Kajewskiella, 8, 10, 12*, 18, 26*, 27, 42, 43*, 48, 52* 

polyantha, 48, 52* 

trichantha, 26*, 42,43* 

Kerianthera, 8, 10, 12*, 18, 26*, 27, 31*, 41*, 49* 

preclara, 26*, 31*, 41*, 42, 48, 49* 

Ladenbergia, 6, 58 

muzonensis, 58, 96 

Larospermia, 5 

Lerchea, 18 

Limnosipanea, 9, 14, 17, 19 

schomburgkii, 19 

Lindenia, 6, 9, 11, 12*, 13, 16, 18 

Mabea, 80 

Macbrideina, 9, 14, 21, 117 

Machaonia, 13 

Macrocnemum P. Browne, 32, 105, 117, 140, 155 

Macrocnemum sensu A. Richard, 140 

candidissimum, 140 

corymbosum, 105, 109*, 113, 140 

cubense, 118, 120, 121 

exsertum, 122, 123 

glabratum, 116 

jamaicense, 140 

longifolium, 140, 152, 155 

microcarpon, 116 

speciosus, 122, 130, 131 

tetrandrum, 140 

tinctorium, 105, 116 

tubulosum, 122, 133, 136 

venosum, 117 

Maguireothamnus, 9, 14, 22 

Malanea grandis, 187 

Malpighiaceae, 57 

Mazaea, 4, 10, 13, 14, 19, 21 

shaferi, 13 

Mazaea group, 13 

Melastomataceae, 30 

Melipona, 54, 64*, 84 

Mimosaceae, 121 

Molopanthera,4, 9, 11, 12*, 14, 18, 20,23,24,25,26*, 27, 

28,29,30,31*,32,33,38,41*,55,192,193 

burchellii, 195 

[colombiana], 192 

paniculata, 26*, 32, 41*, 192, 193, 194*, 195, 196* 

var. paniculata, 26*, 31*, 194*, 195, 196* 

var. burchellii, 41*, 195, 196* 

panniculata, 193, 195 

var. [scabrida], 196 

Morierina, 6, 8, 12*, 53 

montana, 53 

Morindeae, 53 

Mussaenda, 9, 18 

Mussaenda speciosa, 130 

Mussaendeae, 53, 122 

Mussaendopsis, 53 

Naucleeae, 4, 53 

Neblinathamnus, 9, 14, 22 

Neobertiera, 9, 10, 17 

Neomazaea, 13 

Nerstia, 3, 8, 9, 12*, 29 

Ochroma, 147 

Onagraceae, 42 

Osa, 3, 8, 29 

Otocalyx, 13 

Paederieae, 53 

Pallasia, 6 

Parachimarrhis, 4, 8, 10, 12*, 14, 19, 21, 28, 29, 30, 32, 33, 

38,42,43*, 48,51*, 55,56,199,200,201*, 202* 

breviloba, 43*, 48, 51*, 199, 200, 201*, 202*, 202 

Phialanthus, 13, 16 

Phitopis, 7, 13, 14, 22 

Phyllacanthus, 3, 11, 12* 

Phyllomelia, 4, 13, 16, 19 

Picardaea, 4, 8, 10, 11, 12*, 14, 19,20,23,24,25,26*,27, 

28, 29, 30, 31*, 32, 33, 37*, 47*, 48, 52, 55, 117, 

118, 120 

cubensis,26*,31*,37*,47*,48,117,118,119*, 120*, 121 

haitiensis, 117, 118, 120, 121 

Pinckneya L.C.M. Richard, 3, 6, 8, 9, 10, 11, 12*, 14, 17, 

18, 19, 25, 26*, 27, 29, 30, 31*, 32, 37*, 41, 48, 

49*, 52, 53, 54, 55, 121,122 

Pinckneya sensu Saldanha & Allemao, 7 

bracteata (W. Bartram) Rafinesque, 18,23,26*, 31*, 37*, 

49*, 53 

erubescens, 17 

ionantha, 131, 132, 133 

pubens, 53 

rubescens, 17


Pinckneya sensu Saldanha & Allemao (continued) 

viridiflora, 17 

Pinckneyinae, 3, 4, 6, 11, 18, 122 

Pinckneyeae, 3, 18 

Pinkneya, 6 

Placocarpa, 12* 

Plathymenia, 121 

Pogonopus, 4,6,8,9, 10, 11, 12*, 14, 19,23,24,25,26*, 27, 

28,29,30,31*,32, 33,34,37*,38,42, 48,49*, 52, 

53,54,55,56, 117,121,122,123, 131,136, 137 

[amarus], 136 

[bolivianus], 136 

caracasensis, 130 

erythroxylon, 136 

febrifugus, 133, 136 

var. macrosema, 133, 136 

exsertus, 30, 37*, 38, 48, 49*, 55, 56, 122, 123, 124*, 

125*, 126, 127 

[moritzii], 131 

ottonis, 121, 122, 130 

speciosus, 23, 24, 36*, 30, 31*, 37*, 38, 48, 49*, 53, 54, 

55,56,121,122,123,125*, 126,127,128*, 129,133 

subsp. exsertus, 122, 123 

subsp. sandwithianus, 122 

subsp. speciosus, 122, 132, 133 

var. sandwithianus, 122, 132 

var. speciosus, 130 

var. speciosus, 37*, 38, 121, 125*, 127, 128*, 130, 

131,133 

var. sandwithianus, 23, 24,26*, 31*, 37*, 38,48,49*, 

54, 55, 125*, 127, 128*, 129*, 130, 131,132 

[var. ottonis], 131 

[var. panamensis], 127 

tubulosus, 30, 37*, 38, 48, 49*, 55, 56, 121, 122, 123, 

125*, 126, 133, 134*, 136 

[var. ottonis], 131 

[var. panamensis], 127 

[var. boliviensis], 136 

Polarnia, 5 

Polyspia, 5 

Pomazota, 18 

Pomazotoideae, 10 

Portlandia, 3, 6, 8, 9, 12*, 29, 53, 54, 193 

grandiflora, 53 

Portlandia-group, 9 

Portlandia series, 6 

Portlandieae, 6, 10 

Portlandiinae, 3, 6, 8, 9, 10, 11, 18, 29, 42 

Pseudochimarrhis, 53, 137, 140, 141, 169, 175 

barbata, 140, 141, 175, 176 

difformis, 140, 141, 169 

turbinata, 137, 140, 141, 169 

Pseudomussaenda, 8, 10, 18 

Psychotria, 6, 9, 18 

Psychotrieae, 18, 53 

Pteridocalyx, 9, 14, 17, 20 

Raritebe, 197 

Retiniphyllum, 197 

Rogiera, 9, 10, 13 

Roigella, 9, 13 

Rondeletia, 4, 6, 9, 10, 12*, 13, 18, 42, 116 

brandegeeana, 13 

panamensis, 105, 116 

Rondeletia-complex, 13, 14, 20, 22 

Rondeletieae, 3,4,5*, 6, 7,8,9, 10, 11, 12*, 13, 16, 17, 18, 

19, 29,38, 42, 53, 54, 116, 117, 138, 140, 197 

Rondeletiinae, 6, 18, 140 

Rubiaceae, 3, 4, 5, 6, 7, 8, 9, 10, 24, 25, 27, 29, 30, 32, 33, 

52,53,54,56,57, 97, 116, 117,138, 150, 187, 193 

Rubieae, 53 

Rubioideae, 3, 7, 8, 9, 10, 11 

Rustia, 3,4,6,7,8,10,11,12*, 14,16,19,20,23,24,25,26*, 

27, 28, 29, 30, 31*, 32, 33, 34*, 35*, 38,42,44,45*, 

46*,53,54,55,56,57,58,59,78,84,85,88,89,91, 

96,97,98,99, 103,114,116,117,141,186, 199 

alba, 23, 24, 26*, 27,28, 30,33, 35*, 44,45*, 57, 58, 59, 

67, 68*, 69*, 70, 74*, 88 

angustifolia, 24, 25, 28, 30, 33, 35*, 44, 46*, 58, 59, 91, 

92, 92*, 93*, 95 

bilsana, 23, 24, 28, 29, 30, 54, 57, 58, 70, 71*, 72*, 74* 

costaricensis, 23, 28, 30, 31*, 33, 34*, 44, 45*, 54, 57, 

58,59,60*, 61*, 62* 

dressleri, 23, 24,28,29,30,44,54,57,58,59,61*, 65,66* 

ferruginea, 58, 96, 140, 186 

formosa, 23,24,25,26*, 27,28,30,33, 34*, 44,46*, 53, 

57,58,59,70,85, 87*, 88, 89, 91, 92* 

gracilis, 25, 28, 30, 33, 34*, 44, 46*, 57, 58, 59, 89, 91, 

92, 92*, 94*, 95 

haitiensis, 58, 95 

isernii, 58, 96 

longifolia, 58, 85, 96, 197, 199 

mosenii, 86 

occidentalis, 23, 24,25, 26*, 27,28,29,30,31 *, 33, 34*, 

44, 45*, 54, 55, 57, 58, 59, 62, 63*, 64*, 65, 70, 

74*,98 

panamensis, 58, 59, 62 

pauciflora, 25, 96, 99 

pohliana, 58, 86, 88 

rosea, 58, 96 

rubra, 23, 28, 30, 33, 34*, 44, 45*, 54, 57, 59, 72, 74*, 

75*, 76, 80 

schunkeana, 28, 30, 33, 35*, 44, 45*, 54, 55, 57, 58, 59, 

74*,76,78,79*,80,84 

secundiflora, 58, 81, 84 

sellowiana, 58, 85, 88 

simpsonii, 24, 28, 30, 32, 44, 46*, 57, 59, 88, 89, 90*, 

91,92*, 95 

splendens, 76, 81, 82, 84, 199 

thibaudioides, 23, 24, 25, 28, 30, 31 *, 33, 35*, 44,45*, 

54, 55, 57, 58, 59, 74*,76,80,81,82*,83*,84, 

85, 199 

venezuelensis, 23, 24, 28, 33, 35*, 58, 59, 74*, 85 

[veraguensis], 62 

viridiflora, 23, 28, 30, 33, 35*, 44, 46*, 54, 57, 58, 59, 

74*, 76, 77*, 78, 80 

[warscewicziana], 84, 85 

Sambucidia, 5 

Schizocalyx, 13 

Schizolobium, 147 

Schizomussaenda, 18 

Schmidtottia, 3, 8, 12*, 16, 29 

Scolosanthus, 13* 

Sickingia, 7, 16, 105, 117, 140, 141, 186 

ecuadorensis, 137 

erythroxylon, 16, 136 

goudotii, 186 

longifolia, 16, 186 

myriantha, 16


paraensis, 186 

pisoniiformis, 186, 187 

rubra, 17 

tinctoria (Kunth) K. Schumann, 116 


Siemensia, 3, 8, 29 

SimiraAublet, 7,9,10,12*, 14,16,17,18,19,21,38,42,43*, 

105,117,137,138,140,141,185,186,200 

catappiifolia, 17 

cordifolia, 16 

ecuadorensis, 136, 137 

erythroxylon, 136 

fragrans, 16 

glaziovii, 17 

goudotii, 186 

longifolia, 43*, 186 

macrocrater, 17, 20 

paraensis, 186 

pisoniiformis, 186 

rhodoclada, 16 

rubescens, 105, 116 

rubra, 17 

tinctoria Aublet, 116, 185 


Simira group, 16, 17 

Simireae, 5*, 10, 16, 18 

Sipanea, 4, 9, 14, 17, 18, 19 

biflora, 19 

Sipanea group, 17, 19 

Sipaneeae, 4, 5*, 7, 9, 10, 11, 13, 17, 18 

Sipaneopsis, 8, 9, 14, 17, 19, 20, 21 

Solanaceae, 30 

Spathichlamys, 3, 9, 18 

Sphanidia, 5 

Sprucea, 140 

Standleya, 17 

Steenisia, 9, 18 

Stevensia, 9, 14, 16, 21 

Steyermarkia, 9, 14, 17, 19 

Stomandra, 8, 10, 57, 58 

costaricensis, 57, 58, 59, 62 

Suberanthus, 13 

Synisoon, 197 

Syringantha, 9 

Thogsennia, 3, 8, 12*, 29 

Tocoyena, 197 

Tresanthera, 3, 4,6, 7, 8, 10, 11,12*, 14,19,20,23,24,25, 

26*, 27, 28, 29, 30, 31*, 33, 36*, 42, 44, 47*, 54, 

55, 57, 58, 96, 97, 98, 99 

condamineoides, 25, 26*, 28, 31*, 33, 36*, 44, 47*, 58, 

96, 97, 98, 99, 100*, 101, 103, 199 

var. condamineoides, 26*, 28, 31 *, 33, 36*, 44, 47*, 

96, 99,100*, 101 

var. thyrsiflora, 28, 33, 36*, 44, 47*, 99, 101, 102* 

pauciflora, 25, 97, 98, 99 

thyrsiflora, 97, 99, 101 

Trigona, 54, 84 

Urophylloideae, 10 

Vanguerieae, 53 

Viburidia, 5 

Warszewiczia, 6, 9, 14, 20, 53, 116, 187 

schwackei, 20 

Wendlandia, 3, 9, 18 

Wernhamia, 9, 17 

boliviensis, 17 

Wittmackanthus, 6 

Xanthophytum, 18, 42

Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora

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