Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora
Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora
Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora
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Organization for Flora Neotropica<br />
<strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>): <strong>Part</strong> I (<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea, Pogonopus,<br />
Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and Parachimarrhis)<br />
Author(s): Piero G. Delprete<br />
Reviewed work(s):<br />
Source: Flora Neotropica, Vol. 77, <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>): <strong>Part</strong> I (<strong>Rustia</strong>, <strong>Tresanthera</strong>,<br />
Condaminea, Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys,<br />
and Parachimarrhis) (Jun. 25, 1999), pp. 1-225<br />
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica<br />
Stable URL: http://www.jstor.org/stable/4393887 .<br />
Accessed: 17/11/2011 11:32<br />
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FLORA NEOTROPICA MONOGRAPH 77<br />
RONDELETIEAE (RUBIACEAE)-PART I<br />
(<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea, Pogonopus,<br />
Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and<br />
Parachimarrhis)<br />
PIERO G. DELPRETE<br />
Publishedfor<br />
The Organization for Flora Neotropica<br />
by<br />
The New York Botanical Garden Press<br />
Bronx, New York<br />
Issued 25 June 1999
Flora Neotropica<br />
Published for<br />
Organization for Flora Neotropica<br />
by<br />
The New York Botanical Garden Press<br />
Executive Board<br />
Ariane Luna Peixoto, President, UFRRJ, Caixa Postal 74.582, 23851-970 Seropedica, Itaguai,<br />
RJ, Brazil<br />
Ren&e Fortunato, Vice President, Instituto de Recursos Biologicos, I.N.T.A., 1712 Castelar, Buenos<br />
Aires, Argentina<br />
Wm. Wayt Thomas, Executive Director, Ex Officio, Institute of Systematic Botany, The New York<br />
Botanical Garden, Bronx, NY 10458-5126, U.S.A.<br />
Dieter Wasshausen, Secretary, Department of Botany - NHB- 166, Smithsonian Institution, Washington,<br />
DC 20560-0001, U.S.A.<br />
Paul J. M. Maas, Treasurer, Institute of Systematic Botany, State University of Utrecth, Postbus<br />
80.102, 3508 TC Utrecht, The Netherlands<br />
Paul Berry, Botany Department, 132 Birge Hall, 430 Lincoln Drive, University of Wisconsin,<br />
Madison, WI 53706-1381, U.S.A.<br />
Mireya D. Correa A., Departamento de Botanica, Facultad de Ciencias Naturales, Exactas y<br />
Tecnologia, Universidad de Panama, Estafeta Universitaria, Panama, Panama<br />
Jose Rubens Pirani, Departamento de Botanica, Instituto de Biociencias, Universidade de Sao Paulo,<br />
Caixa Postal 11461, 05422-970 Sao Paulo SP, Brazil<br />
Ghillean T. Prance, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K.<br />
Rodolphe Spichiger, Conservatoire et Jardin Botaniques, de la Ville de Geneve, Case Postale 60,<br />
CH-1292 Chambesy/GE, Switzerland<br />
Claudio Delgadillo M. (1996-2001)<br />
Laurence J. Dorr (1996-2001)<br />
Jose R. Pirani (1996-2001)<br />
Editorial Committee<br />
James L. Luteyn, Editor<br />
S. Rob Gradstein, Editor<br />
Julio A. Lombardi (1999-2004)<br />
Michael O. Dillon (1999-2004)<br />
Michael Nee (1999-2004)<br />
ORGANIZATION FOR FLORA NEOTROPICA<br />
Founded and Conducted under the Auspices of UNESCO
<strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)-<strong>Part</strong> I
<strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)-<strong>Part</strong> I<br />
(<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea, Pogonopus,<br />
Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys, and<br />
Parachimarrhis)<br />
PIERO G. DELPRETE<br />
CONTENTS<br />
A bstract/R esum en ........................................................................................................ 2<br />
Introduction to the <strong>Rondeletieae</strong> ............................. ............................... 3<br />
H istorical Sum m ary .................................................................................................. 4<br />
Phylogenetic Relationships ........................................ ................... 10<br />
Current Circumscription of the <strong>Rondeletieae</strong>, Including Notes on<br />
Genera Newly Included, Excluded, or Synonymized ........................................... 11<br />
Key to the Genera of Neotropical <strong>Rondeletieae</strong> ...................................................... 19<br />
Notes on Descriptions, Abbreviations, and Acronyms ............................................ 23<br />
Introduction to the Neotropical Genera Studied .......................................................... 23<br />
General Morphology .............................................................................................. 23<br />
Seed Morphology .......... ............................................................................. 33<br />
Pollen M orphology ................................................................................................... 42<br />
Wood Anatomy .......... .................................................... ................... 52<br />
Presence of Calcium Oxalate Crystals ..................................................................... 54<br />
K aryology ................................................................................................................. 54<br />
Pollination and Other Plant/Animal Interactions ..................................................... 54<br />
H abitat and Ecology ................................................................................................. 55<br />
U ses .......................................................................................................................... 55<br />
Species Concepts ............................................................. ................... 56<br />
Systematic Treatment of the Genera Studied ............................................................... 56<br />
1. <strong>Rustia</strong> . ................................................................................... 56<br />
2. <strong>Tresanthera</strong> ....... ............................................................................................ 96<br />
3. C on dam inea .......................................................................................................... 103<br />
4 . P icarda ea ............................................................................................................. 117<br />
5. Pogonopus ............................................................... .................. 121<br />
6 . C h im arrh is............................................................................................................ 13 7<br />
7. D ioicodendron ...................................................................................................... 187<br />
8. Molopanthera .............................................................. .................. 192<br />
9. D olichodelp hys ..................................................................................................... 196<br />
10. Parachimarrhis ...... ....................................................................................... 199<br />
A cknow ledgm ents ........................................................................................................ 203<br />
Literature Cited ......... ......................................................................... 203<br />
N um erical List of Taxa ................................................................................................ 209<br />
List of Exsiccatae .......... .................................................... ................... 210<br />
Nomenclatural List .............................................................. ................... 217<br />
Index of Local Names .................................. ........................... 220<br />
Index of Scientific Names ............................................................ 221
2 Flora Neotropica<br />
ABSTRACT<br />
Delprete, P. G. (Institute of Systematic Botany, The New York Botanical Garden, Bronx,<br />
NY 10458-5126, U.S.A.). <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)-<strong>Part</strong> I (<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea,<br />
Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera, Dolichodelphys,<br />
and Parachimarrhis). Flora Neotropica Monograph 77: 1-226. 1999.-The tribe<br />
<strong>Rondeletieae</strong>, as here circumscribed, is a predominantly neotropical group of trees, shrubs,<br />
and rarely herbs. In the Neotropics it is represented by approximately 42 genera and 340<br />
species. The present volume is the first part of a series in which the monographic treatment<br />
of the neotropical <strong>Rondeletieae</strong> is planned. For this reason, a general definition and<br />
a taxonomic survey of this group, as well as a list of all the genera, their approximate<br />
number of species and their distribution, and a key to all the genera are included.<br />
This study originated as a revision of the tribe Condamineeae (sensu Hooker), a group<br />
of predominantly neotropical shrubs and trees (except Pinckneya from North America,<br />
and Bikkia and Badusa from the South Pacific), usually delimited by having valvate corollas<br />
and many-seeded capsules. The Condamineeae were subdivided by Hooker into three<br />
subtribes: Condamineinae, Portlandiinae, and Pinckneyinae. As a result of phylogenetic<br />
analyses using morphological characters, the Condamineeae (sensu Hooker) has been<br />
shown to be paraphyletic. The genera of the Portlandiinae have been transferred to the<br />
newly amended tribe Catesbaeeae sensu Delprete, and genera of the Condamineinae and<br />
Pinckneyinae transferred to the <strong>Rondeletieae</strong>, resulting in a complete dismantling of the<br />
Condamineeae. The Sipaneeae (Sipanea, Sipanaeopsis, Limnosipanea, and Neobertiera)<br />
are included in a broad delimitation of the <strong>Rondeletieae</strong> sensu Delprete. Additionally,<br />
Phyllomelia (with bi-seeded indehiscent fruits, i.e., pseudosamaras), recently treated as<br />
genus incertae sedis by Robbrecht, is also included in the <strong>Rondeletieae</strong> s.l., and believed<br />
to be related to Mazaea.<br />
The neotropical genera treated in the present monograph are those of the subtribes<br />
Condamineinae and Pinckneyinae, now included in the <strong>Rondeletieae</strong>, plus a few related<br />
genera: <strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea, Pogonopus, Chimarrhis,<br />
Dioicodendron, Parachimarrhis, Molopanthera, and Dolichodelphys. Forty-six taxa, 40<br />
species, and 6 varieties, are recognized in the 10 genera listed above.<br />
RESUMEN<br />
Delprete, P. G. (Institute of Systematic Botany, The New York Botanical Garden, Bronx,<br />
NY 10458-5126, U.S.A.). <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)-<strong>Part</strong> I (<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea,<br />
Picardaea, Pogonopus, Chimarrhis, Dioicodendron, Molopanthera,<br />
Dolichodelphys, and Parachimarrhis). Flora Neotropica Monograph 77: 1-226. 1999.-<br />
La tribu <strong>Rondeletieae</strong>, como esta aqui circunscrita, es un grupo predominantemente neotropical<br />
de arboles, arbustos y raramente hierbas. En el Neotr6pico la tribu esta representada<br />
por aproximadamente 42 generos y 340 especies. El presente volumen es la primera<br />
parte de una serie en la cual se esta programando el tratamiento monografico de los generos<br />
neotropicales de las <strong>Rondeletieae</strong>. Por lo tanto, se incluyen aqui una descripci6n de la<br />
tribu, un listado de todos los generos, con numero aproximado de especies y su distribuci6n<br />
geografica, y una clave para todos los generos neotropicales de este grupo.<br />
El presente estudio se origino a partir de una revision de la tribu Condamineeae (sensu<br />
Hooker), un grupo predominantemente neotropical (a excepci6n de Pinckneya, endemico<br />
de Norte America, y Bikkia y Badusa endemicos del Pacifico Sur) de arboles y arbustos,<br />
usualmente caracterizado por corolas valvadas y capsulas con muchas semillas. Las Condamineeae<br />
fueron divididas por Hooker en tres subtribus: Condamineinae, Portlandiinae<br />
y Pinckneyinae. Como resultado de un analisis filogenetico usando datos morfologicos,<br />
se demostro que la tribu Condamineeae (sensu Hooker) es un grupo parafiletico. Los<br />
generos de la subtribu Portlandiinae fueron transferidos a la tribu Catesbaeeae, recientemente<br />
enmendada por Delprete, y los generos de las Condamineinae y Pinckneyinae fueron
Introduction to the <strong>Rondeletieae</strong> 3<br />
transferidos a las <strong>Rondeletieae</strong>, causando asi el completo desmembramiento de la tribu<br />
Condamineeae. La tribu Sipaneeae (Sipanea, Sipanaeopsis, Limnosipanea and Neobertiera)<br />
esta tambien incluida en una mas amplia delimitacion de las <strong>Rondeletieae</strong> sensu Delprete.<br />
Adicionalmente, Phyllomelia (con frutos indehiscentes y con dos semillas, i.e., pseudosamaras),<br />
recentemente tratado como genus incertae sedis por Robbrecht, es tambien<br />
incluido en las <strong>Rondeletieae</strong> s.l. y certamente relacionado con Mazaea.<br />
Los generos neotropicales tratados en la presente monografia, tradicionalmente incluidos<br />
en las Condamineineae y Pinckneyinae y ahora transferidos a las <strong>Rondeletieae</strong> (a los cuales<br />
se afiaden unos generos relacionados), son: <strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea,<br />
Pogonopus, Chimarrhis, Dioicodendron, Parachimarrhis, Molopanthera y Dolichodelphys.<br />
Cuarenta y seis taxa (40 especies y 6 variedades) son descritos en los 10 generos<br />
anteriormente listados.<br />
INTRODUCTION TO<br />
THE RONDELETIEAE<br />
The <strong>Rubiaceae</strong> is the fourth largest family of flow-<br />
ering plants, with approximately 650 genera and<br />
12,000 species, with mostly pantropical distribution<br />
and predominantly woody. In the Neotropics it is the<br />
largest dicotyledonous family, with approximately<br />
200 genera and 5000 species. Andersson (1992) cited<br />
about 4500 species, but considering the large amount<br />
of new taxa recently described from material collected<br />
in newly explored neotropical areas, his estimate of<br />
the neotropical species is, in my opinion, too low.<br />
This family is easily recognized by its opposite<br />
(rarely whorled, or exceptionally alternate) leaves<br />
with entire margins, interpetiolar (rarely intrapetiolar)<br />
stipules with colleters on the adaxial side, gamopetalous<br />
corollas, and inferior ovaries (exceptionally halfsuperior<br />
or superior). The latest comprehensive family<br />
classification was proposed by Robbrecht (1988,<br />
1993), where he recognized four subfamilies and<br />
44 tribes. Recent phylogenetic studies based on molecular<br />
data (Bremer et al., 1995; Rova et al., 1997)<br />
seem to indicate that the family should be divided<br />
into three subfamilies: Cinchonoideae, Ixoroideae,<br />
and Rubioideae.<br />
The <strong>Rondeletieae</strong> (subfam. Cinchonoideae), as<br />
here circumscribed, is a group of predominantly neotropical<br />
(except Pinckneya Rich. from North America;<br />
and Aleisanthia Ridl., Glionnetia Tirveng., Spathichlamys<br />
R. Parker, and Wendlandia Bartl. ex DC.<br />
being paleotropical) trees, shrubs, and rarely herbs,<br />
characterized by corollas valvate, contorted or imbricate,<br />
stamens variably inserted at basal, medial, or<br />
distal positions on the corolla tube, anthers dehiscing<br />
longitudinally orporicidally (<strong>Rustia</strong> and <strong>Tresanthera</strong>),<br />
and few- to many-seeded capsules or two-seeded<br />
pseudosamaras.<br />
This study originated as a revision of the Conda-<br />
mineeae (sensu Hooker, 1873) (Delprete, 1996f), a<br />
group of predominantly neotropical shrubs and trees,<br />
usually delimited by having valvate corollas and<br />
many-seeded capsules (Hooker, 1873). The tribe Con-<br />
damineeae was founded by Hooker (1873) and sub-<br />
divided by him into three subtribes: Condamineinae<br />
(as "Eucondamineae"), Portlandiinae (as "Port-<br />
landieae"), and Pinckneyinae (as "Pinckneyeae").<br />
This group has been shown to be paraphyletic<br />
(Delprete, 1995a, 1996d, 1996f; also see Phylogenetic<br />
Relationships in this volume).<br />
Because of their corolla aestivation (imbricate<br />
lobes with reduplicate tubes) and their base chromo-<br />
some number (x = 12, large chromosomes; Kiehn,<br />
1995, and pers. comm.), the genera of the Portlandiinae<br />
(Bikkia Reinw., Catesbaea L., Ceuthocarpus<br />
Aiello, Coutaportla Urb., Coutarea Aubl., Cubanola<br />
Aiello, Hintonia Bullock, Isidorea A. Rich., Nernstia<br />
Urb., Osa Aiello, Phyllacanthus J. D. Hook., Portlandia<br />
P. Browne, Schmidtottia Urb., Siemensia Urb.,<br />
and Thogsennia Aiello) have been included in the<br />
newly emended Catesbaeeae (Delprete, 1995a,<br />
1996d). Ongoing phylogenetic studies based on morphological<br />
and molecular data will definitely clarify<br />
the relationships within the Catesbaeeae-Chiococceae<br />
complex (Delprete et al., in progress).<br />
Based on previous phylogenetic analyses<br />
(Delprete, 1995a, 1996d, 1996f), the results of which<br />
are summarized below (see Phylogenetic Relationships),<br />
the subtribes Condamineinae and Pinckneyinae<br />
(sensu Hooker, 1873) were transferred to the tribe<br />
<strong>Rondeletieae</strong> (sensu Delprete). The inclusion of the<br />
two subtribes in the <strong>Rondeletieae</strong> is supported by their
4 Flora Neotropica<br />
base chromosome number (x = 11, small chromosomes;<br />
Kiehn, 1995, and pers. comm.), many-seeded<br />
capsules, and variable corolla aestivation. The monophyly<br />
of the <strong>Rondeletieae</strong> s.l. is yet to be tested (Rova<br />
et al., in progress). The inclusion of the Condamineeae<br />
(pro parte) in the <strong>Rondeletieae</strong> implies that the valvate<br />
aestivation is not a sufficient character for tribal<br />
delimitation. In addition, the minute flowers of<br />
Chimarrhis (usually reported as valvate), Molopanthera,<br />
Dolichodelphys, and Parachimarrhis have imbricate<br />
aestivation.<br />
The tribe Sipaneeae (sensu Robbrecht, 1988, 1993)<br />
is also here included in the wide circumscription of<br />
the <strong>Rondeletieae</strong>, where they are exceptional only in<br />
their herbaceous habit. Of this group, only Sipanea<br />
Aubl. has been the subject of chromosome studies, for<br />
which Kiehn (1995) reports: "In Sipanea, one diploid<br />
and one tetraploid taxon on a basic number of x = 11<br />
[small chromosomes, pers. comm.] are known. While<br />
chromosomes ofmitotic metaphase are not elongated<br />
as in <strong>Rondeletieae</strong> but rather compact as in Naucleeae,<br />
interphase nuclear structures of Sipanea resemble<br />
those of both <strong>Rondeletieae</strong> and Naucleeae."<br />
Additionally, Phyllomelia (monotypic, endemic to<br />
The present volume is the first part of a planned<br />
series of monographic treatments of neotropical <strong>Rondeletieae</strong>,<br />
as presently circumscribed. Being an introductory<br />
volume, the section Historical Summary introduces<br />
the complex nomenclatural history and<br />
delimitation of the <strong>Rondeletieae</strong> as currently recognized.<br />
At the end of this section, a description of this<br />
tribe is presented, followed by additional notes on the<br />
genera here included in (or previously referred to) the<br />
<strong>Rondeletieae</strong>. A table summarizes all the neotropical<br />
genera of this group, with their approximate number<br />
of species, habits, distributions, habitats, and elevation<br />
ranges. Finally, a Key to Neotropical <strong>Rondeletieae</strong><br />
is presented.<br />
The genera treated in the Systematic Treatment of<br />
this volume are those of the subtribes Condamineinae<br />
and Pinckneyinae, previously treated as part of the<br />
Condamineeae and now included in the <strong>Rondeletieae</strong>,<br />
plus a few genera related to them. An overview of<br />
morphological, anatomical, palynological, and ecological<br />
studies of the genera is also presented. The<br />
Systematic Treatment includes monographs of <strong>Rustia</strong>,<br />
<strong>Tresanthera</strong>, Condaminea, Picardaea, Pogonopus,<br />
Chimarrhis, Dioicodendron, Parachimarrhis, Molop-<br />
anthera, and Dolichodelphys. Finally, the Nomencla-<br />
tural List gives all the accepted taxa, orthographic<br />
variants, and synonyms, as recognized by the author.<br />
HISTORICAL SUMMARY<br />
<strong>Rubiaceae</strong> classification has undergone a series of<br />
rearrangements as scientific knowledge and technology<br />
became more sophisticated. The taxonomic history<br />
of this family, with emphasis on the tribe <strong>Rondeletieae</strong><br />
s.l., is here divided into six time periods.<br />
During the Linnean period (1753-1789) only a few<br />
genera and species were known. The post-Linnean<br />
period (1789-1828) is characterized by the exploration<br />
of tropical areas and the discovery of many new<br />
taxa; Jussieu during this period proposed the first<br />
<strong>Rubiaceae</strong> classification, using locule number and<br />
number of ovules per locule as main characters. The<br />
Candollean period (1829-1872) started with Achille<br />
Richard, who in 1829 presented the first monographic<br />
work dedicated to the whole family. Richard's system<br />
highly influenced the classification proposed by<br />
A. P. de Candolle, which remained the standard reference<br />
until the end of the 1800s. The late 19th cen-<br />
Cuba) with two-seeded pseudosamaras (Spjut, 1994), tury (1873-1900) is marked by the innovative work<br />
is also included in the <strong>Rondeletieae</strong> s.l., and is believed of J. D. Hooker, the first rubiologist who followed<br />
to be related to Mazaea (see notes after <strong>Rondeletieae</strong>, evolutionary theories. Hooker's classification influbelow).<br />
enced the work of the other two great scholars of this<br />
period: Baillon and Schumann. The modem period<br />
(1900-1990) is punctuated by the milestone works of<br />
Bremekamp (1934, 1940, 1954, 1966), Verdcourt<br />
(1958), and Robbrecht (1988). Their classifications<br />
attempted to break away from phenetic systems; in<br />
constructing them they evaluated character evolution,<br />
using total evidence (i.e., morphology, anatomy, cytology,<br />
biogeography, etc.) and defined groups using<br />
sets of characters (vs. single character classifications).<br />
During the last decade of the 20th century (after 1990)<br />
<strong>Rubiaceae</strong> phylogenetic studies evolved quite rapidly,<br />
being influenced by the advent of cladistic methods<br />
and by the science of molecular biology. The various<br />
delimitations of the <strong>Rondeletieae</strong>, Condamineeae,<br />
Sipaneeae, Catesbaeeae, and Chiococceae are summarized<br />
in Figure 1.<br />
Linnean Period (1753-1789)<br />
Linnaeus (1753) simply listed and described the<br />
several genera known to him (a few of them neotro-<br />
pical), which he included in his Tetrandia Monogynia<br />
and Pentrandria Monogynia. In this work he de-<br />
scribed Rondeletia L., the type genus of this tribe.<br />
Adanson (1763) simply divided the <strong>Rubiaceae</strong> [as
Introduction to the <strong>Rondeletieae</strong> 5<br />
SIPANEAE<br />
CaMl.do He mr 3.i3. Selhum.. Verort Brmahmp Rebobrert Brner DdMpre<br />
130 1873 1 191 158 193 1s2 i m tbreataert<br />
RONDELTI|EA| | | | |<br />
-SIA | |<br />
?<br />
'CON.. ........ ' .............. ............... ......................................<br />
CONDAMINE_A_<br />
-o- H U H 11 U H 11 |<br />
C,HIOCOCCEA<br />
.11<br />
I<br />
I I I I<br />
FIG. 1. Diagrammatic representation of the delimitations of the <strong>Rondeletieae</strong>, Sipaneeae, Simireae, Condamineeae<br />
(and its three subtribes), Catesbaeeae, and Chiococceae according to the various authors. Vertical bars indicate tribal delimitation:<br />
<strong>Rondeletieae</strong> (black), Sipaneeae (vertical lines), Simireae (horizontal lines), Condamineeae (white), Catesbaeeae<br />
(dotted), and Chiococceae (oblique lines).<br />
"Aparines"] into two sections: section I with two ovules<br />
per ovary, and section II with more than two ovules per<br />
ovary, without giving any further subdivision.<br />
Post-Linnean Period (1789-1828)<br />
Antoine Laurent de Jussieu (1789, 1807, 1820)<br />
produced the first important <strong>Rubiaceae</strong> classification,<br />
dividing them into four arbitrary sections: Sect. I, with<br />
dicoccous fruits and uniovulate locules; Sect. II, with<br />
bilocular fruits, uniovulate locules, and capsular or<br />
baccate; Sect. III, with bilocular fruits, multiovulate<br />
locules, capsular or baccate; and Sect. IV, with multilocular<br />
fruits, uniovulate locules, baccate or<br />
drupaceous. In the second section he included<br />
Chimarrhis, due to Jacquin's (1763) erroneous observation<br />
that genus had two-seeded capsules.<br />
Rafinesque (1820) proposed a new elaboration of<br />
the Linnean classification for his "Classe Endogynie,"<br />
which he arbitrarily divided into three orders based<br />
on the number of locules and number of seeds per<br />
locule. In this system the members of the <strong>Rubiaceae</strong><br />
were represented as a mixture of previously described<br />
and Rafinesque's odd new genera, spread throughout<br />
his several ill-defined "families" ("Famille[s] Sambucidia,<br />
Dilarnia, Polarnia, Lamospermia, Aparinia,<br />
Cephalanthia, Viburidia"), which were in turn disseminated<br />
within the last two orders (Polyspia and<br />
Sphanidia) of his classification. Needless to say,<br />
Rafinesque's system was never adopted.<br />
Candollean Period (1829-1872)<br />
The two leading <strong>Rubiaceae</strong> classifications (A. Ri-<br />
chard, 1830; A. P. de Candolle, 1830) of this period<br />
were still pervaded by fixed species concepts.<br />
Achille Richard (son of Louis Claude Marie Ri-<br />
chard) presented a new <strong>Rubiaceae</strong> classification of 11<br />
tribes and 154 genera at the Royal Academy of Sci-<br />
ences in Paris on 7 July 1829. As Richard explained<br />
in the preface of this work, during that year Augustin<br />
Pyramus de Candolle was writing the treatment of the<br />
<strong>Rubiaceae</strong> for his Prodromus, and Richard was invited<br />
to submit his unpublished manuscript to be included<br />
in the Prodromus. Richard's Memoire sur lafamille<br />
Rubiacees (the first <strong>Rubiaceae</strong> monograph) went to<br />
press separately, in Paris in 1829, and was eventually<br />
published in December 1830 (and reprinted in 1834).<br />
De Candolle's Prodromus was published in Septem-<br />
ber 1830, anticipating the publication of Richard's<br />
Memoire by three months. Richard's taxa (tribes and<br />
genera) newly published in the Prodromus, were not<br />
or only inconsistently cited by de Candolle, some fol-<br />
lowed by "Rich." and others without any citation. I<br />
agree with Steam's (1957) conclusion that all of the<br />
new taxa and new combinations established by Rich-
6 Flora Neotropica<br />
ard and published in the Prodromus should be cited 2. Portlandiinae [as "Portlandieae. Calycis limbus<br />
as: "A. Richard in de Candolle." Richard (1830), in brevis v. elongatis, lobi nunquam in laminan<br />
his Memoire, founded and described the tribe foliaceam dilatati. Corollae tubus elongatus,<br />
Cinchoneae so as to include those rubiaceous genera saepissime angulatus, lobi reduplicatim valwith<br />
bilocular many-seeded capsules and winged vati"], with Portlandia, Isidorea A. Rich. in<br />
seeds, a group that included nine genera of the Ron- DC., Bikkia, and Morierina Vieill.<br />
deletieae s.l.<br />
3. Pinckneyinae [as "Pinkneyeae. Calycis lobus<br />
Augustin Pyramus de Candolle<br />
unus hic illic in laminam foliaceam<br />
(September 1830)<br />
amplam<br />
largely followed (and sometimes<br />
coloratum<br />
literally reproduced)<br />
petiolatam dilatatus"], with Pinck-<br />
Richard's classification, but with several modifica- neya [as "Pinkneya"] and Pogonopus.<br />
tions and innovations, presenting a system of 13 tribes<br />
and 223 genera. Following Richard, he maintained the In the <strong>Rondeletieae</strong>, Hooker included Warszetribe<br />
Cinchoneae (as "Cinchonaceae"). Adding to wiczia Klotzsch, Pallasia Klotzsch (= Wittmackanthus<br />
Richard's classification, de Candolle (1830) validated Kuntze), Bathysa C. Presl, Elaeagia Wedd., Chalepothe<br />
tribe Hedyotideae proposed by Chamisso and phyllum J. D. Hook., Lindenia Benth. (= Augusta<br />
Schlechtendal (1829), and established the subtribe Pohl), Gloneria Andre (= Psychotria L.), Augusta,<br />
Rondeletiinae [as "<strong>Rondeletieae</strong>"], where he placed Greenea Wight & Am. [as "Greenia"], Deppea Cham.<br />
nine genera of the <strong>Rondeletieae</strong> s.l.<br />
& Schltdl. (Hamelieae), and Carphalea Juss. [as<br />
Endlicher (1838) largely followed de Candolle's "Carphalia"].<br />
classification and divided the tribe Hedyotideae into Baillon (1880) followed both de Candolle's and<br />
the subtribes Hedyotidinae (as "Hedyoteae") and Hooker's classifications but thought that corolla aes-<br />
Rondeletiinae (as "<strong>Rondeletieae</strong>").<br />
tivation alone was insufficient to separate rubiaceous<br />
tribes. Baillon's tendency was to lump similar tribes<br />
Late 19th Century (1873-1900)<br />
and genera superficially, producing a classification<br />
that subdivided the <strong>Rubiaceae</strong> into 13 tribes and 194<br />
Joseph Dalton Hooker (1873) produced the first<br />
<strong>Rubiaceae</strong> classification that followed<br />
genera. He included the Condamineeae and the Ronevolutionary<br />
deletieae in the "Portlandia Series" or "Portlandieae"<br />
concepts, which were proposed by his friend and colleague<br />
Charles Darwin (1859). Hooker's classifica-<br />
(both of them nom. illeg. because the group contains<br />
tion was a major breakthrough in the classification of<br />
Rondeletia). Baillon went into lengthy speculative<br />
this increasingly large family, where he proposed 25<br />
explanations in which he hypothesized relationships<br />
tribes and 337 genera. Hooker was the first to intro- among the genera that he included in the same "series"<br />
or<br />
duce corolla aestivation as a<br />
tribes,<br />
cardinal tribal character<br />
increasing the understanding of this<br />
in constructing a phylogenetic classification. In this<br />
large family and supporting evolutionary theories,<br />
which were still debated<br />
work he established the tribe Condamineeae<br />
during his times.<br />
[as<br />
Herman Karsten<br />
"Condaminieae"], separating it from the <strong>Rondeletieae</strong>.<br />
(1887) commented on the newly<br />
established Condamineeae of J. D. Hooker<br />
He defined the Condamineeae as a group with valvate<br />
(1873:<br />
corollas (vs. imbricate or contorted in the Rondele-<br />
356) as follows (free translation from German):<br />
tieae), and distinguished it from the Hedyotideae in Bentham & [J. D.] Hooker, like their predeceshaving<br />
simple stipules, sometimes bifid, and with sors, believe that the Condamineeae are natuwoody<br />
habit (vs. dentate to setose stipules and herba- ral genera, when they use the deciduous or perceous<br />
habit). Hooker also stated that the Condami- sistent calyx lobes and the basifixed or<br />
neeae and the Hedyotideae were similar in having dorsifixed anthers together with the mode of<br />
capsular fruits with numerous horizontal seeds [al- dehiscence [of the anthers] as principal charthough<br />
Chimarrhis (Condamineeae sensu Hooker) has acters. The related Cascarilla [Wedd. (= Ladenvertical<br />
peltate seeds] that are non-winged to minutely bergia Klotzsch)] shows, however, that each<br />
winged. Hooker (1873) subdivided the Condamineeae character of the calyx lobes appears variable in<br />
into three subtribes:<br />
some of its species and has no generic value.<br />
Condaminea and Chimarrhis cannot be distin-<br />
1. Condamineinae [as "Eucondamineae. lobi<br />
guished from each other in the mode of openaequalis<br />
v. subequales. Corollae tubus teres v.<br />
ing of their capsules alone, therefore [the capsub-0;<br />
lobi simpliciter valvati"], with Conda- sule dehiscence] cannot be treated as an<br />
minea, Chimarrhis, and <strong>Rustia</strong> (incl. Henlea<br />
important character.<br />
and <strong>Tresanthera</strong>)
Introduction to the <strong>Rondeletieae</strong> 7<br />
Karl Schumann (1888a, 1889, 1891) presented a A few years later Bremekamp (1940) restated his<br />
classification of 21 tribes and 346 genera that influ- skepticism about the <strong>Rubiaceae</strong> classifications availenced<br />
<strong>Rubiaceae</strong> systematics throughout the 20th cen- able at that time: "All botanists acquainted with the<br />
tury. His work was influenced by all his predecessors, family <strong>Rubiaceae</strong> will agree that the present subdivimost<br />
strongly by Hooker (1873). Following Hooker's sion [cf. Schumann, 1891] is far from satisfactory and<br />
classification, Schumann (1888a, 1891) maintained that more of its tribes are either artificial or ill-defined<br />
the Condamineeae [as "Cinchonoideae-Cinchoninae- or both." In this little-known paper he predicted the<br />
Condamineeae"] separate from the <strong>Rondeletieae</strong>, but course of future <strong>Rubiaceae</strong> studies during the 20th<br />
he did not recognize the subtribal ranks proposed by century, declaring that fruit succulence should be re-<br />
Hooker. In addition, Schumann (1888a, 1891) pro- garded as being of secondary importance, and that the<br />
duced the first comprehensive keys to the 10 genera primary taxonomic characters were to be found in<br />
that he included in the Condamineeae and to the 15<br />
genera of his <strong>Rondeletieae</strong>. Of the genera here included presence or absence of hairs in the corolla<br />
in the <strong>Rondeletieae</strong>, Schumann<br />
throat; the<br />
(1888b)<br />
place where the stamens are inserted<br />
presented<br />
in the<br />
detailed discussions of Sickingia Willd. [= Simira tube; the presence or absence of floral<br />
Aubl.], Chimarrhis, Pinckneya sensu Allemao & dimorphism; the uni- or bisexuality of the flow-<br />
Saldanha<br />
ers; the insertion and<br />
(Saldanha, 1872) [= Simira], <strong>Tresanthera</strong>,<br />
shape of the placenta; the<br />
number of<br />
Henlea [= <strong>Rustia</strong>], <strong>Rustia</strong>, and Phitopis J. D. Hook.<br />
ovary cells and the presence or absence<br />
of false septs [septa]; the nature of the<br />
Modern Period<br />
stipules; the position of the inflorescences, etc.<br />
(1901-1990)<br />
These characters, however, are at present but<br />
During the first half of the 20th century, system- imperfectly known, and therefore a satisfactory<br />
atists strived to construct phylogenetic classifications subdivision can be given, the genera themselves<br />
by attempting to detect "primitive" or "derived" char- will have to be studied in more detail.<br />
acters. The concept was taken to extremes by [Bremekamp, 1940: 172]<br />
Hutchinson (1926), who divided the Angiosperms into<br />
Verdcourt<br />
two main, supposedly phylogenetic, branches: herba-<br />
(1958), strongly influenced by<br />
ceous and woody. Nevertheless, in his key to<br />
Bremekamp's early work, was the second rubiologist<br />
tropical to<br />
West African <strong>Rubiaceae</strong> (Hutchinson &<br />
classify the <strong>Rubiaceae</strong> on the basis of total evidence,<br />
Dalziel,<br />
1931) he maintained corolla aestivation, number of<br />
adding new sets of characters, such as chromosomes,<br />
ovules per locule, and fruit fleshiness as cardinal tribal<br />
pollen grains, hairs, stipules, raphides, seed albumen,<br />
and<br />
characters for this<br />
embryos, and constructing the <strong>Rubiaceae</strong> classifamily.<br />
fication<br />
In 1934 Bremekamp published a highly influenusing<br />
sets of characters. He divided the <strong>Rubiaceae</strong><br />
into three subfamilies<br />
tial, and at that time quite controversial, article. In it<br />
(Rubioideae,<br />
he criticized Baillon's and Schumann's systems for<br />
Cinchonoideae, and Guettardoideae) and 28 tribes,<br />
being constructed on groups formed by a dichotomous<br />
using presence or absence of raphides (calcium oxclassification<br />
based on single characters, and noted<br />
alate), more or less developed embryos, and presence<br />
or absence of albumen in the seeds as main characthat<br />
several of their groups were paraphyletic. He<br />
ters. He combined the Condamineeae with the Rondeclared<br />
that the wings of certain seeds were not significant<br />
taxonomic characters, contrary to Baillon's<br />
deletieae, stating:<br />
opinion, and doubted that <strong>Rubiaceae</strong> classifications The <strong>Rondeletieae</strong> are well-characterized by<br />
based on ovule number and ovules per locules were their capsular fruits containing large numbers<br />
natural. Bremekamp was also the first to group cer- of minute horizontal seeds. The testa cells are<br />
tain genera according to their pollination mechanism pitted and the aestivation not valvate. The Con-<br />
(the "ixoroid pollination syndrome"). In the same damineeae are very similar but the aestivation<br />
paper, Bremekamp placed Chimarrhis in the tribe is valvate and the seeds often winged. In both<br />
Chimarrheae, but without giving any description or the habit is mainly arborescent and the stipules<br />
delimitation of the group, so this tribal name is as a are entire or at the most bifid. I would combine<br />
nomen nudum; he apparently abandoned this tribal these two groups into one tribe and consider<br />
name because it never appeared again in his later them to be subtribes at the most [subtr.<br />
work. In addition, he separated the tribe Sipaneeae Condamineinae Verdcourt]. In the Cinchoneae<br />
from the <strong>Rondeletieae</strong>, principally because of its her- placed next to them the aestivation character<br />
baceous habit (Sipaneeae is here returned to the Ron- is considered valueless by all previous workdeletieae).<br />
ers because if used it would divide an other-
8 Flora Neotropica<br />
wise compact tribe-all three types ofaestiva- inae are worthy of recognition at tribal level." This<br />
tion occurring in it. I do not think this one char- was probably the first suggestion that the Condamiacter<br />
is sufficient to separate the <strong>Rondeletieae</strong> neeae sensu Hooker (1873) were a paraphyletic asand<br />
Condamineeae particularly as two types of semblage.<br />
aestivation occur in the <strong>Rondeletieae</strong>. Garcia Kirkbride (1982) presented a linear system<br />
[Verdcourt, 1958: 245]<br />
for the <strong>Rubiaceae</strong> to be used for floristics. In this work<br />
she constructed two hypothetical phylogenetic trees,<br />
In his last paper on <strong>Rubiaceae</strong> classification, based on the presence or absence of albumen and the<br />
Bremekamp (1966) analyzed Verdcourt's classifica- presence or absence of raphides, treating the most<br />
tion, with which he partly agreed. In contrast to his common state of these two characters as the primiprevious<br />
classification (Bremekamp, 1934), he di- tive one. After her rather speculative phylogeny, she<br />
vided the <strong>Rubiaceae</strong> into 8 subfamilies and 43 tribes. decided to adopt the same subfamilial and tribal clas-<br />
In agreement with Hooker (1873) and Schumann sification proposed by Verdcourt (1958), including the<br />
(1891), but contrary to Verdcourt (1958), he main- Condamineeae in the <strong>Rondeletieae</strong>, but she listed the<br />
tained the Condamineeae separated from the Ronde- taxa with the same linear arrangement presented by<br />
letieae, stating: "The <strong>Rondeletieae</strong> differ from the Bremekamp (1954b, 1966) "because it better illus-<br />
Condamineae [sic] in the aestivation of the corolla trates the subfamilial phylogeny of the <strong>Rubiaceae</strong>"<br />
lobes, which in this tribe is either imbricate or con- (Garcia Kirkbride, 1982).<br />
torted. However, as these two types of aestivation are, Robbrecht (1988), in his comprehensive treatment<br />
in contrast to imbricate and valvate, but rarely met Tropical Woody <strong>Rubiaceae</strong>, divided the family into 4<br />
with in nearly related genera, it is not impossible that subfamilies (Rubioideae, Antirheoideae, Ixoroideae,<br />
further study will lead to splitting up of this tribe [Ron- and Cinchonoideae) and 44 tribes (and estimated 637<br />
deletieae]." In his general discussion Bremekamp genera and 10,700 species). In this classification he<br />
(1966) also added: "With regard to his [Verdcourt's maintained the views of Hooker (1873) and<br />
(1958)] reduction of the Condamineeae to the Ron- Bremekamp (1966), and maintained the Condamineeae<br />
deletieae I feel rather skeptical, but I admit that both separated from the <strong>Rondeletieae</strong>, without recognizing<br />
tribes are very imperfectly known and deserve a any subtribal divisions. He described the Condamithrough<br />
revision."<br />
neeae as follows: "Trees or shrubs. Stipules mostly<br />
Aiello (1979) followed Bremekamp's (1966) clas- interpetiolar, entire, rarely bifid. Raphides absent.<br />
sification, and treated seed insertion and position as Inflorescences mostly terminal and many-flowered.<br />
key tribal and generic characters. She placed Corolla aestivation valvate, rarely imbricate. Ovary<br />
Portlandia and some of the genera that she separated 2-locular, with many horizontal ovules in each locfrom<br />
Portlandia in the Condamineeae, because of ule. Capsules with many horizontal seeds. Exotestal<br />
their horizontal seeds; transferred Siemensia from the cells with pitted thickening along the inner tangential<br />
Condamineeae to the Hedyotideae (subfam. Rubioideae) walls. Pollen mostly 3-colporate, rarely porate<br />
because of their raphides, multicellular hairs, and (Bikkia)." He listed the following genera under the<br />
numerous tiny seeds; and transferred Coutarea Aubl. tribe Condamineeae: Badusa A. Gray, Bikkia,<br />
to the Cinchoneae because of its winged vertical seeds. Ceuthocarpus, Chimarrhis, Condaminea, Cubanola<br />
Finally, she treated Hintonia Bullock, Ceuthocarpus, Aiello, Dioicodendron, Flexanthera [= Simira],<br />
Coutaportla, Cigarilla Aiello [= Nernstia Urb.], and Isidorea, Kerianthera J. H. Kirkbr. [Isertieae],<br />
Schmidtottia as genera of uncertain position, and de- Morierina, Nernstia, Osa, Parachimarrhis, Picarclared<br />
that "none of the above placements can be made daea, Pinckneya, Pogonopus, Portlandia, <strong>Rustia</strong>,<br />
with great certainty since a large amount of work re- Schmidtottia, Siemensia, Stomandra [= <strong>Rustia</strong>],<br />
mains to be accomplished with regard to tribal rela- Thogsennia, and <strong>Tresanthera</strong>, and tentatively included<br />
tionship within the <strong>Rubiaceae</strong>."<br />
Kajewskiella Merr. & Perry and Pseudomussaenda<br />
Ridsdale (1982) reported: "In aestivation the sub- Wernham.<br />
tribe Portlandiinae differs from the remainder of the The <strong>Rondeletieae</strong> were described by Robbrecht<br />
Condamine[e]ae; Coutarea, Exostema (Pers.) Humb. (1988) as follows: "Trees or shrubs, rarely (Sipane-<br />
& Bonpl., Hintonia, and Schmidtottia differ from this opsis Steyerm.) herbaceous. Stipules entire, rarely<br />
subtribe only in the orientation of the seeds and their bifid. Raphides absent. Further little differing from the<br />
mode of attachment. However, the remainder of the Condamineeae: corolla lobes imbricate or more rarely<br />
Cinchoneae is also heterogeneous in this respect contorted. Flowers frequently heterostylous. Pollen<br />
(Ridsdale & Friskus, unpublished observations). Fur- mostly 3-colporate." In this tribe he included<br />
ther study might as well indicate that the Portlandi- Acrobotrys K. Schum & K. Krause, Acunaeanthus
Introduction to the <strong>Rondeletieae</strong> 9<br />
Borhidi, Koml. & M. Moncada, Aleisanthia, Arach-<br />
nothryx Planch. [= Rondeletia s.l.], Augusta, Bathysa,<br />
Blandibractea Werham [= Simira], Chalepophyllum,<br />
Cuatrecasasiodendron Standl. & Steyerm., Dendro-<br />
sipanea Ducke, Eizia Standl., Elaeagia, Glionnetia,<br />
Gloneria Andre [= Psychotria L.], Greenea, Holsti-<br />
anthus Steyerm., Javorkaea Borhidi & Koml.,<br />
Lindenia [=Augusta], Macbrideina Standl., Maguire-<br />
othamnus Steyerm., Neblinathamnus Steyerm., Pteri-<br />
docalyx Wernham, Rogiera Planch. [= Rondeletia<br />
s.l.], Roigella [= Rondeletia s.l.], Rondeletia, Simira,<br />
Sipaneopsis, Spathichlamys, Steenisia Bakh.f.,<br />
Stevensia Poit., Warszewiczia, and Wendlandia.<br />
Following Bremekamp (1966), Robbrecht provi-<br />
sionally maintained the Sipaneeae (with Limno-<br />
sipanea, Sipanea, and Steyermarkia, and treating<br />
Neobertiera as a dubious genus) separate from the<br />
<strong>Rondeletieae</strong> because of their herbaceous to<br />
subfrutescent habit, but noted that "a revision of the<br />
<strong>Rondeletieae</strong>/Condamineeae<br />
complex may well show<br />
that this is not justified."<br />
The Last Decade of the 20th Century<br />
The last decade of the 20th century has seen the<br />
advent, and the first applications in the <strong>Rubiaceae</strong>, of<br />
a new methodology and a new branch of science that<br />
revolutionized the apparently stable system proposed<br />
by Robbrecht (1988): cladistic analyses and molecular<br />
biology. In addition, during this decade the <strong>Rubiaceae</strong><br />
has finally received the increased attention of<br />
the botanical community, as shown by the organization<br />
of two international <strong>Rubiaceae</strong> congresses and by<br />
the considerable proliferation of botanists dedicated<br />
to <strong>Rubiaceae</strong> systematics.<br />
Andersson and Persson (1991) published the first<br />
<strong>Rubiaceae</strong> cladistic analysis in which, using morphological<br />
characters, they attempted to circumscribe the<br />
tribe Cinchoneae. <strong>Part</strong>ly as a result of this analysis,<br />
the genera Cigarilla (= Nernstia), Coutarea, Exostema,<br />
Molopanthera, Syringantha Standl., and Wernhamia<br />
S. Moore (= Simira) were transferred from the<br />
Cinchoneae to the Condamineeae. Andersson and<br />
Persson (1991) commented on these transfers as follows:<br />
"Coutarea and Exostema seem to have their true<br />
relationship with Portlandia and related genera.<br />
Portlandia is presently referred to the tribe Condamineeae<br />
[sensu Robbrecht, 1988], to which Coutarea<br />
and Exostema may thus be referred. Unfortunately, it<br />
seems that this might only create new problems, since<br />
a relationship between Condaminea and the Portlandia<br />
group is not supported by our analysis." Their<br />
comments support the inclusion of the "Portlandiagroup"<br />
in the Catesbaeeae sensu Delprete (1996d).<br />
Bremer and Jansen (1991) presented the first cladistic<br />
analysis of the <strong>Rubiaceae</strong> using molecular data<br />
(chloroplast DNA restriction site mutations) obtained<br />
from 33 representative genera from 18 tribes (sensu<br />
Robbrecht, 1988). In their cladograms, Pogonopus<br />
and Pinckneya shared the same clade with Calycophyllum<br />
DC. (Calycophylleae; cf. Andersson &<br />
Persson, 1991) and Mussaenda L. (Isertieae sensu<br />
Robbrecht, 1988), and the same basal clade with<br />
Erithalis P. Browne and Chiococca P. Browne, Exostema,<br />
Coutarea, and Catesbaea. The results of their<br />
study suggested for the first time a relationship between<br />
the subtribe Portlandiinae (Condamineeae) with<br />
the tribe Chiococceae. This is also supported by several<br />
morphological similarities (i.e., filaments attached<br />
at the base of corolla tube, anthers basifixed,<br />
stigmatic area consisting of two narrow lines, etc.).<br />
Bremer (1992) presented a cladistic analysis using<br />
20 morphological characters of genera in subtribe<br />
Portlandiinae (Condamineeae) and tribe Chiococceae.<br />
She concluded that the former group should be included<br />
in the latter. Many morphological characters<br />
were not considered in her analysis: fruit morphology,<br />
number and position of ovules (many horizontal or<br />
imbricate in Portlandiinae vs. two pendulous in<br />
Chiococceae s.str.), placentation (central in Portlandiinae<br />
vs. apical in Chiococceae s.str.), anther shape and<br />
attachment (linear and basifixed in Portlandiinae vs.<br />
elongate and dorsifixed in Chiococceae).<br />
At the First International Conference on the Systematics<br />
of the <strong>Rubiaceae</strong> (4-6 October 1993, St.<br />
Louis, Missouri; cf. Taylor, 1995), Bremer (1996)<br />
presented a phylogenetic analysis using rbcL molecular<br />
sequences that indicated that, in contrast to<br />
Robbrecht's (1988, 1993) classifications, the <strong>Rubiaceae</strong><br />
are better divided into three major groups,<br />
namely, the Cinchonoideae s.str., Ixoroideae s.l., and<br />
Rubioideae. During this conference, Delprete (1993)<br />
presented the results of a preliminary phylogenetic<br />
analysis of 43 genera using 44 morphological characters<br />
of all Condamineeae (sensu Robbrecht, 1988),<br />
all Chiococceae (sensu Robbrecht, 1988), several<br />
<strong>Rondeletieae</strong>, two Cinchoneae (Cinchona L. and<br />
Joosia H. Karst.), and Coffea L. (Coffeae) as<br />
outgroups. He concluded that at the tribal level the<br />
Portlandiinae (Hooker, 1873) should be separated<br />
from both the Condamineeae s.str. and the Chiococceae<br />
s.str., and instead be included in the Catesbaeeae<br />
(a tribe treated as incertae sedis by Robbrecht,<br />
1988). For further information about this analysis see<br />
Phylogenetic Relationships, below.<br />
Robbrecht (1993), following the conclusions presented<br />
by Delprete (1993), treated the Portlandiinae<br />
as an informal "Portlandia-group" within the Conda-
10 Flora Neotropica<br />
mineeae, but had reservations about including the PHYLOGENETIC RELATIONSHIPS<br />
group in the Catesbaeeae (sensu Hooker). The genera<br />
listed under the Condamineeae by Robbrecht<br />
(1993) were Chimarrhis, Condaminea, Dioicodendron,<br />
Flexanthera [= Simira], Kerianthera [Isertieae],<br />
Parachimarrhis, Picardaea, Pinckneya, Pogonopus,<br />
<strong>Rustia</strong> (incl. Stomandra), <strong>Tresanthera</strong>, and tentatively<br />
Kajewskiella Merr. & Perry, with Pseudomussaenda<br />
Wernham transferred to the Isertieae. Robbrecht<br />
(1993) made several changes in the genera that he<br />
previously included in the <strong>Rondeletieae</strong> (Robbrecht,<br />
1988). Following Lorence (1991), Robbrecht (1993)<br />
treated Arachnothryx, Rogiera, and Javorkaea as synonyms<br />
of Rondeletia, and also added Mazaea Krug<br />
& Urb. He maintained the Sipaneeae (to which he<br />
added Neobertiera Wernham) and separated the<br />
Simireae (here included, and proposed to be closely<br />
related to Parachimarrhis) from the <strong>Rondeletieae</strong>.<br />
At the Second International <strong>Rubiaceae</strong> Conference<br />
(13-15 September 1995, Meise, Belgium), Bremer<br />
(1996) presented a second phylogenetic analysis using<br />
rbcL molecular sequences that supported her previous<br />
results (Bremer, 1995) concerning the monophyletic<br />
origin of the <strong>Rubiaceae</strong> and their subdivision<br />
into three main lineages. Delprete (1995a, 1996d)<br />
presented a second set of tribal-level phylogenetic<br />
analyses using morphological data, from 170 species<br />
of 44 genera, including all Condamineeae and Catesbaeeae<br />
sensu Hooker and selected Chiococceae, <strong>Rondeletieae</strong>,<br />
Cinchoneae, and Coffeae. According to the<br />
results of these analyses (Delprete, 1995a, 1996d), the<br />
two subtribes Condamineinae and Pinckneyinae were<br />
transferred into a broader, tentative delimitation of the<br />
<strong>Rondeletieae</strong>, which is adopted, with few modifications,<br />
in the present treatment. He also included the<br />
subtribe Portlandiinae (Condamineeae sensu Hooker,<br />
1873) in the Catesbaeeae. The tribe Catesbaeeae<br />
(sensu Hooker, 1873) has nomenclatural priority over<br />
the subtribe Portlandiinae (also, Portlandieae Baillon<br />
is a nomen illegitimum because it includes Rondeletia);<br />
therefore, the newly emended tribe should be called<br />
"Catesbaeeae J. D. Hooker emend. Delprete" (Delprete,<br />
1996d). An abbreviated version of these analyses is<br />
presented in Phylogenetic Relationships, below.<br />
Classification at subfamilial and tribal levels in the<br />
<strong>Rubiaceae</strong> has been debated since its establishment,<br />
and up to the present day no overall consensus among<br />
workers has been reached. As certain taxonomic characters<br />
are carefully studied, or new ones are added to<br />
phylogenetic analyses, the macrosystematics of the<br />
<strong>Rubiaceae</strong> have been reexamined and modified accordingly.<br />
The number of recognized subfamilies within the<br />
<strong>Rubiaceae</strong> varied according to systematists' interpretation<br />
of the usefulness and taxonomic importance of<br />
certain characters. Using similar data sets, Verdcourt<br />
(1958) proposed three subfamilies (Rubioideae,<br />
Cinchonoideae, and Guettardoideae) whereas<br />
Bremekamp (1966) divided the <strong>Rubiaceae</strong> into the<br />
largest number of subfamilies ever proposed, eight:<br />
Cinchonoideae, Urophylloideae, Pomazotoideae,<br />
Gleasonioideae, Guettardoideae, Ixoroideae, Rubioideae,<br />
and Hillioideae. The most recent comprehensive<br />
classification using morphological, anatomical,<br />
and cytological data was proposed by Robbrecht<br />
(1988, 1993), who recognized four subfamilies<br />
(Cinchonoideae, Rubioideae, Antirheoideae, and<br />
Ixoroideae) and 44 tribes (a compromise between<br />
Verdcourt's and Bremekamp's classifications).<br />
Bremer (Bremer et al., 1995; Bremer, 1996) presented<br />
phylogenetic analyses using rbcL molecular sequences<br />
showing evidence for recognition of only<br />
three subfamilies (merging the Antirheoideae within<br />
the Ixoroideae s.l.), which was later supported by<br />
trnL-F (Rova et al., 1997) and rps 16 (Andersson,<br />
unpubl. data) molecular sequences.<br />
The phylogenetic analyses using molecular and<br />
morphological data presented by Bremer (1992),<br />
Bremer and Eriksson (1992), and Bremer and Struwe<br />
(1992) supported Bremekamp's and Verdcourt's opinions<br />
about the ambiguous usefulness of certain flower<br />
and fruit characters. In my opinion, fruit and flower<br />
characters have often been superficially or erroneously<br />
interpreted in trying to detect phylogenetic relationships.<br />
In order to avoid the repetition of erroneous<br />
observations, morphological characters and characters<br />
states should be coded after direct observation of fresh<br />
Rova et al. (1997) presented a phylogenetic analysis<br />
using trnL molecular sequences of more than 100<br />
species representative of the most tribes in<br />
Robbrecht's (1988, 1993) classification. These analyses<br />
supported the rbcL phylogenies of Bremer et al.<br />
(1995) and Bremer (1996), and rpsl6 phylogenies of<br />
Andersson (in progress, unpubl. data) in dividing the<br />
<strong>Rubiaceae</strong> in three main groups: Cinchonoideae s.str.,<br />
Ixoroideae s.l., and Rubioideae.<br />
or preserved material, instead of being reported from<br />
previous literature. The taxonomic significance of<br />
characters such as number of ovules per locule, placentation,<br />
mesocarp fleshiness, seed insertion and<br />
position within the fruit, dioecy, heterostyly, and corolla<br />
aestivation should be evaluated with accurate<br />
anatomical and morphological studies, and compared<br />
with phylogenies obtained with alternative sets of data<br />
(i.e., molecular, biochemical, cytological data).
Introduction to the <strong>Rondeletieae</strong> 11<br />
Recently, tribal rearrangements have occurred ses. In all analyses <strong>Rustia</strong> and <strong>Tresanthera</strong> were<br />
within subfam. Cinchonoideae, and mostly among the placed as sister genera on one clad,, supported by<br />
Cinchoneae, <strong>Rondeletieae</strong>, Condamineeae, Cates- three synapomorphies (8:1 = leaves with pellucid<br />
baeeae, and Chiococceae. The Condamineeae and the glands; 11:2 = presence of secundiflorous panicles;<br />
Sipaneeae have been historically treated either as sis- 42:1 = testa with ridges and small pits). Augusta and<br />
ter tribes (Hooker, 1873; Schumann, 1891; Lindenia were consistently placed as sister taxa on one<br />
Bremekamp, 1966) or as included (Baillon, 1880; clade, supported by testa sculpturing with small pits<br />
Verdcourt, 1958) in the <strong>Rondeletieae</strong>. The Chiococ- to ? smooth (42:0), which justifies Kirkbride's<br />
ceae were treated as a tribe (Hooker, 1873; Schumann, (1997b) recent inclusion of Lindenia in Augusta.<br />
1891; Bremekamp, 1934, 1966; Verdcourt, 1958; These analyses demonstrated that the Condami-<br />
Robbrecht, 1988) defined by two-seeded fleshy fruits neeae (sensu Hooker) were paraphyletic, and their<br />
until Bremer (1992) included the Portlandiinae (with three subtribes were rearranged and delimited accordmany-seeded<br />
capsules). Catesbaea and Phyllacanthus ingly. The Condamineinae and the Pinckneyinae,<br />
were separated from the Gardenieae by Hooker (1873) which included the genera treated in the present monoto<br />
establish the Catesbaeeae; these two genera were graph (<strong>Rustia</strong>, <strong>Tresanthera</strong>, Condaminea, Picardaea,<br />
referred to the Gardenieae (Ixoroideae) (Grisebach, Pogonopus, Pinckneya, Chimarrhis, Dioicodendron,<br />
1861; Baillon, 1880; Schumann, 1891), treated as a Molopanthera, and Dolichodelphys), were merged in<br />
separate tribe near the Gardenieae (Hooker, 1873; a broad delimitation of the tribe <strong>Rondeletieae</strong>, result-<br />
Verdcourt, 1958), or with uncertain relationship ing in the complete dismantling of tribe Condami-<br />
(Robbrecht, 1988).<br />
neeae.<br />
Monophyly and phylogenetic relationships of the According to these analyses, the subtribe Port-<br />
Condamineeae, <strong>Rondeletieae</strong>, Chiococceae, and landiinae (of the former Condamineeae sensu Hooker)<br />
Catesbaeeae (sensu Robbrecht, 1988) were tested in was included within the newly emended Catesbaeeae<br />
various cladistic analyses by Delprete (1993, 1995a, (treated by Robbrecht [1988] as incertae sedis), for<br />
1996d) using morphological data. The final analyses which a revised tribal delimitation was presented<br />
involved 170 species of 44 genera of all Condami- (Delprete, 1996d). The inclusion of the Portlandiinae<br />
neeae, all Catesbaeeae s.str. and selected Chiococceae, into the emended Catesbaeeae is supported by the<br />
<strong>Rondeletieae</strong>, Cinchoneae, and Coffea (Coffeae). karyological studies of Kiehn (1995, and pers.<br />
Forty-four morphological characters were scored from comm.), in which they are reported to be x = 12 and<br />
direct observation of herbarium specimens and/or liv- very large.<br />
ing material, and entered into a data matrix. Three In these analyses the Chiococceae and the Catesphylogenetic<br />
analyses were performed (using PA UP baeeae (both sensu Delprete, 1996d) appeared to be<br />
3.1 [Swofford, 1993]), each with a different outgroup: closely related, and in one analysis they were mapped<br />
the first using Cinchona and Joosia, the second using as sister groups. Nevertheless, recent studies using<br />
Coffea, and the third using Cinchona, Joosia, and trnL-F sequence data (Rova, Andersson, Delprete &<br />
Coffea simultaneously. A strict consensus tree and a Albert, in progress) indicated that Catesbaeeae and<br />
majority rule tree were produced for each analysis. Chiococceae (both sensu Delprete, 1996d) form a<br />
The consensus trees were produced from PA UP and strongly supported monophyletic group.<br />
were then transferred to MacClade (Maddison &<br />
Maddison, 1992) for character mapping and interpre- CURRENT CIRCUMSCRIPTION OF THE<br />
tation of state changes on the cladogram. Finally, RONDELETIEAE, INCLUDING NOTES ON GENERA<br />
synapomorphies and parallel homoplasies ("parallel- NEWLY INCLUDED, EXCLUDED, OR SYNONYMIZED<br />
isms") were mapped on the majority rule consensus<br />
trees obtained in each analysis. The majority rule con- The current definition and delimitation of the Ronsensus<br />
tree using Cinchona, Joosia, and Coffea as deletieae are tentative, and its monophyly has yet to<br />
outgroup is shown in Figure 2 (for detailed informa- be tested. The description of this group is intended for<br />
tion see Delprete, 1996d). In this analysis, the this floristic treatment and is not meant to be strictly<br />
Chiococceae, Portlandiinae, and the Catesbaeeae phylogenetic. Table I summarizes the neotropical<br />
formed a monophyletic group supported by one genera of <strong>Rondeletieae</strong>, and gives for each the apsynapomorphy<br />
(44:0 = pollen exine echinate), with proximate number of species, occurrence in neotrothe<br />
Catesbaeeae nested within the Portlandiinae. The pical countries, habitat, and elevation ranges.<br />
rest of the Condamineeae (Condamineinae and The following notes are necessary for clarification<br />
Pinckneyinae sensu Hooker) were interspersed with of the taxonomic position, delimitation, and recent<br />
the selected members of the <strong>Rondeletieae</strong> in all analy- synonymy of certain taxa that have at some point been
12 Flora Neotropica<br />
-'------ I'--------ID<br />
213:01 M:0<br />
8:1 1: 2 42:1<br />
,________<br />
:<br />
|,<br />
2o:2<br />
3:2 i<br />
|<br />
CUB<br />
DAD<br />
CAT |<br />
It---------r<br />
"- 26:3 3 :1 - - -. ..T<br />
.PAR<br />
lobes absent; 35:3 =stigmatic surface linear I2:i3 along ,,.2 11 style; II 36:0 * =ovules 1 * 4:3 * pendulous; 37:1 =ovules -<br />
I= one =fruitM per locule; 38:1<br />
ss<br />
FOG<br />
CO .<br />
CHI<br />
BAT<br />
rr ~ ~ * BAUG<br />
JOO<br />
BAT<br />
COT =Coutaportla, NER=Nernstia, CUB Cubanola, THO =Thogsennia, CATC=Catesbaea, PHY Phyllacanthus, CEU<br />
FIG. 2. Majority rule consensus tree from the phylogenetic analysis of all genera of'the Condamineeae and Catesbaeeae<br />
(sensu Delprete): Catesbaeeae (black), Chiococceae (white) and <strong>Rondeletieae</strong> s.l. (oblique lines). Key to generic acronyms:<br />
RUS=<strong>Rustia</strong>, TRE =<strong>Tresanthera</strong>, CON =Condaminea, P00 =Pogonopus, PIN =Pinckneya, DOL=Dolichodelphys, DIO<br />
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<br />
=Ceuthocarpus, SCH =Schmidtottia, BIK--Bikkia, MOR-Morierina, EXO =Exostema, BAD =Badusa, CHI Chiococca,<br />
Joo41:2 =seed laterally flattened; 42:1 =testa ridged and withng strmall pits; 43:1 =testalr a indicated with dashed lines. Char-<br />
7:177,ERI Erithalis, fi Au-4. 1996).<br />
SCO Scolosanthus, PLA Placocarpa, CHO Chione, ALL Allenanthus, indenia, AUG<br />
entire marge indicated by double bars. Numbers indicate character and character states. Vertical bars indicate tribal delimitationg.
Introduction to the <strong>Rondeletieae</strong> 13<br />
referred to the Sipaneeae, Condamineeae, and Ron- 1991), to which its only species was transferred as R.<br />
deletieae sensu Robbrecht (1988, 1993), but are now brandegeeana Lorence.<br />
excluded or not treated in the first part of the Ronde- Acunaeanthus Borhidi, Koml. & Moncada was<br />
letieae. The following discussion is organized into treated as distinct from Rondeletia by Delprete<br />
informal groups according to observations stemming (1996d). Acunaeanthus is similar to Mazaea in havfrom<br />
this research. These informal groups do not im- ing septicidal, oblong capsules, 4-merous flowers,<br />
ply any taxonomical status.<br />
tubular corollas, and imbricate corolla lobes; these<br />
characters are also shared with Rondeletia. Acunae-<br />
The Bathysa Group<br />
anthus differs from Mazaea in having terminal inflo-<br />
The two species of rescences<br />
Schizocalyx (Weddell, 1854;<br />
(vs. axillary), corollas pinkish white (vs.<br />
Standley, 1929) were recently shown to be conspe- orange), ovary with 15-30 ovules per locule (vs. 2-6<br />
cific and were transferred to ovules<br />
Bathysa by Delprete per locule), and winged seeds (vs. unwinged<br />
(1997b), as B. bracteosa (Wedd.) Delprete. The seeds). The two taxa are closely related and probably<br />
calycophylls of Schizocalyx have been proven to be congeneric, but further studies are necessary to ascerof<br />
little taxonomic significance, and this genus is tain their<br />
syn- relationships with Rondeletia.<br />
onymous with Bathysa.<br />
Phitopis was described by Hooker (187 la), who<br />
The Mazaea Group<br />
stated that it had the same habit as Hippotis Ruiz & Mazaea was published by Krug and Urban<br />
Pav. (tribe Hippotieae; see Garcia Kirkbride, 1981), (1897a). As explained by Robbrecht and Bridson<br />
the first name being an anagram of the other.<br />
(1993), Neomazaea Krug & Urb. (1897b) is a nomen<br />
Schumann (1888b) analyzed the position of Phitopis<br />
superfluum, and is treated as a synonym of Mazaea.<br />
and tentatively included it in the <strong>Rondeletieae</strong>. The type species of Ariadne Urb. was originally de-<br />
Standley (1931 d) added one species to this genus and scribed by Standley (1918) as Neomazaea shaferi<br />
maintained it in the <strong>Rondeletieae</strong> (Standley, 1936). Standl., and was later transferred by Urban (1922,<br />
Phitopis is here kept in the <strong>Rondeletieae</strong> as closely 1923) to his newly established Ariadne. Delprete<br />
related to Bathysa, with which it might be congeneric. (1999a) recently compared the monotypic Ariadne<br />
Bathysa was described by Presl (1845), based on with Mazaea and treated them as congeneric, and cre-<br />
Coffea stipulata Vell. With recent transfers (Delprete, ated the new combination Mazaea shaferi (Standl.)<br />
1996b, 1997b) and the discovery of new taxa, this<br />
Delprete. Ariadne is treated as a synonym of Mazaea<br />
genus now comprises approximately 16 species dis-<br />
(incl. Neomazaea Krug & Urb.; see Robbrecht &<br />
tributed in Central and South America. An account<br />
Bridson, 1993) because of its axillary inflorescences,<br />
of the Brazilian species in this genus was recently 4-merous flowers, imbricate corolla lobes, 1-3 ovules<br />
completed by Germano Filho (1993).<br />
per locule, and narrowly oblong capsules.<br />
The monotypic Phyllomelia was founded by<br />
The Rondeletia Complex<br />
Grisebach (1866), who placed it between Machaonia<br />
The generic limits of Rondeletia and its closely and Phialanthus (Chiococceae), probably because of<br />
related taxa are in a state of flux. In the present treat- the two-seeded indehiscent fruits. Phyllomelia was<br />
ment, a Rondeletia-complex is adopted which includes placed in the Ixoreae by Standley (1934) and treated<br />
Arachnothryx (Planchon, 1849), Javorkaea (Borhidi as genus incertae sedis by Robbrecht (1988, 1993).<br />
& Jarai-Komlodi, 1983), Rogiera (Planchon, 1849), Because of the axillary, few-flowered, cymose inflo-<br />
Roigella (Borhidi & Fernmndez, 1981a) and Suber- rescences with capilliform rachis, 4-5(-6)-merous<br />
anthus. The morphological characters used to segre- corollas, filaments inserted in the middle of the cogate<br />
these genera from Rondeletia s.str. (Planchon, rolla tube, anthers narrowly oblong, and absence of<br />
1849; Steyermark, 1967,1974; Borhidi & M. Fernndez, raphides in the blades, I concluded that Phyllomelia<br />
1981a, 1981b; Borhidi et al., 1980) have been shown is closely related to Mazaea (Delprete, 1999a). Both<br />
to be variable among species of Rondeletia s.str. genera are endemic to the serpentine soils of Sierra<br />
(Lorence, 1991). The inclusion of the above listed taxa del Rosario and Sierra de los Organos, Prov. Pinar del<br />
in the Rondeletia-complex does not necessarily as- Rio, Cuba. Phyllomelia is exceptional to the Rondesume<br />
their reduction to synonymy under Rondeletia. letieae s.l. by having indehiscent 2-seeded fruits with<br />
The Rondeletia-complex is here adopted for the prac- persistent rotate corolla, which are released as distical<br />
reasons of the presentation in this treatment. persal units, called "pseudosamaras" by Spjut (1994).<br />
The monotypic Otocalyx Brandegee has been For additional information about Phyllomelia, see<br />
shown to be congeneric with Rondeletia (Lorence, Delprete (1999a).
Table I<br />
Summary of the neotropical <strong>Rondeletieae</strong> (35 genera, ca. 340 species). Genera in boldface are<br />
No. of<br />
Genera species Countries" Habitat<br />
Acrobotrys I Co Cloud forests<br />
Acrosynanthus 6 Cu, J Pine forests, serpentine barrens<br />
Acunaeanthus 1 Cu Serpentine formations<br />
Augusta 4 M, G, Be, Ho, ES, N, CR, Pa, Among rocks in turbulent creeks<br />
Br (and South Pacific)<br />
Bathysa 16 CR, Pa, Co, V, Gu, E, Pe, Br, Lowlands of Central America, cloud forests, Amazoni<br />
Bo rain forests, Brazilian Atlantic forest<br />
Chalepophyllum 3 V, Gu Wet savannas and plateaux of the Guayana shield<br />
Chimarrhis 14 Cu, J, Ha, LA, Co, V, Gu, E, Caribbean seasonal forests, Andean slopes and lowlan<br />
Pe, Br, Bo Amazonian rain forests<br />
Condaminea 1(2?) CR, Pa, Co, V, E, Pe, Br, Bo Deciduous lowland forests, cloud forests, and rain<br />
forests<br />
Cuatrecasasiodendron 2 Co Colombian cordilleras<br />
Dendrosipanea 3 V, Br Flooded and non-flooded savannas<br />
Dioicodendron 1 Co, V, E, Pe Andean seasonal forests, and cloud forests<br />
Dolichodelphys 1 Co, V, E, Pe Andeail cloud forests, and rain forests<br />
Elaeagia 15 M, G, Be, Ho, Cu, CR, Pa, Co, Dry and wet seasonal forests, Cuban Sierra Maestra<br />
V, Gu, E, Pe, Br, Bo thickets, South American cloud forests<br />
Holstianthus 1 V Venezuelan mesetas<br />
Limnosipanea 5 Pa, Co, V, Br Llanos and wet savannas, pond margins and standing<br />
water<br />
Macbrideina 1 Co, E, Pe Amazonian slopes of the Andes and lowland Amazoni<br />
rain forest<br />
Maguireothamnus 4 V, Gu, Br Tepuis of the Guayana shield<br />
Mazaea 2 Cu, Ha Pine woodlands and serpentine barrens
Molopanthera 1 Br Seasonal and Atlantic forests<br />
Neblinathamnus 4 V, Br Cerro de la Neblina<br />
Neobertiera 1 Gu Lowland rain forest<br />
Parachimarrhis 1 Co, E, Br Amazonian rain forest<br />
Phitopis 2 Co, (E?), Pe Amazonian slopes of the Andes<br />
Phyllomelia 1 Cu Serpentine barrens<br />
Picardaea 1 Cu, Ha, DR Seasonal forests<br />
Pogonopus 3 M, G, ES, CR, Pa, Co, V, Pe, Seasonal and evergreen deciduous forests, and Andean<br />
Br, Bo, Ar cloud forests<br />
Pteridocalyx 2 Gu Lowland rain forests<br />
Rondeletia complex ca. 150 US, M, G, Be, Ho, ES, N, CR, Deciduous and evergreen seasonal forests; mostly<br />
Pa, Cu, J, Ha, DR, PR, VI, Caribbean,few species in Andean cloud forests (rarely<br />
LA, Co, V, Gu, E, Pe, Bo lowland Amazonian forests)<br />
<strong>Rustia</strong> 14 G, N, CR, Pa, Co, V, E, Pe, Br Andean cloud forests, seasonal forests, Amazonian<br />
slopes of the Andes, swampy coastal areas of Pacific<br />
coast, Brazilian Atlantic forests<br />
Simira 37 M, G, Be, Ho, ES, N, CR, Pa, Seasonal lowland forests, Andean cloud forests and<br />
Co, V, Gu, E, Pe, Br, Bo Amazonian lowland rain forests Brazilian Atlantic for<br />
Sipanea 18 CR, Pa, Co, V, TT, Gu, Br, Bo Guayana shield, and Caribbean coast of Central and<br />
South America, in poorly drained moist soil<br />
Sipaneopsis 8 Co, V, Br South American savannas<br />
Stevensia 9 Ha, DR Forested thickets on lateritic or serpentine soils<br />
Steyermarkia 1 M, G Coastal Caribbean lowlands<br />
<strong>Tresanthera</strong> 1 V, TT Rain forest of coastal cordillera<br />
Warszewiczia 7 N, CR, Pa, Co, V, Gu, E, Pe, Lowland rain forests, cloud forests, and Amazonian ra<br />
Br, Bo forests<br />
'Ar, Argentina; Ba, Bahama Archipelago; Be, Belize; Bo, Bolivia; Br, Brazil; Ca, Cayman Islands; Co, Colombia; CR, Costa R<br />
ES, El Salvador; G, Guatemala; Gu, The Guianas; Ha, Haiti; Ho, Honduras; J, Jamaica; LA, Lesser Antilles; M, Mexico; N, N<br />
United States; TT, Trinidad and Tobago; V, Venezuela; VI, Virgin Islands.
16 Flora Neotropica<br />
Stevensia Poit. was described by Pierre Antoine is exceptional in having 2-seeded pseudosamaras.<br />
Poiteau (1804). Since then, several species have been Future flowering collections will shed light on the<br />
added and this genus is currently represented by nine identity and systematic relationships of this taxon.<br />
species endemic to Haiti and the Dominican Republic.<br />
Liogier (1995) recently published an updated de-<br />
The Simira Group<br />
scription of Stevensia and a key to all the species.<br />
Acrosynanthus Urb. was described by Urban<br />
Simira was established by Aublet (1775), and<br />
(1913) as a monotypic genus with 6-merous flowers. Sickingia by Willdenow (1801), with Sickingia<br />
Additional species described by other authors and erythroxylon Willd. and Sickingia longifolia Willd.<br />
himself (Standley, 1918; Urban, 1923; Howard &<br />
Baillon (1879) wrongly reduced Sickingia to a sub-<br />
Proctor, 1958) proved that this taxon has 4-5(-6)genus<br />
of Chimarrhis (see Excluded Species in the<br />
merous flowers.<br />
treatment of Chimarrhis, below). Bremekamp (1954a)<br />
convincingly demonstrated that Sickingia and Simira<br />
are congeneric, and transferred the two Willdenowian<br />
The Augusta Group<br />
species to Simira. Steyermark (1972), following<br />
Lindenia was described by Bentham (1841), and Bremekamp's conclusion, transferred all the South<br />
a detailed account of the taxonomic history and mono- American species of Sickingia to Simira. Bremekamp<br />
graphic treatment of this taxon was published by Dar- (1966) also separated Simira from the <strong>Rondeletieae</strong><br />
win (1976). Augusta was established by Pohl (1828- because of the morphology and insertion of its seeds,<br />
1829), under which he described five species endemic establishing the tribe Simireae. Robbrecht (1988) into<br />
Brazil. The complex taxonomic history of the Bra- cluded the tribe Simireae in the <strong>Rondeletieae</strong> but a few<br />
zilian taxa of Augusta has been summarized by years later (Robbrecht, 1993) reestablished the tribe.<br />
Delprete (1997a), who treated Pohl's five species as In the current treatment the Simireae are included in<br />
synonyms ofAugusta longifolia (Spreng.) Rehder. On the <strong>Rondeletieae</strong> according to a cladistic analysis usthe<br />
basis of its rheophytic habit, fruit dehiscence, and ing morphological characters (Delprete, 1996d). Reinflorescence<br />
position and architecture, Kirkbride cent accounts of Simira from selected Brazilian states<br />
(1997b) included this taxon in Augusta, as subgen. were published by Peixoto (1981, 1982) and Barbosa<br />
Lindenia. Kirkbride attributed the differences in the and Peixoto (1989).<br />
floral morphology of the two taxa to a pollination Flexanthera was established by Rusby (1927). In<br />
syndrome. With Kirkbride's inclusion, Augusta is the protologue he stated that Flexanthera is "allied to<br />
represented by four species of rheophytic shrubs dis- <strong>Rustia</strong> [but without discussing their similarities],...<br />
tributed in Central America, Brazil, and islands of the from which it differs in its smaller flowers, reflexed<br />
South Pacific (Fiji and New Caledonia).<br />
and recurved dehiscent anthers [poricidal in <strong>Rustia</strong>]<br />
and deciduous calyx-limb and in the character of the<br />
Eosanthe<br />
placentae." Standley (1930a, 1931b) transferred the<br />
two known species of Flexanthera to Sickingia (Ron-<br />
Eosanthe was established by Urban (1923), who deletieae) without further comments, and Steyermark<br />
related it to Phyllomelia. Eosanthe is a monotypic (1972) transferred the species of Sickingia to Simira.<br />
genus endemic to the serpentine soils of Sierra de Flexantherafragrans Rusby and F. subcordata Rusby<br />
Cristal, Cuba (Borhidi, 1991; Delprete, 1999a). Its 4- are synonyms with Simirafragrans (Rusby) Steyerm.<br />
lobed persistent calyx functions in a similar manner and Simira cordifolia (J. D. Hook.) Steyerm., respecas<br />
the pterophyllous rotate calyx of Phyllomelia but, tively, and Flexanthera is synonymous with Simira.<br />
because of its general habit (small multi-stemmed The monotypic Exandra Standl. was described by<br />
shrubs; cf. Borhidi, 1991), ridged corolla tube and Standley (1923). This taxon was later treated by<br />
thick-coriaceous leaves, Eosanthe resembles Schmidt- Steyermark (1972) as congeneric with Simira, to which<br />
ottia (Catesbaeeae sensu Delprete, 1996d). On the it was transferred as S. rhodoclada (Standl.) Steyerm.<br />
other hand, the sheathing stipules, axillary inflores- The monotypic Holtonia Standl. was established<br />
cences, persistent 4-lobed calyx, filaments not by Standley (1932) with a species that he previously<br />
connated to the corolla tube, and 2-seeded indehiscent described in Sickingia (S. myriantha; Standley,<br />
fruits of Eosanthe resemble those of Phialanthus 1930a). He suggested that Holtonia is closely related<br />
(Chiococceae sensu Delprete, 1996d). In conclusion, to Chimarrhis and therefore belonged in the Condabecause<br />
of its resemblance to both Schmidtottia and mineeae (sensu Hooker). Holtonia has recently been<br />
Phialanthus, Eosanthe is here tentatively placed in the transferred to Elaeagia (Burger & Taylor, 1993) and<br />
Catesbaeeae-Chiococceae-complex, within which it a new combination [E. myriantha (Standl.) C. M.
Introduction to the <strong>Rondeletieae</strong> 17<br />
Taylor & Hammel] created, a decision with which I subglobose), both characters of little taxonomic sigagree.<br />
The suggested relationship with Chimarrhis nificance. General evidence strongly suggests that the<br />
(Standley, 1932) was appropriate, and corresponds to two taxa are congeneric, but the two species of<br />
my idea (Delprete, 1996b) that Elaeagia and Bathysa Pteridocalyx are known only from single flowering<br />
are closely related to Chimarrhis.<br />
specimens. Their mature fruits are unknown, and<br />
Wernhamia (Moore, 1922), traditionally placed in therefore fruiting collections are necessary to confirm<br />
the Cinchoneae and transferred to the Condamineeae their identity. Due to its probable synonymy with<br />
by Andersson and Persson (1991), has recently been Pteridocalyx, Neobertiera is not included in the key<br />
synonymized with Simira by Delprete and Nee (1997). to neotropical <strong>Rondeletieae</strong> given below.<br />
The taxa Wernhamia boliviensis S. Moore and Simira<br />
catappifolia (Standl.) Steyerm. are synonyms of<br />
Standleya and Bradea<br />
Simira macrocrater (K. Schum.) Steyerm.<br />
Blandibractea (Wernham, 1917), after careful The Brazilian endemics Standleya Brade and<br />
morphological comparison, was also treated as con- Bradea Standl. are closely related, and both belong<br />
generic with Simira by Delprete (1998c), B. to the Hedyotideae. They are similar in having tetbrasiliensis<br />
being synonymous with S. glaziovii (K.<br />
ramerous flowers, campanulate corollas, stamens in-<br />
Schum.) Steyerm.<br />
serted about the middle of the corolla tube, linear to<br />
ovate calyx lobes, few-seeded capsules, and ovoid<br />
seeds attached to a stalked<br />
The<br />
placenta. Standleya was<br />
Sipanea Group<br />
listed in the <strong>Rondeletieae</strong> by Robbrecht (1988, 1993)<br />
Limnosipanea, Sipanea, and Steyermarkia were and Delprete (1996d), and Bradea in the Hedyotideae<br />
transferred by Bremekamp (1934) from the Rondele- by Robbrecht (1988, 1993). Standleya differs from<br />
tieae to his newly founded tribe Sipaneeae. The only Bradea in having herbaceous habit (vs. subfruticose)<br />
morphological element that makes Sipanea and and unwinged seeds with tuberculate testa (vs. shal-<br />
Limnosipanea exceptional within the <strong>Rondeletieae</strong> is lowly winged with smooth testa). The tribal delimitheir<br />
herbaceous habit. Sipaneopsis and Neobertiera tation of the Hedyotideae, and their supposed relation-<br />
Wernham have habits that range from herbaceous (10 ships with the <strong>Rondeletieae</strong> are still uncertain, and<br />
cm tall) to subfrutescent (to 2-4 m tall), and could be need to be carefully analyzed.<br />
interpreted as the phylogenetic links between the<br />
Sipaneeae and the <strong>Rondeletieae</strong> s.str. Robbrecht<br />
Eizia<br />
(1988, 1993) maintained the Sipaneeae provisionally<br />
separate from the <strong>Rondeletieae</strong> because of their her- Eizia was listed by Delprete (1996d) in the Ronbaceous<br />
habit. In agreement with Verdcourt (1958) deletieae, but Lorence and Dwyer's (1988) observaand<br />
Steyermark (1974), the tribe Sipaneeae is here tions and recent personal studies have convinced me<br />
included in a broad circumscription of the Rondele- that this monotypic genus is better placed in the<br />
tieae.<br />
Hamelieae.<br />
Neobertiera and Pteridocalyx<br />
Pinckneya<br />
The monotypic Neobertiera was described by Pinckneya (Richard, 1803) is a monotypic genus<br />
Wernham (1917), who erroneously stated that its fruits endemic to the southwestern United States. It is the<br />
were berries and placed it in the Hamelieae. only North American genus included in the Rondele-<br />
Steyermark (1967: 289-290) corrected Wernham's tieae. Two Brazilian species, P. rubescens Allemao<br />
errors, declaring that the fruits of Neobertiera are & Saldanha and P. viridiflora Allemao & Saldanha,<br />
many-seeded capsules, and transferred the genus to were attributed to Pinckneya (Allemao & Saldanha da<br />
the <strong>Rondeletieae</strong>. The similarities between Gama, in Saldanha da Gama, 1872), and a third Bra-<br />
Neobertiera and Pteridocalyx Wernham (1911, 1913) zilian species, P. erubescens, was added to this geare<br />
striking, both having dichasial-cymose inflores- nus by Glaziou [in herb.]. Schumann (1888) transcences<br />
terminating in monochasial branchlets, con- ferred the Allemao & Saldanha species to Sickingia<br />
torted corolla aestivation, many-seeded capsules, [as S. rubra (Martius) K. Schum. and S. viridiflora<br />
subglobose seeds with reticulate exotesta, and both (Allemao & Saldanha) K. Schum.], and synonymized<br />
coming from low-elevation Guyanan rain forests; the Glaziou species under S. glaziovii K. Schum.<br />
Pteridocalyx differs from Neobertiera by the presence Steyermark (1972) transferred these three species to<br />
of calycophylls and by narrow, elongated fruits (vs. Simira, as S. glaziovii (K. Schum.) Steyerm., S. rubra
18 Flora Neotropica<br />
(Martius) Steyerm., and S. viridiflora (Allemao &<br />
Saldanha) Steyerm., respectively, leaving the one remaining<br />
species, P. bracteata (W. Bartr.) Raf., as a<br />
North American endemic. Additional information can<br />
be found in Delprete, 1996c.<br />
Steenisia (5 spp.) from Borneo, and Wendlandia (ca.<br />
56 spp.) widespread in Indomalesia. Augusta (including<br />
Lindenia) is the only genus present in both the<br />
Neotropics and Paleotropics, with one species in Central<br />
America, one endemic to Brazil, and two species<br />
in the Pacific Islands. The paleotropical Gloneria was<br />
Kerianthera<br />
listed by Delprete (1996d) in the <strong>Rondeletieae</strong>, but it<br />
is here excluded because recent personal observations<br />
Kerianthera J. H. Kirkbr. is a monotypic genus demonstrated that this genus is synonymous with<br />
endemic to the Brazilian Amazon. It was placed in the<br />
Psychotria L. (Psychotrieae).<br />
Condamineeae by Kirkbride (1985b) but, after phylogenetic<br />
and morphological studies, was transferred<br />
to the Isertieae by Delprete (1996e).<br />
<strong>Rondeletieae</strong> (A. P. de Candolle) Miquel, Flora<br />
Nederl. Indie 2: 130, 156. 1856. Rondeletiinae A.<br />
The Paleotropical Genera<br />
P. de Candolle (as "<strong>Rondeletieae</strong>," tribus<br />
Pseudomussaenda (5 spp., endemic to Africa) was<br />
Hedyotideae), Prodr. 342, 401. 1830. <strong>Rondeletieae</strong><br />
A. P. de Candolle ex Reichenbach, stat. non indic.,<br />
placed in the Condamineeae-Pinckneyinae by Der Deutsche Botaniker 1: 77. 1841.<br />
Wernham (1916) and later transferred by Verdcourt<br />
Type genus.<br />
Rondeletia Linnaeus.<br />
(1958) to the Isertieae. This transfer has recently been<br />
supported by Puff et al. (1993), who suggested a close Condamineeae J. D. Hooker (as "Condaminieae"), Gen.<br />
relationship to Schizomussaenda H. L. Li and PI. 8, 12. 1873. Condamineinae J. D. Hooker (as<br />
Mussaenda. Pseudomussaenda, Schizomussaenda, "Eucondaminieae," nom. inadmiss.), Gen. P1. 12.<br />
Mussaenda, and Kerianthera are exceptional within 1873. Type genus. Condaminea A. P. de Candolle.<br />
the Isertieae in having capsular fruits and Pinckneyinae J. D. Hooker (as "Pinckneyeae"), Gen. PI.<br />
calycophylls.<br />
12. 1873. Type genus. Pinckneya L. C. M. Richard<br />
Kajewskiella (endemic to the Solomon in<br />
Islands)<br />
A. Michaux.<br />
was established by Merrill and Perry (1947). It was Chimarrheae Bremekamp, nom. nud., Rec. Trav. Bot.<br />
Neerl. 31: 253. 1934.<br />
tentatively placed in<br />
Chimarrhidoideae<br />
the Condamineeae by Jansen<br />
Rafinesque<br />
(as "Sous-famille<br />
(1978) because of its valvate aestivation, absence of<br />
Chimarhidia"), Ann. G6n. Sci.<br />
Phys. 6: 84. 1820. Type genus. Chimarrhis Jacquin.<br />
raphides, and seed shape and position. Tange (1995) Simireae Bremekamp ex S. Darwin, Taxon 25: 606.<br />
found raphides in both species of Kajewskiella, which 1976. Simireae Bremekamp, sine descr. lat., Acta<br />
are in turn "indistinguishable from those found in Bot. Neerl. 15: 22. 1966. Type genus. Simira Aublet.<br />
Xanthophytum [Reinw. ex Blume]. These observa- Sipaneeae Bremekamp (as "Sipaneae"), Rec. Trav. Bot.<br />
tions lead to the conclusion that Kajewskiella belongs Neerl. 31: 253. 1934. Type genus. Sipanea Aublet.<br />
to the tribe Hedyotideae as defined by Robbrecht<br />
(1988), where together with Pomazota [Ridl.], Shrubs to tall canopy trees, rarely herbs or sub-<br />
Xanthophytum and Lerchea [L.] probably forms a shrubs; wood whitish, yellowish, or reddish; bark ofnatural<br />
group." (Tange, 1995). After studying flower ten containing quinine-related compounds. Raphides<br />
and seed morphology of Kajewskiella, I agree with absent. Stipules mostly entire, rarely bifid (four stipu-<br />
Tange that this genus should be included in the lar units in Condaminea), mostly interpetiolar or rarely<br />
Hedyotideae.<br />
intrapetiolar (Elaeagia), frequently with colleters on<br />
Badusa (Pacific Islands; cf. Soejarto et al., 1996) the adaxial side secreting resinous compounds, perwas<br />
transferred from the Cinchoneae to the Conda- sistent to readily caducous. Leaves opposite, decusmineeae<br />
(subtr. Portlandiinae) by Ridsdale (1982). sate (rarely in whorls of 3-4 leaves per node in a few<br />
Delprete (1996d) treated Badusa as related to the herbaceous taxa), petiolate to sessile; blades folia-<br />
Catesbaeeae sensu Delprete. Recent additional infor- ceous to thick-coriaceous; domatia variably present.<br />
mation about this genus can be found in Soejarto et Inflorescences terminal or axillary, often with 2-8<br />
al., 1996.<br />
orders of branching. Flowers hermaphroditic (dioe-<br />
The paleotropical genera belonging to the Ronde- cious in Dioicodendron), mostly actinomorphic (zyletieae,<br />
as presently circumscribed, are Aleisanthia (2 gomorphic in Molopanthera, and subzygomorphic in<br />
spp.) from the Malay Peninsula, Glionnetia (1 sp.) Cuatrecasasiodendron), 4-6-merous. Calyx persisfrom<br />
the Seychelles Islands, Greenea (12 spp.) from tent or caducous; lobes often minute, sometimes foli-<br />
Indomalesia, Spathichlamys (1 sp.) endemic to Burma, ose, frequently expanded into calycophylls. Corolla
Introduction to the <strong>Rondeletieae</strong> 19<br />
white, red, green, or yellow, membranous to fleshy;<br />
aestivation valvate, contorted, or imbricate. Stamens<br />
mostly as many as corolla lobes, inserted near the base,<br />
on the medial zone, or near the orifice of corolla tube;<br />
anthers included or exserted, oblong to narrowly el-<br />
liptic to button-shaped, 2-locular, commonly opening<br />
by longitudinal slits or rarely by terminal pores (<strong>Rustia</strong><br />
and <strong>Tresanthera</strong>), dorsifixed near the base or around<br />
the middle, introrse. Pollen exine reticulate or foveolate<br />
(not echinate). Style branches present, with stigmatic<br />
surface smooth to verrucate. Ovary inferior,<br />
bilocular, with few to many ovules per locule attached<br />
to a central placenta, or exceptionally 1 ovule per locule<br />
apically attached (Phyllomelia). Fruits woody,<br />
loculicidal or septicidal capsules, mostly opening<br />
basipetally (opening in central portion in Picardaea),<br />
or exceptionally pseudosamaras (Phyllomelia); pla-<br />
centa central, rarely apically incomplete, or shortly<br />
stalked. Seeds horizontal, imbricate, or peltate and<br />
vertical; winged, with lateral (Simira and<br />
Parachimarrhis) or concentric wing, or unwinged<br />
(apically inserted, ellipsoid-ovoid in Phyllomelia).<br />
The <strong>Rondeletieae</strong> and Condamineeae have traditionally<br />
been described as having many-seeded capsules.<br />
Nevertheless, the inclusion of Phyllomelia (related<br />
to Mazaea, see below), brings a new fruit type<br />
into the delimitation of the <strong>Rondeletieae</strong>: the<br />
pseudosamara (Spjut, 1994). In Phyllomelia the 2-<br />
seeded indehiscent fruits, with a persistent, rotate<br />
calyx, detach at maturity and function as a dispersal<br />
unit. I regard this fruit type as being the result of a<br />
derived set of characters that contributed to the adaptation<br />
to wind dispersal into this tribe.<br />
KEY TO THE GENERA OF NEOTROPICAL RONDELETIEAE S.L.<br />
1. Herbs to small subshrubs, usually 15 seeds per fruit; seeds angular, polyhedral; mouth of the<br />
corolla without triangular projections.<br />
4. Anthers exserted beyond the corolla; leaves in whorls of 3 or more per node (but leaves in<br />
pairs in L. schomburgkii J. D. Hook.) (Amazon basin) ............................................................. Limnosipanea<br />
4. Anthers included within the corolla; leaves in pairs (mostly South America, except S. biflora L.f.<br />
in Central America) ............................................................................. Sipanea<br />
1. Small shrubs to large canopy trees, >1 m tall.<br />
5. Stipules 4 per node, 2 intrapetiolar and 2 interpetiolar (Costa Rica to Bolivia) .................................... Condaminea<br />
5. Stipules 2 per node.<br />
6. Stipules intrapetiolar or calyptrate, mostly copiously exuding resin when still in bud, in several<br />
species solidifying into a globose mass at terminal bud (Central and South America, and E. cubensis<br />
Britt. endemic to Cuba) .............................................................................. Elaeagia<br />
6. Stipules interpetiolar, mostly scantly exuding resin when still in bud (copiously exuding resin in<br />
Chalepophyllum), not solidifying into a globose mass at terminal bud.<br />
7. One or more calyx lobes of some flowers expanded into 1 or 2 foliose extensions (calycophylls).<br />
8. Inflorescences axillary, subterminal, in pairs at the last two nodes below the apical bud,<br />
caducous at fruit maturity; seeds peltate on the central placenta, with central hilum (a few<br />
South American species with calycophylls) ............................................................. Chimarrhis<br />
8. Inflorescences terminal and solitary, persistent after fruit maturity; seeds not peltate on a<br />
central placenta, with lateral hilum.<br />
9. Corollas 2.5-8 cm long, long-tubular.<br />
10. Corolla tube with a ring of hairs at base inside; corolla lobes glabrous within;<br />
seeds not winged (mountains of Central and South America) .................................. Pogonopus<br />
10. Corolla tube without a ring of hairs inside; corolla lobes pubescent within;<br />
seeds winged (swamp areas of SE United States) ....................................................... Pinckneya<br />
9. Corollas 2 cm long; wood usually turning reddish when<br />
exposed to air (calycophylls present in several South American species) ...................... Simira
20 Flora Neotropica<br />
11. Seeds smaller,
Introduction to the <strong>Rondeletieae</strong> 21<br />
mouth of the tube, partially exserted; corolla tubes<br />
green and lobes red throughout anthesis; capsules<br />
thin-woody, oblong, 2.5-3.5 cm long; seed tetrahedral,<br />
1-2 mm long, unwinged (Amazonian slopes of the<br />
Andes from Colombia to Peru) ........................................ Macbrideina<br />
23. Corollas 2 cm<br />
long; wood usually turning reddish when exposed to<br />
air (Central and South America) . ........ ......................... Simira<br />
26. Seeds small, 2-3.5 mm long; capsules small,
22 Flora Neotropica<br />
entire margin; stipules obovate to ellipticobovate,<br />
15-30 mm long, foliaceous; seeds<br />
prominently winged (Colombia)<br />
......................................... Cuatrecasasiodendron<br />
33. Corollas actinomorphic, 4-5-lobed; the<br />
innermost lobe equal in size to the other,<br />
with entire margin; the outer 1 to 2 lobes<br />
undulate-crenulate; stipules deltoid to<br />
lanceolate, 3 cm long.<br />
35. Corollas narrowly campanulate, red; corolla tube narrowly imbutiform<br />
(Cerrado and Atlantic forests of Brazil) ... Augusta subgen. Augusta<br />
35. Corollas hypocrateriform, white; corolla tube narrowly tubular.<br />
36. Calyx lobes linear; seeds with lateral hilum (truncate at apex),<br />
not winged, horizontally attached to the central placenta<br />
(Central America and South Pacific Islands) .. Augusta subgen. Lindenia<br />
36. Calyx lobes oblong-obovate; seeds with central hilum,<br />
with shallow concentric wing, vertical, peltately attached<br />
to the central placenta (Venezuelan tepuis) ........... Maguireothamnus<br />
34. Corollas 2 x 1 cm.<br />
38. Ovary 4-locular, with 5-6 ovules per locule; mature<br />
fruits dividing into 4 cocci (Colombia) .............................. Acrobotrys<br />
38. Ovary 2-locular, with numerous ovules per locule (>6);<br />
mature fruits not dividing into 4 cocci (but old capsules of<br />
Chalepophyllum<br />
splitting into 4 parts, though not into 4 cocci).<br />
39. Inflorescences axillary (not subterminal), at the axils<br />
of the last 5-7 nodes below the apical bud, in congested<br />
cymules of 1-4 flowers.<br />
40. Corollas hypocrateriform, white (the tubes<br />
sometimes suffused with pinkish-reddish tinge<br />
at base); resinous exudates from young stipules<br />
copious; calyx lobes narrowly oblong to narrowly<br />
lanceolate, subcoriaceous, unequal, 1-3 mm wide,<br />
fully expanded before anthesis; seeds with a narrow<br />
marginal expanded wing, 3-4 mm long (high<br />
elevations of Venezuela and Guianas)......... Chalepophyllum<br />
40. Corollas narrowly campanulate, salmon-red;<br />
resinous exudates from young stipules scant;<br />
calyx lobes lanceolate, membranous, subequal,<br />
4-6 mm wide,expanding after anthesis; seeds<br />
unwinged, to 1 mm long (high elevations of<br />
Amazonian Venezuela) ...................................... Holstianthus<br />
39. Inflorescences terminal, in corymbose panicles of<br />
9-40 flowers.<br />
41. Inflorescences 10 cm long; flowers subcampanu-<br />
late; calyx lobes widely ovate; stipules readily<br />
caducous (Colombia and Peru) ................................. Phitopis
Introduction to the Neotropical Genera Studied 23<br />
NOTES ON DESCRIPTIONS, (20-30 m) with much expanded buttresses in South<br />
ABBREVIATIONS, AND ACRONYMS American species. Some Amazonian species of<br />
Chimarrhis (C. glabriflora, C. hookeri, C. barbata,<br />
In the preparation of herbarium specimens, veg- and C. turbinata) can grow into forest giants, 40-50<br />
etative and reproductive parts are subjected to minor m tall and 2 m diam.<br />
shrinkage. Measurements given in keys and descrip- The tree architecture of this group has been studtions<br />
have been obtained from such specimens. Col- ied in only a few species of Chimarrhis. Halle et al.<br />
ors are from personal observations and label data, (1978) reported that C. cymosa (from the Lesser<br />
unless otherwise indicated. The abbreviation "BA"<br />
Antilles) has a Petit model of branching, while C.<br />
represents the basal angle from the leaf margin to the<br />
microcarpa (from eastern Amazon) has an Aubreville<br />
midrib of the leaf blade, and "L/W" symbolizes the model ("Terminalia branching"). I have observed C.<br />
length to width ratio of the leaf blades. Nomenclature<br />
glabriflora and C. hookeri (both from western Amaof<br />
leaf domatia follows Robbrecht (1988: fig. 12), and zon forest), and both have Petit's model.<br />
that of vestiture types follows Radford et al. (1974: Large buttresses are present only in the large trees<br />
140-141). In the descriptions of inflorescence archi- of the Amazonian species of Chimarrhis, extending<br />
tectures, "L/A" is the ratio of the total length (L) to to 4 m in height near the trunk and expanding outthe<br />
length of the branched portion (A). The general ward 8-10 m from the trunk (per. obs.). Additionally,<br />
shape and vestiture of the disk, and whether the disk at least two arborescent species of <strong>Rustia</strong> (R. alba and<br />
exceeds the calyx, refer to mature capsules, unless R. rubra) can develop small buttresses.<br />
otherwise indicated. The term "microscopically (40x)" Lateral roots are occasionally present in the genmeans<br />
that the structures or characteristics referred to era studied. <strong>Rustia</strong> occidentalis, R. thibaudioides, and<br />
can be seen using a dissecting microscope at 40x<br />
Pogonopus speciosus (all multistemmed shrubs growmagnification.<br />
All specimens cited, including types, ing in proximity to streams or standing freshwater)<br />
have been seen unless indicated by "n.v." immediately have been personally observed to produce lateral roots<br />
following the citation. Specimens cited as "B*" were at the basal nodes of the external branches. The three<br />
destroyed in the Berlin herbarium during the Second<br />
species of Pogonopus are very ornamental shrubs due<br />
World War (Hiepko, 1987). Herbarium acronyms to their colourful calycophylls, and their propagation<br />
follow Holmgren et al., 1990. Authors of plant names could be easily accomplished through cuttings of<br />
have been abbreviated throughout the text according young woody branches.<br />
to Brummitt & Powell, 1992. Genera included in the<br />
present systematic treatment are in boldface in the key.<br />
Leafy Branchlets<br />
INTRODUCTION TO THE<br />
Leafy branchlets are here referred to as the terminal<br />
NEOTROPICAL GENERA STUDIED portion of the vegetative shoots of the last nodes<br />
(commonly known as "twigs"). Leafy branchlets are<br />
GENERAL MORPHOLOGY commonly terete in all genera, with Dioicodendron<br />
being exceptional in having subtetragonal leafy<br />
Habit<br />
branchlets. Young leafy branchlets vary among spe-<br />
The Condamineeae sensu Hooker (Robbrecht, cies of the same genus in being either thin and woody<br />
1988, 1993) is a paraphyletic group of genera repre- or thick and semisucculent. I consider these two catsented<br />
by shrubs, small trees, and tall canopy trees egories of vegetative shoots as a reliable taxonomic<br />
(i.e., Chimarrhis). The genera Pogonopus, Condaminea, character, which is quite consistent at the specific<br />
Picardaea, and <strong>Rustia</strong> are most commonly found as level, mostly in Chimarrhis and secondarily in <strong>Rustia</strong><br />
shrubs. Several species of <strong>Rustia</strong> (R. costaricensis, R. and <strong>Tresanthera</strong>.<br />
dressleri, R. alba, R. bilsana, R. rubra, R. viridiflora,<br />
R. venezuelensis, and rarely R. formosa), Dioico-<br />
External Indumentum<br />
dendron, Dolichodelphys, and Molopanthera are usually<br />
encountered as medium-sized trees 10-15 m tall The indumentum of the genera here treated, when<br />
(exceptionally to 20-30 m). <strong>Tresanthera</strong> is usually present, consists of unicellular, nonglandular hairs. It<br />
encountered as pyramidal single-stemmed trees. is extremely variable among and within populations<br />
Trees of Chimarrhis vary from medium-sized [10- of the same species (<strong>Rustia</strong> thibaudioides, R. alba,<br />
20(-30) m tall] in Central America and the Caribbean pers. obs.; Condaminea corymbosa, pers. obs.; Pogon-<br />
Islands (C. parviflora, C. latifolia, C. cubensis, C. opus speciosus var. sandwithianus, pers. obs.; Pinckjamaicensis,<br />
and C. cymosa) to large canopy-trees neya bracteata, pers. obs.) and even among the foli-
24 Flora Neotropica<br />
age of a single individual (<strong>Rustia</strong> thibaudioides, pers.<br />
Domatia<br />
obs.; Condaminea corymbosa, pers. obs.; Pogonopus Domatia are here defined as presence of hairs,<br />
speciosus, pers. obs.; Chimarrhis hookeri, pers. obs.). correlated or not with a pit or a crypt, in the axils of<br />
For the above reasons the occurrence, density, and<br />
the secondary veins, or rarely of the tertiary veins of<br />
quality of the indumentum have been largely considthe<br />
leaf blades. Domatia are commonly associated<br />
ered characters of little taxonomical significance. with the presence of mites. In many of the species of<br />
Almost all the genera studied have species with wide the genera studied, larval and adult stages of unidengeographic<br />
distributions that have been described as tified mites have been found.<br />
a complex of species based on their variable The general classification of leaf-domatia in the<br />
indumentum. Standley (1918, 1921, 1930a, 1930b,<br />
present treatment (and in the terminology used in the<br />
1931 a, 1931b, 1931 c, 1934a, 1934b) and Steyermark<br />
species descriptions) is in accordance with Robbrecht's<br />
(1963, 1964b, 1965, 1967, 1972, 1974, 1975) are<br />
typology (1988: 50).<br />
among the many botanists who have especially em-<br />
Leaf domatia have not been found (and are most<br />
phasized the density and quality of the indumentum<br />
probably absent) in <strong>Tresanthera</strong>, Condaminea, Dioof<br />
vegetative parts in describing new species and new<br />
icodendron, Molopanthera, and Dolichodelphys; and<br />
varieties of <strong>Rubiaceae</strong>. Nevertheless, in some very are found to be variably absent or present in <strong>Rustia</strong>,<br />
special cases, where a good number of collections<br />
Pogonopus, Chimarrhis, and Picardaea. The absence<br />
were available, and very distinct forms of indument or presence, morphology, and vestiture ofdomatia was<br />
were consistently present and were correlated with a<br />
studied in detail and found to be very variable within<br />
particular area, I have used vestiture as a secondary<br />
species, within populations of the same species, and<br />
taxonomic character at the varietal rank.<br />
even within the same individual. Domatia are therefore<br />
considered a character of secondary taxonomical<br />
Leaves<br />
importance, to be used, at the most, at the specific level.<br />
The leaves of woody <strong>Rubiaceae</strong> are mostly oppo- Nevertheless, several species of Chimarrhis dissite,<br />
rarely ternate or whorled, with stipules constantly play a certain degree of consistency in the type of<br />
present, and entire blades; these three vegetative fea- domatia, and some species can be recognized solely<br />
tures make the family easy to recognize even with by the presence of their characteristic domatia (see<br />
sterile material. Opposite leaves are usually decussate. Fig. 58). The most typical domatia are those of C.ja-<br />
In the genera here treated, the leaves are opposite, maicensis, unique in the genus in being tuft-pit domatia,<br />
decussate, and petiolate. The leaves of Condaminea with hairs departing from the veins and minutely pitcorymbosa<br />
are exceptional, varying from sessile (in ted at the axils of secondary veins, and rarely of ter-<br />
Central America) to petiolate (in South America) (see tiary veins. Chimarrhis cubensis has minutely winged<br />
Fig. 47). Petioles are mostly adaxially flattened to veins, forming "pocket-domatia" in axils of secondconcave,<br />
thickened at the base, and apulvinate. How- ary veins. In C. latifolia, C. brevipes, C. gentryana, and<br />
ever, in two species of <strong>Rustia</strong> (R. occidentalis and R. C. barbata the blades and the veins are entirely glathibaudioides)<br />
I have observed rudimentary pulvini brous, and no domatia have been observed. In the rest<br />
on petioles of young leaves, which become thicker of the species of Chimarrhis, domatia were variably<br />
(sometimes corky) and nonfunctional on older leaves. present or absent and of various kinds (see Fig. 58).<br />
Blades are always entire and mostly stiff-foliaceous,<br />
but coriaceous in Condaminea, and semicoriaceous<br />
Foliar Pellucid Glands<br />
in <strong>Tresanthera</strong> and in some species of <strong>Rustia</strong> (R. Solereder (1893), in his Anatomischen Charakdressleri,<br />
R. alba, R. bilsana, R. venezuelensis, R. teristik und zur Systematik, reported the presence of<br />
formosa, R. simpsonii, R. angustifolia, and variably "inneren Drilsen" [internal druses] in the <strong>Rubiaceae</strong>,<br />
so in R. thibaudioides).<br />
which he found previously described as "intercellu-<br />
Venation of all the genera studied is brochido- lare Secretbehilter" ["intercellular secretion-containdromous,<br />
with secondary veins ascending, arcuate, ers"], "secretfiihrende" [secretory cells] of differing<br />
and anastomosing near the blade margins. Bacterial forms and various contents, and also as "eingenthiimnodules<br />
are found in blades of <strong>Rustia</strong> (R. occidentalis), liche Drisen" ["characteristic druses"] present in<br />
Pogonopus (P. speciosus var. sandwithianus, being some taxa in the form of secretory cells. Such secrepunctate<br />
on the adaxial surface due to bacteria), and tory glands have recently been called "pellucid<br />
Chimarrhis (C. glabriflora and C. hookeri). To my glands" (Steyermark, 1974; Dwyer, 1980), "transluknowledge,<br />
these are the first reports of bacterial nod- cent dots" (Robbrecht, 1988), or "pellucid-dots"<br />
ules in neotropical <strong>Rubiaceae</strong>.<br />
(Burger & Taylor, 1993).
Introduction to the Neotropical Genera Studied 25<br />
In the <strong>Rubiaceae</strong>, schizogenous secretory cavities, biaceae. Unfortunately, stipules are sometimes readily<br />
filled with a resinous secretion, have been found only caducous, and in that case they can only be found at<br />
in the leaves of the closely related genera <strong>Rustia</strong> and the youngest nodes (and sometimes only at the apical<br />
<strong>Tresanthera</strong> (which are also unique in the family in bud). I found that caducous or persistent stipules are<br />
having poricidal anthers). To my knowledge no de- consistent within each genus of the Condamineeae<br />
tailed work has been done on the anatomy of such sensu Hooker (except for Chimarrhis; see below). For<br />
secretory structures, nor has any attempt been made descriptive purposes the stipule persistency is here<br />
to identify the content of their cavities (Delprete et divided into three degrees: persistent (present after<br />
al., in progress). The only overview on pellucid-punc- leaves of the same node fall off), readily caducous<br />
tate leaves was presented by Solereder (1893: 279; (present only at the vegetative bud), and caducous<br />
free translation from German) as follows: (present during leaf development but caducous before<br />
leaves fall<br />
Intercellular secretory space, in the form of<br />
off). Of the genera here presented, the<br />
schizogenous secretory cavities filled with<br />
stipules are readily caducous in <strong>Rustia</strong> and Tressticky<br />
[resinous] secretion, lined with a thin<br />
anthera, caducous in Condaminea and Pinckneya, and<br />
walled epithelium, are found in <strong>Rustia</strong> persistent in Pogonopus, Picardaea, Dioicodendron,<br />
(incl.<br />
Henlea) and <strong>Tresanthera</strong>, which were united Molopanthera, and Dolichodelphys. Chimarrhis is<br />
by Bentham and Hooker [1873] in a exceptional to the group in having most species with<br />
single genus<br />
(<strong>Rustia</strong>), which I already noted briefly in stipules readily caducous (subgen. Chimarrhis), and<br />
my paper on Hymenocnemis in Bot. Centralbl.<br />
a few species with persistent stipules (subgen.<br />
1891. These secretory cavities occur in the spe-<br />
Pseudochimarrhis).<br />
cies of <strong>Rustia</strong> and <strong>Tresanthera</strong> in large leaves, Stipular position is also an important taxonomic<br />
usually as translucent points, which were alcharacter<br />
in the <strong>Rubiaceae</strong>. All the genera here treated<br />
ready described by Karsten [1858, 1861; cf.<br />
have two interpetiolar stipules, except for Conda-<br />
Karsten, 1858-1869] for <strong>Rustia</strong> in general, for minea, which is commonly described as having four<br />
<strong>Tresanthera</strong> condamineoides and Henlea interpetiolar and intrapetiolar stipules. The nature of<br />
splendens [= <strong>Rustia</strong> thibaudioides], and the<br />
by stipules in Condaminea is rather ambiguous, be-<br />
Bentham [1844] for Exostem[m]a occidentale ing all basally connate and not truly intrapetiolar (Fig.<br />
justly transferred to <strong>Rustia</strong> by Hemsley [1881 ]), 3F-H), because no stipules are positioned between the<br />
and by Schumann (1889, 1891) [incorrectly de- stem and the petioles. The stipules of Condaminea<br />
scribed as "characteristic fat bodies"] for could also be interpreted as two interpetiolar stipules<br />
<strong>Rustia</strong>. I have described the secretory cavities highly dissected and connate at the base. To underfor<br />
certain available species, namely T. conda- stand fully the position of stipules in Condaminea,<br />
mineoides, R. angustifolia, R.formosa, R. gra- developmental studies are needed.<br />
cilis, R. occidentalis, R. secundiflora, R. The most common shape of the stipules is trianwarczewicziana<br />
[two former = R. thi- gular with entire margins (Fig. 3), varying from delbaudioides],<br />
as well as R. pauciflora [= T. toid to narrowly triangular, with the exception of<br />
pauciflora] described by me, which Schumann Condaminea (narrowly lanceolate and foliose) and<br />
[in contrast to Bentham & Hooker (1873)] Dioicodendron (obovate stipules). Stipule size in the<br />
treated in a distinct genus, <strong>Tresanthera</strong> genera here studied varies from small and scale-like<br />
pauciflora.<br />
(Picardaea) to small and deltoid (Pogonopus, Pinckneya,<br />
Molopanthera, and Chimarrhis turbinata) to<br />
I examined all the species of <strong>Rustia</strong> (incl. Henlea) medium-size and narrowly triangular (<strong>Rustia</strong>, most<br />
and <strong>Tresanthera</strong> using light microscopy, and all of<br />
species of Chimarrhis, and Dolichodelphys) to large<br />
them have pellucid glands. Such glands represent a<br />
and foliose (<strong>Tresanthera</strong>, Condaminea, Chimarrhis<br />
good field character to easily identify these two rubicymosa).<br />
aceous genera. The density and size of the pellucid<br />
glands in the blades is variable (rarely consistent<br />
within the same species), and for the most part it is<br />
Colleters<br />
not a reliable taxonomic character.<br />
The term colleters is "derived from the Greek Colla<br />
[= glue], referring to the sticky excretion from this<br />
Stipules<br />
structure," and "they frequently produce a mixture of<br />
Stipules are the paramount feature of the family terpenes and mucilage" (Esau, 1965). Colleters are<br />
and represent one of the easiest and most apparent commonly defined as "pluricellular mucilage-secretfield<br />
characters for recognizing members of the Ru- ing structures occurring inside stipules and on other
26<br />
A<br />
'<br />
Flora Neotropica<br />
FIG. 3. Stipule and colleter variation in the genera referred to the Condamineeae (sensu Hooker, 1873). A-C. <strong>Rustia</strong>.<br />
A. R. occidentalis (Delprete 5267, TEX), adaxial view. B. R. alba (Delprete & Verduga 6420, TEX), adaxial view. C. R.formosa<br />
(Kirkbride 5431, F), adaxial view. D. Kerianthera preclara (Prance et al. 24293, NY), adaxial view. E. <strong>Tresanthera</strong><br />
condamineoides var. condamineoides (Steyermark & Rabe 96163, US), adaxial view. F-H. Condaminea corymbosa (Delprete<br />
& Webster 6104, TEX). F. View from above showing stipular insertion. G. Detail of base of stipule showing colleters in<br />
adaxial view. H. Habit of stipules. I, J. Pinckneya bracteata (Delprete & Sharif 6472, TEX). I. Adaxial view. J. Lateral<br />
view of stipule insertion (white area is petiole attachment). K, L. Pogonopus speciosus var. sandwithianus (Delprete &<br />
Apreza 6359, TEX). K. Adaxial view. L. Lateral view of stipule insertion and tuft of hairs in stipules' axils (white area is petiole<br />
attachment). M. Picardaea cubensis (Shafer 7850, NY), adaxial view. N. Kajewskiella trichantha (Kajewski 1667, A),<br />
adaxial view. O-Q. Chimarrhis. 0. C. cymosa (Webster et al. 9205, DAV), adaxial view. P. C. glabriflora (Delprete &<br />
Verduga 6423, TEX), abaxial view. Q. C. turbinata (Irwin et al. 55443, GH), adaxial view. R. Dioicodendron dioicum<br />
(Hammered et al. 5233, NY), adaxial view. S. Dolichodelphys chlorocrater (Ramirez & Cardenas 1017, COL), adaxial view. T.<br />
Molopanthera paniculata var. paniculata (Mori et al. 10328, NY), adaxial view. All drawn to the same scale, except G.
Introduction to the Neotropical Genera Studied 27<br />
organs" (Robbrecht, 1988). Additional information<br />
Inflorescences<br />
about colleter structure and function can be found in The terminology and classification of inflores-<br />
Esau, 1965 (311); Halle, 1967 (95-97); Lersten, cences in the present work has been heavily influenced<br />
1974a, 1974b, 1975; Robbrecht, 1988 (64-68); and by the recent work of Weberling (1992). More spe-<br />
Rogers, 1987.<br />
cifically, Weberling (1977) presented a comprehen-<br />
Colleters in the <strong>Rubiaceae</strong> are most commonly sive review of the generalized typology of <strong>Rubiaceae</strong><br />
found inside of the stipules, usually at the basal por- inflorescences, and diagrammatic representations can<br />
tion, and are also frequently found inside the calyx, be found in Delprete (1996d: fig. 2B). When differat<br />
the base of the calyx lobes or alternate to them. ent interpretations on inflorescence morphology were<br />
For the group of genera under study, colleters have presented by various authors, Weberling's (1977,<br />
been consistently found inside the stipules at the basal 1992) conclusions were used.<br />
portion.<br />
Inflorescence position, in the genera studied, is<br />
When apical stipules are about to open, their in- most commonly terminal, with the exception of<br />
ternal colleters begin to secrete a resinous material Chimarrhis (axillary and subterminal). I consider the<br />
which is believed to protect the young vegetative buds axillary inflorescences of Chimarrhis to be the result<br />
against herbivory. The presence of"sticky" resinous of extreme reduction of the last terminal nodes<br />
exudates in vegetative buds has been observed in (Weberling, 1977, 1992; Robbrecht, 1988).<br />
<strong>Rustia</strong>, Condaminea, Pogonopus, and Chimarrhis. I <strong>Part</strong>icular attention should be called to the infloconsider<br />
absence or presence, arrangement, and gen- rescences ofPogonopus, Molopanthera, and Dioicoeral<br />
morphology of stipular colleters significant taxo- dendron, which have been variously described as ternomic<br />
characters, especially when used at the generic minal or terminal and lateral. Standley (1918, 1930a,<br />
and specific levels (Fig. 3).<br />
1931 b, 1931 c) described the inflorescence of Pogon-<br />
In <strong>Rustia</strong>, stipular colleters are present in a trian- opus as "cymes arranged into terminal leafy panicles";<br />
gular area on the base in R. occidentalis (Fig. 3A), in Steyermark (1974) as [literally translated from Spana<br />
single basal row in R. alba (Fig. 3B), or as a narrow ish] "densiflorous subcorymbose-cymose with<br />
basal area in R.formosa (Fig. 3C). In Picardaea (Fig. plurichothomous ramification"; Dwyer (1980) as "ter-<br />
3M) stipular colleters are present in a single basal row. minal paniculate and with small lateral cymes,"<br />
In <strong>Tresanthera</strong> (Fig. 3E) colleters are long-slender and though he described P. speciosus as having "inflorespresent<br />
on the base. In Condaminea (Fig. 3G) colleters cences terminal and axillary and thyrsoid-cymose";<br />
are present in an irregular basal area on each of the Burger and Taylor (1993) [as in Dwyer, 1980] defour<br />
stipular units. In Pogonopus (Fig. 3K,L) the scribed Pogonopus with "inflorescences terminal,<br />
colleters are in one to two irregular basal rows and paniculate and subcorymbose-cymose," but P.<br />
intermixed with sericeous hairs. In Chimarrhis (Fig. speciosus with "inflorescences terminal or axillary."<br />
30-Q) the position of the colleters is variable, but In my opinion (and according to Weberling, 1992),<br />
usually present on a basal area in C. cymosa (Fig. 30), Pogonopus, Molopanthera, and Dioicodendron have<br />
in a single basal row in C. glabriflora (Fig. 3P), or on terminal frondose paniculate inflorescences, with cya<br />
densely packed narrow basal area in C. turbinata mose branches subtended by foliose bracts, similar to<br />
(Fig. 3Q). In Dioicodendron (Fig. 3R) the stipular foliage leaves. The basal "leaves" (pherophylls) subcolleters<br />
are sparsely distributed in the basal area and tending the inflorescence branches are identical in size<br />
intermixed with golden densely pubescent hairs. In and shape to foliage leaves, and are gradually smaller<br />
Dolichodelphys (Fig. 1S) the colleters are sparsely toward the distal portion of the inflorescence. In gendistributed<br />
on the basal portion of the stipule. In eral, the presence of"leaves" (pherophylls) in termi-<br />
Molopanthera (Fig. 3T) only a few colleters are nal inflorescences of the <strong>Rubiaceae</strong> has led the above<br />
present at the base of the stipules and intermixed with cited workers to incorrectly describe the infloresgolden-sericeous<br />
hairs. [In Pinckneya (Fig. 3I,J) stipu- cences as lateral and cymose instead of terminal and<br />
lar colleters are sparsely distributed in a basal trian- paniculate. This argument was convincingly made by<br />
gular area. In Kerianthera (Fig. 3D) the colleters are Troll (1950) and again by Weberling (1977, 1992).<br />
sparsely distributed in an ovate area up to half the Inflorescence position of Chimarrhis is unique<br />
length of the stipules and (as in Pogonopus) are in- among the genera studied in having what I call "axiltermixed<br />
with gold-sericeous hairs. In Kajewskiella lary subterminal inflorescences," being borne in pairs<br />
(Fig. 3N) the colleters are long-slender, in a basal area in the leaf axils on the last terminal nodes. In my field<br />
one-third the length of the stipule and intermixed with observations of Amazonian species (i.e., C. glabrisericeous<br />
hairs].<br />
flora, C. hookeri, C. barbata, and C. duckeana), the
28 Flora Neotropica<br />
apical shoot stops growing during anthesis and fruit<br />
maturation, and the axillary inflorescences are<br />
abscissed just after seed dispersal (sometimes even<br />
before capsules dehiscence), immediately resuming<br />
apical shoot growth (see phenology of C. glabriflora).<br />
In some species growing in Central America and the<br />
Caribbean region, where the climate is markedly seasonal,<br />
the inflorescences sometimes remain attached<br />
to the branchlets up to one year after seed dispersal<br />
(C. parviflora, C. cymosa, and C. microcarpa). The<br />
axillary-subterminal inflorescence position of<br />
Chimarrhis is a good taxonomic character that easily<br />
separates it from related genera.<br />
The inflorescence morphology of the taxa presented<br />
here is quite variable. The terminal panicle is<br />
the most common, with architecture variable among<br />
and within genera. The panicles of <strong>Rustia</strong> vary from<br />
many-branched pyramidal and once- to thrice-compound<br />
(R. costaricensis, R. alba, R. bilsana, R. rubra,<br />
R. viridiflora, R. venezuelensis, R. formosa, R. simpsonii)<br />
to few-branched and reduced (R. occidentalis,<br />
R. dressleri, R. gracilis) to few-branched and longpedunculate<br />
(R. angustifolia) to racemoid or thyrsoid<br />
secundiflorous (R. schunkeana, R. thibaudioides).<br />
ers are perfect in all genera examined, except in Dio-<br />
icodendron, which has individuals with either male<br />
or female flowers only.<br />
In most of the genera studied the flowers are acti-<br />
nomorphic, except for Condaminea and Molopan-<br />
thera. The anthers of Condaminea are reflexed toward<br />
the lower base of the corolla tube and all open upward,<br />
forming a landing area for visitors/pollinators. The<br />
flower buds of Molopanthera are zygomorphic, curving<br />
upward; its anthers and corolla lobes are longer in<br />
the lower portion of the minute flowers; the two lower<br />
stamens have filaments curved upward, and their anthers<br />
often remain connected by the cuneate extensions<br />
present at both ends (forming a cup-shaped unit).<br />
The flowers of this group are usually presented in<br />
an erect position, with the exception of some pendulous-flowered<br />
species of <strong>Rustia</strong> (R. thibaudioides, R.<br />
schunkeana, R. viridiflora, and R. rubra; see Delprete,<br />
1996a). The floral morphology in <strong>Rustia</strong> is very diversified,<br />
in response to the process of coevolution<br />
with a variety of pollinators. This speciation has probably<br />
been facilitated by <strong>Rustia</strong>'s poricidal anthers,<br />
which can be variably exserted or included, adapting<br />
to the several pollination syndromes found in this<br />
<strong>Tresanthera</strong> inflorescences vary from densely thyrs- genus. In R. costaricensis the flowers are minute,<br />
oid panicles (T. condamineoides var. thyrsiflora) to erect, and exserted spreading anthers, and are pollipauciflorous<br />
thyrses (T. condamineoides var. conda- nated by small bees (lap-pollination). The fleshy flowmineoides).<br />
Condaminea have terminal, long-pedun- ers ofR. occidentalis, R. bilsana, and R. dressleri have<br />
culate, corymbose panicles, once- to twice-compound. anthers only half-exserted and convex toward the cen-<br />
Pogonopus, Dioicodendron, and Molopanthera have ter, which are instead buzz-pollinated by bees (perterminal<br />
frondose panicles with corymbose branches sonally observed in R. occidentalis).<br />
(tertiary thyrsoid in Molopanthera). Picardaea has<br />
short-pedunculate, corymbose, trichotomous panicles<br />
Calyces<br />
terminating with triflorous cymules. Dolichodelphys All the genera studied have gamosepalous calyces<br />
has laxly branched panicles with many secondary cy- positioned at the apices of the ovaries. The general<br />
mose branches.<br />
shape of the calyces varies from small-cupular to ex-<br />
Inflorescence branches of the genera studied are comtremely<br />
reduced, but this character is consistent at<br />
monly opposite to subopposite and decussate, and the most at the specific level. Calyx lobes vary from easrachis<br />
is usually basally terete and decussately comily<br />
distinguishable to extremely reduced (i.e., a wavy<br />
pressed distally, where the lateral branches are inserted.<br />
margin) or absent. The calyces are reduced to a mi-<br />
Inflorescence bracts subtending primary and secnutely<br />
undulate margin (sometimes with barely visondary<br />
branches are consistently present, and vary ible lobes) in <strong>Rustia</strong>, <strong>Tresanthera</strong>, Pogonopus,<br />
from leaf-like (pherophylls) to extremely reduced and Chimarrhis, and Dioicodendron. In Condaminea,<br />
deltoid, to narrowly linear. The bracteoles subtend- Picardaea, Molopanthera, Dolichodelphys, and<br />
ing flowers are not constantly present, are always Parachimarrhis the calyces are small-cupular and<br />
minute, and vary from deltoid to linear.<br />
vary from truncate to obviously lobed.<br />
The persistence of the calyces in mature fruits is<br />
Flowers<br />
an important taxonomic character for the taxonomic<br />
The floral morphology of the genera here treated delimitation of the genera here examined. The calyis<br />
rather variable. Flowers are protandrous in all the ces are consistently persistent in mature capsules of<br />
genera except for Chimarrhis, where the receptive <strong>Rustia</strong>, <strong>Tresanthera</strong>, Chimarrhis, Dioicodendron,<br />
stigmas protrude above the still-closed corollas (for Dolichodelphys, and Parachimarrhis, and caducous<br />
more details see discussion under Chimarrhis). Flow- in those of Condaminea, Pogonopus, and Picardaea.
Introduction to the Neotropical Genera Studied 29<br />
Calycophylls<br />
Calycophylls are defined as the expansion of one<br />
or more calyx lobes into colorful foliose units that serve<br />
mainly to attract visitors/pollinators, and probably to<br />
effect seed dispersal in mature capsules (Delprete,<br />
1966b). The oft-used term "semaphyll" was coined by<br />
Leppik (1956) and refers to "all colored leaves ofplants,<br />
like petals, sepals, bracts, ligulate flowers, etc., which<br />
serve to attract pollinators," and therefore is not specific<br />
to calyx lobes. In the <strong>Rubiaceae</strong> the presence of<br />
calycophylls is usually correlated with capsular fruits,<br />
variably occurring in many genera (Robbrecht, 1988;<br />
Delprete, 1996b), and which I treated as a secondary<br />
taxonomic character at the generic and specific ranks.<br />
A survey of the presence and general morphology of<br />
calycophylls in <strong>Rubiaceae</strong> was recently published by<br />
Classen-Bockhoff (1996). As discussed elsewhere<br />
(Delprete, 1996b, 1996c, 1996e, 1997b, 1998b, 1998c),<br />
some genera that were previously separated from their<br />
sister taxa based on the presence of calycophylls need<br />
to be reconsidered and probably returned to their origividuals<br />
growing in undercanopy and shady places<br />
have white to cream-white calycophylls, while the<br />
individuals in sunny exposures have pink to pale-lilac<br />
calycophylls.) In Chimarrhis the calycophylls are<br />
cream-white to greenish white (as in Calycophyllum).<br />
In Pogonopus (and Pinckneya), as the capsules<br />
approach maturity the calycophylls gradually lose<br />
their color, drying out, becoming brown, and are<br />
sometimes eliminated by breaking off of their driedup<br />
fragile stalks, or persisting on the mature capsules.<br />
(I speculated that in some cases the dried-up<br />
calycophylls of Pogonopus remain attached to the<br />
mature capsules while the seeds are released, probably<br />
assisting in the dispersal of the minute seeds by<br />
shaking the capsules as a result of wind turbulence.)<br />
In Parachimarrhis several flowers per inflorescence<br />
have a small calycophyll. The calycophylls of<br />
this genus are white to cream-white and 7-13 mm<br />
long, with the abaxial side covered with a dense mat<br />
of glandular hairs. The function of this mat of hairs,<br />
not seen in any other rubiaceous genera, is unknown.<br />
nal taxonomic position. For example, Chimarrhis has<br />
been historically described as lacking calycophylls, but<br />
Corollas<br />
the newly described C. gentryana (Delprete, 1996b) dis- The corollas of this group vary considerably in<br />
plays an abundance of cream-colored calycophylls (for size, shape, and thickness among and within genera<br />
which it has been often confused with Calycophyllum), (Fig. 2), and are commonly actinomorphic (except<br />
which are also sporadically found in C. brevipes, C. Condaminea and Molopanthera). The Condamineeae<br />
barbata, and C. turbinata. Calycophylls are typically (sensu Hooker) have been classically distinguished<br />
pigmented (white, cream, pink, or red) and have pal- from the <strong>Rondeletieae</strong> in having valvate aestivation,<br />
mate venation. In some species of Chimarrhis (C. but careful observation showed that only a few genbarbata<br />
and C. turbinata) bracts are present as foliose era of this group have truly valvate corollas. All the<br />
expansions in the terminal portions of the inflores- genera placed in the subtribe Portlandiinae sensu<br />
cences, not to be confused with calycophylls, because Hooker (Bikkia, Ceuthocarpus, Coutaportla, Couthey<br />
are extrafloral, not colored, and pinnately veined. tarea, Cubanola, Hintonia, Isidorea, Nernstia, Osa,<br />
Pogonopus consistently has inflorescences with a Portlandia, Schmidtottia, Siemensia, and Thogsennia)<br />
multitude of calycophylls. Because this genus has have imbricate lobes with reduplicate corolla tubes.<br />
corymbose inflorescences, the calycophyllous flow- and have been removed to the newly amended tribe<br />
ers are positioned on the older external flowers of the Catesbaeeae (Delprete, 1996d).<br />
inflorescences (the central flowers of the ultimate Valvate aestivation is commonly defined as nonbranchlets).<br />
In Pogonopus the number of calycophyl- overlapping floral parts with continuous contact along<br />
lous flowers per inflorescence is highly variable, rang- their margins. In the genera studied, valvate aestivaing<br />
from 2-3 to 20-30.<br />
tion is correlated with fleshy corollas. Typical valvate<br />
The main function of calycophylls is obviously to aestivation (without induplications or reduplications)<br />
attract potential pollinators. In Pogonopus, whose<br />
most probable pollinators are hummingbirds, the<br />
calycophylls (and flowers) are dark pink to bright red<br />
(whereas in Pinckneya, whose most probable pollinators<br />
are butterflies, the calycophylls [and flowers]<br />
are cream-white to pale-pink). In Pogonopus (as in<br />
Pinckneya) the color intensity of calycophylls varies<br />
according to the solar exposure of the inflorescences,<br />
being pink in shaded individuals and deep-pink to<br />
dark-red in sunny exposures. (In Pinckneya the indiis<br />
present in some species of <strong>Rustia</strong> (i.e., R. occidentalis,<br />
R. dressleri, and R. bilsana) and in Dioicodendron.<br />
Valvate aestivation with reduplicate corolla<br />
lobes is present in the remaining species of <strong>Rustia</strong>,<br />
and consistently in <strong>Tresanthera</strong>, Condaminea,<br />
Picardaea, and Pogonopus (and Pinckneya). Chimarrhis<br />
and Dolichodelphys have usually been reported<br />
to have valvate aestivation (and for this reason were<br />
previously included in the Condamineeae sensu<br />
Hooker), but have instead corolla lobes narrowly im-
30 Flora Neotropica<br />
bricate. Molopanthera shares with Chimarrhis and<br />
Parachimarrhis narrowly imbricate corollas (Fig.<br />
2N), the latter differing by having flower buds curved<br />
upward (and unequal stamens).<br />
Corolla shapes and sizes are also quite variable<br />
(Fig. 4). In <strong>Rustia</strong> corollas vary from minute and cam-<br />
panulate (R. costaricensis, Fig. 4B) to medium-sized<br />
and campanulate (Fig. 4A) to long and narrowly tu-<br />
bular (Fig. 4C). In <strong>Tresanthera</strong>, Condaminea, and<br />
Picardaea the corollas are medium-sized and cam-<br />
panulate (Fig. 4D-F), while in Pogonopus (Fig. 4G)<br />
(and Pinckneya, Fig. 4H) they are medium-sized and<br />
long-tubular. Chimarrhis (Fig. 4J,K), Dioicodendron<br />
(Fig. 4L,M), and Molopanthera (Fig. 4N) have mi-<br />
nutely campanulate flowers (creamy-white and fra-<br />
grant). Dolichodelphys is unique in this group in hav-<br />
ing corollas that vary greatly in size (from 3 cm long<br />
down to a few millimeters long) and shape (from tu-<br />
bular to urceolate) depending on their position in the<br />
inflorescences (see Dolichodelphys below).<br />
Fleshiness of the corolla is variable among species<br />
of <strong>Rustia</strong>, from fleshy (R. occidentalis, R. simpsonii,<br />
and R. angustifolia) to semifleshy (R. dressleri, R.<br />
alba, R. bilsana, R. rubra, R. viridiflora, R. schunke-<br />
ana, R. thibaudioides, R. formosa) or thin-delicate (R.<br />
gracilis). Corolla thickness varies also among species<br />
of Pogonopus, being either semifleshy (P. speciosus)<br />
or membranaceous (P. exsertus and P. tubulosus). In<br />
Pinckneya the corollas are semifleshy, whereas in<br />
<strong>Tresanthera</strong>, Condaminea, and Picardaea the corol-<br />
las are thick-fleshy (and all with similar pigmenta-<br />
tion). The minute corollas of Chimarrhis, Dioico-<br />
dendron, and Molopanthera are relatively thick with<br />
respect to their dimensions (a few millimeters).<br />
Dolichodelphys has thinly membranous corollas.<br />
Corolla color is variable among the species of<br />
<strong>Rustia</strong>, being white (R. costaricensis, R. dressleri<br />
[with yellow lobes], R. alba, R. simpsonii), pinkish<br />
white (R. thibaudioides, R.formosa, R. gracilis), red<br />
(R. rubra [with green lobes]), green (R. viridiflora),<br />
greenish yellow (R. schunkeana), and purple to flesh-<br />
red outside and white-perlaceous inside (R. occi-<br />
dentalis, R. angustifolia, and R. bilsana). The fleshy<br />
corollas of <strong>Tresanthera</strong>, Condaminea, and Picardaea<br />
are red outside and white-perlaceous inside. The tu-<br />
bular flowers of Pogonopus are deep-pink to red [and<br />
those of Pinckneya a creamy-white with red streaks].<br />
The minute flowers of Chimarrhis, Dioicodendron,<br />
and Molopanthera are cream-white to pale-green. The<br />
membranous corollas of Dolichodelphys are variable<br />
in size, shape, and color, ranging from whitish cream<br />
to yellowish green to pale yellow.<br />
The genera reported to have a sweet fragrance are<br />
the minutely flowered trees of Chimarrhis, Dioico-<br />
dendron, Molopanthera, and Parachimarrhis, which<br />
have flowers that are cream-white and bee-pollinated.<br />
At least two species of <strong>Rustia</strong> have sweet-fragrant<br />
flowers: R. alba and R.formosa. Pogonopus (and Pinckneya)<br />
have a sweetish scent that is barely perceptible<br />
to humans.<br />
Corollas of the genera studied have an internal ring<br />
of hairs, usually located toward the base of the corolla<br />
tube (near the orifice in minute flowers), close<br />
to filament attachment (the only exception being<br />
Pinckneya, where the internal ring of hair is absent).<br />
The function of the internal ring of hairs (usually basal<br />
in position) is believed to be protection against possible<br />
nectar-robbers that might enter through the orifice<br />
of the corollas. This strategy obviously does not<br />
protect the flowers against wasps, bumblebees, and<br />
other insects that rob the flowers by lateral incisions,<br />
below the ring of hairs.<br />
Stamens<br />
Stamens in the group studied are most commonly<br />
exserted, and the anthers open by longitudinal slits,<br />
except for the poricidal anthers of <strong>Rustia</strong> and <strong>Tresanthera</strong>.<br />
Anther presentation is exceptional in <strong>Rustia</strong>,<br />
which also includes a few pendulous-flowered species<br />
with narrowly tubular corollas: R. thibaudioides<br />
(completely included), R. viridiflora (included but<br />
with tips excluded), and R. schunkeana (partially exserted).<br />
Anther shapes are commonly broadly elliptic<br />
to narrowly elliptic in all genera, but are long and<br />
narrow in the pendulous-flowered species of <strong>Rustia</strong>.<br />
<strong>Rustia</strong> and <strong>Tresanthera</strong> are unique among the genera<br />
presented in having poricidal anthers, but they<br />
differ from each other in being apical and biporate in<br />
the former, and lateral and with a common pore in the<br />
latter. The modality of anther opening also differs in<br />
the two genera. The anthers of <strong>Rustia</strong> open by reflexing<br />
the apical deltoid flap placed in a dorsal (abaxial)<br />
position, and leaving the two cylindrical thecae open<br />
at their apical ends. In <strong>Tresanthera</strong> the two thecae are<br />
apically fused and curved toward the center; they open<br />
by reducing their curvature, causing the lateral common<br />
pore below the apex to expand in a ventral<br />
(adaxial) position. <strong>Rustia</strong> occidentalis (see details<br />
under this species) has been personally observed to<br />
be buzz-pollinated by bees, which is the first official<br />
report of such specialized pollination within the <strong>Rubiaceae</strong><br />
(most common in the Solanaceae and Melastomataceae).<br />
Other species, R. dressleri and R. bilsana,<br />
with fleshy campanulate corollas and half-exserted anthers,<br />
are also probably buzz-pollinated by bees. The<br />
tubular pendulous flowers of <strong>Rustia</strong> thibaudioides have<br />
included anthers and are pollinated by hummingbirds.
Introduction to the Neotropical Genera Studied 31<br />
A<br />
C<br />
tern<br />
FIG. 4. Flower variation<br />
in the genera referred<br />
to the Condamineeae<br />
(sensu Hooker, 1873). A-C. <strong>Rustia</strong>. A. R. occidentalis<br />
(from fresh material and Delprete 6329, TEX). B. R. costaricensis (Gomez-Laurito 11698, F). C. R. thibaudioides (from<br />
fresh material and Delprete & Fagua 6378, TEX). D. <strong>Tresanthera</strong> condamineoides var. condamineoides (Sandwith 1757,<br />
K). E. Picardaea cubensis (Wright 2661, GH). F. Condaminea corymbosa (from fresh material and Delprete & Verduga<br />
6412, TEX). G. Pogonopus speciosus var. sandwithianus (from fresh material and Delprete & Apreza 6359, TEX). H.<br />
Pinckneya bracteata (from fresh material and Delprete & Sharif 6474, TEX). I. Kerianthera preclara (Santos & Coelho<br />
759, K, NY). J, K. Chimarrhis glabriflora (from fresh material and Delprete & Verduga 6424, TEX). J. Flower bud with<br />
receptive stigma protruding above still closed corolla. K. Later stage (after J), with anthers exserted above open corolla and<br />
without style (abscissed). L, M. Dioicodendron dioicum. L. Female flower (Barklay et al. 3424, US). M. Male flower<br />
(Cuatrecasas 8665, US). N. Molopanthera paniculata var. paniculata (Williams & Assis 6861, GH), curved floral bud<br />
(zygomorphic).
32 Flora Neotropica<br />
Peculiar anther extensions were observed in <strong>Rustia</strong><br />
and Molopanthera. In R. simpsonii the base of the<br />
anthers have papillose spheroid extensions.<br />
In M. paniculata the flower buds are bent upward,<br />
and when the flowers open the two lower filaments<br />
often remain bent upward, with two pairs of adjacent<br />
anthers connected to each other by their pointed ex-<br />
tensions at both ends (forming two cup-shaped units),<br />
with the fifth filament remaining free (see Fig. 85C).<br />
The peculiar floral morphology of Molopanthera<br />
seems to be a mechanism of pollen projection, which<br />
has been reported by Schumann (1891: 10) but not<br />
personally observed. Apparently the pollen is shed,<br />
when the flower is still closed, into a spherical mass,<br />
and the single filament projects this mass onto the<br />
pollinators (most probably bees). [A detailed descrip-<br />
tion of such pollen projection mechanism in Poso-<br />
queria has been explained by Halle (1967: fig. 40)].<br />
Styles<br />
Styles in these genera are terete, glabrous, exserted<br />
beyond the corollas, and terminated by two style<br />
branches (stigmas). Most species of Chimarrhis are<br />
exceptional in having styles minutely costate (C.<br />
parviflora, C. latifolia, C. cubensis, C. jamaicensis,<br />
C. cymosa, C. microcarpa, C. speciosa, C. turbinata,<br />
and C. duckeana) or strigose to puberulent (C.<br />
turbinata, C. barbata, C. duckeana, and C. gentryana).<br />
When the flowers are presented in a horizontal<br />
or oblique position, the styles are curved upward in<br />
<strong>Rustia</strong>, Condaminea, Chimarrhis, and Pogonopus<br />
(and Pinckneya). The stigma branches are generally<br />
ovate-oblong, with stigmatic surfaces minutely (40x)<br />
papillose. In Chimarrhis, the stigmas are exserted (and<br />
receptive) beyond the still-closed corollas (protogynous).<br />
Dioicodendron is exceptional in being dioecious;<br />
in its female flowers the styles are exserted<br />
beyond the corollas, and the stigmas are arcuate with<br />
their tips still connected at maturity. None of the genera<br />
studied are heterostylous [Chimarrhis was erroneously<br />
reported as such by Urban (1899)].<br />
Ovaries<br />
In the genera studied the ovaries are inferior and<br />
bilocular, with axile placenta on which numerous<br />
ovules are horizontally (vertically in Chimarrhis and<br />
Molopanthera) inserted. The ovaries are topped with<br />
a nectariferous disk, composed mainly of parenchyma<br />
cells. In some genera (<strong>Rustia</strong>, <strong>Tresanthera</strong>, Chimar-<br />
rhis) the shape, persistence, and vestiture of the caly-<br />
ces in mature capsules proved to be useful taxonomic<br />
characters at the specific and varietal levels. Shape<br />
and vestiture of the disks in the generic and specific<br />
descriptions below are referred to as they appear in<br />
mature capsules.<br />
Fruits<br />
In the genera here studied, fruits are multiseeded,<br />
bilocular, woody capsules with a central placenta. In<br />
early stages the exocarp and mesocarp of immature<br />
fruits are green and fleshy, becoming brown and woody<br />
at maturity. Mesocarp fleshiness has been considered<br />
by many botanists (Hooker, 1873; Baillon, 1880;<br />
Schumann, 1889, 1891; Verdcourt, 1958; Bremekamp,<br />
1966; Steyermark, 1974; Robbrecht, 1988, 1993) to<br />
be a key character in establishing tribal and generic<br />
delimitations. It has been recently demonstrated<br />
(Bremer, 1992; Bremer & Eriksson, 1992; Bremer &<br />
Struwe, 1992) that fleshiness of the mesocarp in mature<br />
fruits has evolved independently several times<br />
within the <strong>Rubiaceae</strong>. Nevertheless, its total dismissal<br />
as a taxonomic character would deprive systematists<br />
of a useful source of data (Delprete, 1996d).<br />
Size and shape of capsules are rather consistent<br />
within the same genus, offering a good practical aid<br />
for immediate generic identification. The shapes and<br />
dimensions of capsules are quite variable among genera:<br />
<strong>Rustia</strong> globular, obovoid to turbinate, 4-23 x 4-<br />
9 mm; <strong>Tresanthera</strong> ellipsoid to obovoid, 15-24 x 8-<br />
12 mm; Condaminea obovoid-oblong, 14-40 x 6-11 mm;<br />
Picardaea oblong, 15-19 x 6.5-8 mm; Dolichodel-<br />
phys narrowly cylindrical, 14-23 x 3.5-5 mm;<br />
Pogonopus obovoid to ellipsoid, 5-10 x 4-9 mm (and<br />
Pinckneya bilobed-subglobose, 10-25 x 15-25 mm);<br />
Chimarrhis obovoid to turbinate, 1.5-12 x 1.5-6 mm;<br />
Dioicodendron obovoid to turbinate, 3-4.5 x 2-3.5 mm;<br />
Molopanthera bilobed-subglobose, 2-3 x 4-5 mm;<br />
and Parachimarrhis, biglobular, 3-4.5 x 1.5-2.5 mm.<br />
The mode of capsular dehiscence is a valuable<br />
character in establishing generic boundaries among<br />
the genera studied. Mature capsules of Chimarrhis,<br />
Dioicodendron, and Dolichodelphys have septicidal<br />
dehiscence, whereas <strong>Rustia</strong>, <strong>Tresanthera</strong>, Picardaea,<br />
Condaminea, and Pogonopus (and Pinckneya) have<br />
loculicidal dehiscence. Picardaea has a unique dehiscence<br />
pattern, splitting loculicidally but with the two<br />
halves remaining attached at the apex and base (no<br />
apical dehiscence), as in Macrocnemum (under which<br />
it was originally described).<br />
I recently introduced the term "secondary dehiscence"<br />
(Delprete, 1995a, 1996d), referring to the splitting<br />
that occurs in advanced stages of capsule maturity,<br />
usually after and perpendicular to primary<br />
dehiscence. Secondary dehiscence may or may not<br />
occur, depending on the genus, usually occurring only<br />
at the apices of old capsules; it is a character of some
Introduction to the Neotropical Genera Studied 33<br />
taxonomic importance at the generic level. Second- examined and mounted on SEM stubs using a light<br />
ary dehiscence has been observed in old capsules of microscope. The mounted samples were gold-coated<br />
some species of <strong>Rustia</strong>, <strong>Tresanthera</strong>, Pogonopus, with a Ladd sputter-coater, and then studied, mea-<br />
Chimarrhis, and Dioicodendron. Secondary dehis- sured, and photographed using a Phillips 515 scancence<br />
has not been found (and is probably absent) in ning electron microscope (SEM) housed at the Cell<br />
Condaminea, Picardaea, and Dolichodelphys. Research Center of the University of Texas at Austin.<br />
The dimensions of the seeds are given first by the<br />
SEED MORPHOLOGY<br />
ranges of length and second by the ranges in width.<br />
Voucher specimens are cited in the explanations of<br />
Seed morphology has been considered for a long the microphotographs.<br />
time to be of cardinal importance in <strong>Rubiaceae</strong> systematics<br />
(Hooker, 1873; Baillon, 1880; Schumann, <strong>Rustia</strong> (Figs. 5A-F, 6A-H). Seeds of irregular size<br />
1888a, 1888b, 1889, 1891; Verdcourt, 1958; and shape, flattened, 0.67-2.9 x 0.25-1.33 mm; acute<br />
Bremekamp, 1966; Steyermark, 1974; Robbrecht, at base; acute, truncate, or uncinate at distal end; testa<br />
1988, 1993). Recent investigations (Bremer, 1992; shallowly reticulate, cells elongated and longitudinally<br />
Bremer & Eriksson, 1992; Delprete, 1995a, 1996d) oriented, interspaces smooth to sparsely granular.<br />
have shown that the importance given to seed mor- Species of which seeds were available were observed<br />
phology (i.e., endosperm fleshiness, presence of to have the following dimensional ranges: R. costawings,<br />
etc.) in the classification of this family needs ricensis (Fig. 5A,B) 0.75-1 x 0.25-0.4 mm; R. occireevaluation.<br />
Exotesta sculpturing is an important dentalis (Fig. 5E) 1.06-1.26 x 0.67-0.83 mm; R. alba<br />
taxonomic trait at the generic and specific levels. The (Fig. 6F) 2.25-2.9 x 0.65-0.75 mm; R. rubra (Fig.<br />
presence of wings (testal outgrowths) in seeds was 5F) 1.16-1.5 x 0.33-0.53 mm; R. viridiflora (Fig. 6E)<br />
considered a very important taxonomic character by 0.87-1.27 x 0.33-0.83 mm; R. schunkeana (Fig. 6A)<br />
Baillon (1879), Schumann (1888a, 1888b, 1889, 1.17-1.67 x 1-1.33 mm; R. thibaudioides (Fig. 6C,D)<br />
1891), and Andersson and Persson (1991), but 1.4-1.5 x 0.75-1.25 mm; R. venezuelensis (Fig. 6G,H)<br />
Bremekamp (1966) and Verdcourt (1958) considered 0.93-1 x 1 mm; R.formosa (Fig. 5C) 1.4-1.6 x 0.48it<br />
to be of little importance. Bremer and Eriksson 0.5 mm; R. angustifolia (Fig. 6B) 1.33-1.5 x 0.4-0.73<br />
(1992) concluded that winged seeds are "not predomi- mm; R. gracilis (Fig. 5D) 0.87-1.16 x 0.43-0.67 mm.<br />
nant in any of the six groups" of their cladistic analysis,<br />
which "may indicate that winged seeds are more <strong>Tresanthera</strong> (Fig. 7A-C). Seeds generally ob-<br />
'evolutionary plastic' than the fleshy fruits." As a triangular, flattened, 1-1.5 x 0.33-0.53 mm; acute at<br />
result of a phylogenetic analyses using morphologi- base, truncate at distal end; testa reticulate, cells eloncal<br />
characters, Delprete (1995a, 1996d) concluded that gated and longitudinally oriented, interspaces smooth to<br />
the presence and general shape of seed wings is de- sparsely granular at margins. The seeds of this monopendent<br />
on the insertion of the seeds (whether later- typic genus have been measured as follows: T.<br />
ally or ventrally attached) on the placenta and their condamineoides var. condamineoides (Fig. 7C) 1.33-1.5<br />
final position at maturity (horizontal or imbricate). x 0.37-0.53 mm; T. condamineoides var. thyrsiflora (Fig.<br />
Seed wings of different genera should not be assumed 7A,B) 1-1.2 x 0.33-0.5 mm.<br />
to be homologous, because of their different origins<br />
and developments. When properly used, seed mor- Condaminea (Fig. 7D-H). Seeds generally obphology<br />
provides a good set of taxonomic characters long, irregularly 3-4-angular, 0.57-0.77 x 0.13-0.33<br />
for inferring the phylogeny of the <strong>Rubiaceae</strong>. mm; acute at base, truncate at distal end; testa shal-<br />
In most genera, the seeds are laterally attached and lowly reticulate, cells elongated and longitudinally<br />
horizontally positioned within the capsules (Fig. 5A), oriented, interspaces smooth. The seeds of C. corymexcept<br />
for Chimarrhis and Molopanthera, where the bosa (Fig. 7D-F) are 0.57-0.77 x 0.13-0.33 mm. The<br />
seeds are peltate on the placenta and umbilically at- dubiously placed C. elegans have seeds trapezoid (Fig.<br />
tached. In Chimarrhis the placenta is central and ver- 7H), 1.3-1.87 x 0.9-1.33 mm, testa densely reticutical,<br />
while in Molopanthera the placenta is globular late, cells variously oriented, and granular interspace.<br />
and supported by a short stalk in a central position. In All the previous characters place this species within<br />
most genera, seeds are minute and of irregular shape <strong>Rustia</strong>, but its floral morphology is similar to that of<br />
(but in Parachimarrhis are hemi-elliptic in outline, with Condaminea, where it is tentatively maintained.<br />
lateral linear hilum) and are adapted to wind dispersal.<br />
Seeds for SEM studies were removed from mature Picardaea (Fig. 8F). Seeds generally oblong to<br />
capsules of herbarium specimens. The seeds were first trapezoid, irregularly 3-4-angular, 0.5-0.7 x 0.2-0.4
34 Flora Neotropica<br />
i~ d.J Cri " ... . . ; lOlllslil~i . ......====================================.<br />
. . .. `~~~l~~''' .::ri :.<br />
,<br />
.El;$ifii~ c~~$~llsl3.... .-..l<br />
.<br />
...<br />
1s 74 I | _<br />
o.<br />
"-'~ ,.^ ~... . ... . . :~~-w~ _~<br />
:. ':" '<br />
5 : __ BF 0.3r / 1F m : i 29 F.. _~~~~~~~~~~~~~~~~~~<br />
?::-:?:: ..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
_e~~~~~~:i- ? *_<br />
mm;~_ B-F 0, mm.I8_^. s<br />
FIG. 5. SEM photomicrographs of seeds of <strong>Rustia</strong>. A, B. R. costaricensis (Croat 22191, MO). A. Capsule in longitudi-<br />
:.q<br />
_f~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i#<br />
I:<br />
A _<br />
_1_.?"iF s~; re??~~<br />
_<br />
~~8r* :e>.<br />
a t a - E~, i<br />
i,. 4l.~<br />
* t .,.*? ... 's<br />
.<br />
FIG. 5.SM htomcrgrph ofses ofRsi.A .R otriess(ra 29,M).A asl nlniui<br />
na etin hoigsed aerlyatahd B ed.C .oroa(Iwne a. 8881, MO),U sed.R.gaiis(lzo<br />
1493 M),sed.E.R.ocidntli (elree 32, EX, ee. . . uba 8021,.;? Ster US) Scal~Y~~ se.,; e br: 3~ A,<br />
mm; B-F,?" ~ 0. mm.""" ~ I<br />
::~~l..?<br />
mm; acute at base, truncate at distal end; testa irregu-<br />
larly reticulate, cells mostly with irregular shape,<br />
without particular orientation, interspaces densely<br />
granular. Even though this genus is usually placed<br />
close to Condaminea, its seed morphology closely<br />
resembles that of Pogonopus.<br />
Pogonopus (Fig. 8A-E). Seeds generally oblong<br />
to trapezoid, irregularly 3-5-angular, 0.38-0.75 x<br />
0.23-0.49 mm; acute at base, truncate at distal end;<br />
testa irregularly reticulate, cells mostly with irregu-<br />
lar shape and undulate margins, generally longitudi-<br />
nally oriented, interspaces densely granular. The three
Introduction to the Neotropical Genera Studied<br />
-i<br />
35
36 Flora Neotropica<br />
_16;<br />
%<br />
___~~~~~r<br />
.6. .:A. ,::.'.,::' .<br />
~.,~.<br />
~~ ~'~..~~~~~~~~~W.<br />
.' ...?-.~'..<br />
'>B~..<br />
'<br />
~il<br />
t; .~-~ ~I'. . ' ~ ... '...........<br />
GiE9~~~~4 ~.~1 ~ ~ %1 A<br />
~5?FsI~i7:, ?t:,,,~...~. ~<br />
-LAW<br />
~'"~'-'"~:::'~ ~:~:"'~<br />
~~~~~S~~~~~~~C ~ ~ ~ ~ ~<br />
~ii,<br />
' ~<br />
~?. ~.~~,~.-~.. . .,,.?',fi~ . .<br />
aIl&j ?<br />
....~_,~~"?:I;- ???~:~<br />
? !~ ......<br />
!~ . .~.~ . .<br />
~~~~~uh ~~~~~~~~~~~4.<br />
/<br />
. .. ~~ ....<br />
FIG. 7. SEM photomicrographs of seeds of <strong>Tresanthera</strong> and Condaminea. A, B. <strong>Tresanthera</strong> condamineoides var.<br />
thyrsiflora (Bernardi 5829, K). A. Seed. B. Close-up of seed coat. C. <strong>Tresanthera</strong> condamineoides var. condamineoides<br />
(Broadway 3076, GH), seed. D-G. Condaminea corymbosa. D. Seed (Delprete et al. 6049, TEX). E. Seed (Schunke 10512,<br />
MO). F. Close-up of seed coat (Schunke 10512, MO). G. Seed (Fosberg 28672, US). H. Condaminea elegans (Foster &<br />
Vivar 13333, MO), seed. Scale bars: A, C-E, G, H, 0.3 mm; B, F, 0.01 mm.<br />
~!<br />
"~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
FIG. 7 SEM potomirograps of seds o Tresatheraand Codamine. A, . Tresntheracondaineoids var<br />
rsfor ed .Coeufse d C.i ot Tresanhrodaiedsvr.cnanods<br />
(Broaday 307, GH),seed. -G. Codamine corymosa. D Seed Delprte eta.64,TX.E.Se Shne152,~<br />
MO). F Closeup fsedcat Shne102,M)G.Se(oser287,U)H.Cnarnnaelgn<br />
F ostr<br />
Via 333 - Osed E cl br:A , G ,0. m ,F,00 m
Introduction to the Neotropical Genera Studied 37<br />
...'{<br />
. a<br />
PIP -', .<br />
MM"'.:.ls<br />
FIG. 8. SEM photomicrographs of seeds of Pogonopus, Picardaea, and Pinckneya. A. Pogonopus speciosus var. sandwithianus<br />
(Delprete & Apreza 6359, TEX), cross section of capsule. B. Pogonopus exsertus (Gentry 6720, NY), seed. C. Pogonopus<br />
speciosus var. speciosus (Pittier 11672, US), seed. D. Pogonopus speciosus var. sandwithianus (Delprete & Apreza 6359, TEX),<br />
seed. E. Pogonopus tubulosus (Knrukoff 10167, A), seed. F. Picardaea cubensis (Liogier & Liogier 26174, NY), seed. G, H.<br />
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.
38 Flora Neotropica<br />
species and two varieties have been measured as fol-<br />
low: P. exsertus (Fig. 8B) 0.38-0.58 x 0.23-0.35 mm;<br />
P. speciosus var. speciosus (Fig. 8C) 0.6-0.67 x 0.31-<br />
0.49 mm; P. speciosus var. sandwithianus (Fig. 8D)<br />
0.44-0.64 x 0.35-0.42; P. tubulosus (Fig. 8E) 0.51-<br />
0.75 x 0.29-0.47. The seeds of this genus are typical<br />
in their irregular cells with undulate margins and<br />
densely granular interspace.<br />
Chimarrhis (Figs. 9A-H, 10A-F). Seeds with very<br />
irregular outline, with deeply fringed concentric wing<br />
and hilum in central position, peltately attached to a<br />
central placenta and vertical within the capsule, 0.49-<br />
2.38 x 0.33-1.5 mm; testa deeply reticulate, cells of<br />
irregular shape in the central portion and tendentiously<br />
elongated in the outer portion (forming a narrow concentric<br />
wing) and radially oriented; interspaces<br />
smooth or with foveolate thickening.<br />
Chimarrhis is exceptional within the <strong>Rondeletieae</strong><br />
s.l. because of its inflorescence position, flower, fruit,<br />
and seed morphology. Chimarrhis duckeana (Figs.<br />
10B, 10F), C. turbinata (Fig. 10D), C. barbata, and<br />
exceptionally C. gentryana (Fig. 10E) have seed wings<br />
that tend to elongate in opposite directions, vaguely<br />
resembling the bipolar wings of the genera of the<br />
Cinchoneae (sensu Andersson & Persson, 1991).<br />
The thickenings present within the interspaces of<br />
the primary reticulation are not consistent within the<br />
species, being variable even among seeds of the same<br />
capsule, and cannot be used as a reliable taxonomic<br />
character. Nevertheless there is a tendency in C. latifolia<br />
(Fig. 9B), C. barbata, and C. turbinata (Fig. 10D)<br />
to develop cell thickenings largely parallel to each other<br />
wings at each corer of the body, hilum in central position,<br />
0.5-0.95 x 0.4-0.55 mm; testa shallowly reticulate,<br />
cells of irregular shape and without any particular<br />
orientation; interspaces finely and densely reticulate.<br />
Dioicodendron is considered to be closely related to<br />
Chimarrhis, but the seeds of the former (Figs. 9A-H,<br />
10A-F) differ from those of the latter in being ovoid in<br />
general shape, with extremely narrow wings (when<br />
present), and finely granulate interspaces.<br />
Molopanthera (Fig. 1 1B). Seeds with very irregu-<br />
lar outlines, with deeply fringed concentric wing and<br />
hilum in central position (Fig. 1 B), peltately attached<br />
to globular placental extensions (one each locule), 1 x<br />
2 mm; testa shallowly reticulate, cells elongated with<br />
radial orientation; interspaces with foveolate-reticu-<br />
late thickenings.<br />
The seeds of Molopanthera are extremely similar<br />
to those of Chimarrhis (Fig. 9A-H, 10A-F) in being<br />
attached by a central hilum and peltate to the placenta,<br />
with an irregularly fringed concentric wing, and hav-<br />
ing cells with foveolate thickenings. The two genera,<br />
however, differ in their flower, fruit, and inflorescence<br />
morphology.<br />
Dolichodelphys (Fig. 12A,B). Seeds ovoid, rectangular<br />
to trapezoid, 3-5-angular, unwinged and with<br />
hilum in lateral position (Fig. 12A), laterally attached<br />
to the central placenta, 0.1-0.3 x 0.01-0.15 x 0.01-<br />
0.1 mm; testa reticulate, cells 4-5 sub-isolateral without<br />
particular orientation; thickening ovate in cross<br />
section (Fig. 12B); interspaces smooth.<br />
and perpendicular to the<br />
This dubious<br />
longer margins ("scalariform"<br />
monotypic genus has been placed in<br />
thickenings). In C. brevipes (Fig. 10A), C. duckeana<br />
the Gardenieae (Robbrecht, 1988, 1993) or confused<br />
(Fig. 10B,F), C. turbinata (Fig. 10D), and C.<br />
with <strong>Rustia</strong><br />
gentry-<br />
(Schumann & Krause, 1908c; Standley,<br />
ana (Fig. 10E), the marginal thickenings have 1930b;<br />
spo-<br />
Steyermark, 1974). Based on its seed, flower,<br />
radic minute echinate<br />
and fruit<br />
processes.<br />
morphology, Dolichodelphys obviously does<br />
The dimensional ranges of the seeds of the<br />
not<br />
spebelong<br />
to the Gardenieae, and is included in the<br />
cies directly observed are as follows: C.<br />
<strong>Rondeletieae</strong> s.l.<br />
parviflora<br />
(Delprete, 1996d).<br />
(Fig. 9A) 0.5-0.88 x 0.38-0.5 mm; C. latifolia (Fig. 9B)<br />
0.49-0.78 x 0.33-0.58 mm; C. cubensis (Fig. 9C) 0.58- Parachimarrhis (Fig. 12G). Seeds hemi-elliptic in<br />
0.87 x 0.48-0.77 mm; C. cymosa (Fig. 9D) 0.56-0.78 x outline, dorsiventrally flattened, laterally winged, and<br />
0.56-0.67 mm; C. glabriflora (Fig. 9E,F) 0.48-0.68 x lateral hilum (Fig. 12G), laterally attached to a cen-<br />
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<br />
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<br />
(Fig. 9H) 0.7-0.95 x 0.5-0.75 mm; C. turbi- eral without particular orientation (Fig. 10G),<br />
nata (Fig. O1D) 0.75-1.95 x 0.75-1.25 mm; C. ducke- interspaces smooth.<br />
ana 1-2.83 x 0.83-1.5 mm (Fig. 10B,F); C. gentryana This monotypic genus was treated as closely related<br />
(Fig. 10E) 0.95-1.13 x 0.58-0.88 mm; C. brevipes to Chimarrhis by Ducke (1922, hence the name) or com-<br />
(Fig. O1A) 0.85-1.05 x 0.63-1 mm.<br />
pared to both Chimarrhis and Elaeagia, but included<br />
in the Condamineeae because of its valvate aestivation<br />
Dioicodendron (Fig. 11A). Seed bodies ovoid-sub- (Steyermark, 1965). The seed morphology of Paraorbicular<br />
and 3-5-angular, sometimes with narrow chimarrhis is similar to that of Simira (Fig. 12E,F) in
Introduction to the Neotropical Genera Studied<br />
____I~ ~ ti~<br />
, .<br />
-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
A lir I 7 5 -<br />
-v~it ;_;.i-\ _<br />
i- >; w f s o w<br />
t fF h # ft~~~~~~~~"i<br />
_t ig,. , w t _<br />
o. S~h*8t<br />
~~~~~?- 't- 's e * . >=<br />
.I<br />
O _ w<br />
F<br />
_ }j.~~~~~~~~~~<br />
:' .i:- f ,''k} !t<br />
FIG. 9. SEM photomicrographs of seeds of Chimarrhis. A. C. parviplora (Jimenez 3418, F), seed. B. C. tatifolia (Allen<br />
5762, GH), seed. C. C. cubensis (Wright 1622, BR), seed. D. C. cymosa (Wilbur 8137, TEX), seed. E, F. C. glabriplora<br />
(Schunke 10653, F). E. Seed. F. Close-up of seed coat. G. C. hookeri (Ancuash 504, MO), seed. H. C. speciosa (Steyermark<br />
95211, NY), seed. Scale bars: A-E, G. H. 0.3 mm; F. 0.02 mm.<br />
Pt_ }<br />
F<br />
39
40 Flora Neotropica<br />
? .<br />
, I[~<br />
'-.~ ~.-i- ~~ .i<br />
.<br />
.~ ,~.., ~'.<br />
,,.~' t~ l,,1 "1'"."',<br />
5.~.~ ~:,~.. ...~.:a . ,a. u<br />
-._<br />
-.- ~ ~ ~ ~ ~ ~ ~ ..<br />
,..i~..:..-,:.,?,<br />
ii e. --~<br />
~..<br />
(Vquez Arevalo 8969, TEX). C. duckeana (Nascimento et al INPAWWF .302, NY). Scale bars: A-F, 0.3 .<br />
FIG. 10. SEM photomicrographs of seeds of Chimarrhis. A. C. brevipes (Maguire d& Politi 27971, US). B. C. duckeana<br />
(Fr6es 22596, NY). C. C. turbinata (Museum personnel 9 713, P). D. C. turbinata (Silva & Rosario 3 732, NY). E. C. gentryana<br />
S<br />
:. ~~:.. ~,..-. .~.<br />
;.*L.i ___I .~,-~.....' ~?-~I ~,~<br />
':'"''"i' 'c<br />
FIG. 10. SEM p ~ hotmirogr ofseso ph Ciari. A .beie Mgie&Plt 77,U) ..dce
Introduction to the Neotropical Genera Studied<br />
~ag ti '""~: ~t<br />
..?-.. ... . -. --- --<br />
dendron - S colombianum Y ~ (Sanchez a et?al. 6,M, _ _ s ...S ,? . E,.<br />
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. .<br />
..<br />
~ ' :"~"Y''"::~'"' ......~~~~~~f<br />
totype), seed. C, D. Keriantherapreclara (Bisby et:a! P18128, NY). C. Seed. D. Close-up of seed coat. E, F. Cuatrecasasio-<br />
A.Docdendron doomioicum ( Wrancke 1383, UC). , seed. B. Molos-panthsera pncult a t. ucheli b (Warming3 E s. n.n , F, lee-mm<br />
'<br />
. ' 'f...<br />
?' ,<br />
FIG~ %~, ~~~~,, SMpooirgah ?<br />
iioedo,* .....~her, fseso<br />
:oooreo<br />
n uteasidro..<br />
A. Dioicoendron diicum ( Wudock 1383 UC), see. B, Moloonthero l:. pnicltlotovar. bttrhellii ( orming s~., F,<br />
its lateral orbicular wing and extensive lateral hilum,<br />
and I think the two genera are closely related.<br />
[Pinckneya (Fig. 8G,H). Seeds with a broad thin lunate-orbicular<br />
wing (Fig. 8G), dorsiventrally flattened,<br />
body of seed 2-3 mm long, the wing 1.5-6.4 x 2.5-<br />
10.3 mm; orbicular at distal end; testa irregularly reticulate,<br />
cells mostly elongate without particular orientation,<br />
interspaces secondarily reticulate (Fig. 8H)<br />
with thickenings parallel to each other and perpendicu-<br />
41
42 Flora Neotropica<br />
lar to the two longer margins. This genus is closely<br />
related to Pogonopus (Fig. 8A-E), but the seed mor-<br />
phology of each of these two genera is very distinct.]<br />
[Kerianthera<br />
(Fig. 11C,D). Seeds with very irregu-<br />
lar outline (Fig. 11C), with deeply fringed bilateral<br />
orbicular wing, dorsiventrally flattened, 3.8-5.8 x<br />
1.3-3.5 mm; testa shallowly irregularly reticulate,<br />
cells slightly elongate without particular orientation,<br />
interspaces secondarily irregularly reticulate, with<br />
subcircular secondary interspaces (Fig. 11D).]<br />
POLLEN MORPHOLOGY<br />
Aiello (1979) published a general overview of the<br />
pollen for most genera of the Condamineeae (sensu<br />
Hooker), and several genera assigned to the Cinchoneae<br />
and <strong>Rondeletieae</strong>. Aiello divided the pollen<br />
into three main sculpture types: type I, with echinate<br />
exine; type II, with foveolate exine; and type III, with<br />
reticulate exine. The genera of the Condamineeae<br />
(sensu Hooker) with echinate exine (subtr. Portlandiinae)<br />
have been transferred to the Catesbaeeae (sensu<br />
Delprete, 1996d). Pollen of types II and III display<br />
various kinds of intergradation, as the exine of some<br />
taxa ranges from densely reticulate to foveolate, and<br />
the distinction between the two types does not hold<br />
up. For this reason, no particular pollen sculpturing<br />
categories have been proposed (or maintained) in the<br />
present study, and the pollen is described below for<br />
each genus.<br />
The pollen grains of the genera studied were all<br />
[Cuatrecasasiodendron (Fig. 11E,F). Seeds tetragonal<br />
in outline, dorsiventrally flattened, central body<br />
circular (0.1-0.2 mm diam.), with subentire concentric<br />
square wing and hilum in lateral position (Fig.<br />
1 1E), laterally attached to the central placenta, 0.25-<br />
0.45 x 0.23-0.45 mm; testa shallowly reticulate, cells<br />
elongated with peltate orientation departing from the tricolporate, and with reticulate-foveolate exine. None<br />
hilum; interspaces with foveolate thickenings (Fig. of the genera observed has dimorphic pollen, because<br />
1 1F). This genus has been placed in either Condami- none of them are heterostylous.<br />
neeae (sensu Hooker) or <strong>Rondeletieae</strong>. In my opin- As reported in previous studies (Muller, 1979;<br />
ion this genus is related to Rondeletia (s.l.), and is Blackmore & Barnes, 1986; Pacini, 1986, 1990), the<br />
therefore not included in the present study.] pollen grains of flowering plants are subjected to<br />
changes in size and shape due to mechanical properties<br />
[Kajewskiella (Fig. 12C,D). Seeds ovoid commonly called harmomegathy (Wodehouse,<br />
(sometimes<br />
angular), unwinged and with hilum in lateral<br />
1935). The pollen grains were usually prolate in shape<br />
at release and became<br />
position (Fig. 12C), laterally attached to the central<br />
spheroid upon contact with<br />
placenta, 0.35-0.6 x<br />
moisture<br />
0.25-0.35 mm<br />
(or a stigmatic surface). The colpi of the<br />
(K. trichantha);<br />
testa shallowly reticulate, cells multilateral without pollen grains were invaginated at the release stage (as<br />
normally released from mature anthers), and turned<br />
particular orientation; interspaces densely granularinside<br />
out<br />
foveolate and interconnected in parallel rows upon contact with moisture and during ger-<br />
(Fig. 12D).<br />
mination.<br />
This dubious genus was included in the<br />
Also, the exine reticulation at the polar and<br />
present<br />
study because it was previously compared to Xantho- equatorial regions was more expanded when the pollen<br />
was<br />
phytum (Hedyotideae), Dolicholobium spheroid than when it was prolate. For the<br />
(Cinchoneae),<br />
above reasons the<br />
or Bikkia (Condamineeae s.l.) by Merrill and equatorial measurements in previ-<br />
Perry<br />
ous literature are<br />
(1947), included in the Condamineeae<br />
variable, depending on the stages of<br />
(sensu Hooker)<br />
the<br />
by Jansen (1979), and tentatively maintained pollen grains (and their laboratory preparation, i.e.,<br />
by<br />
Robbrecht (1988, 1993). Because of its seed acetolysis) when measured. Because the process of<br />
morphology<br />
and presence of raphides, Tange (1995) trans- acetolysis also causes the pollen grains to expand<br />
ferred Kajewskiella to the Hedyotideae, a decision equatorially and assume a spheroid shape, the equatorial<br />
measurements were made at full<br />
with which I expansion, in<br />
concur.]<br />
the spheroid stage.<br />
Pollen-connecting threads were observed in sev-<br />
[Simira (Fig. 12E,F). Seeds hemi-elliptic in out- eral species of <strong>Rustia</strong> and <strong>Tresanthera</strong>. The nature of<br />
line, dorsiventrally flattened, laterally winged and the threads was not investigated, and their origin is<br />
with hilum in lateral position (Fig. 12E,F), laterally uncertain; nevertheless, because they were still present<br />
attached to a central placenta and in two ranks each after acetolysis, it is suspected that they are filiform<br />
locule; testa finely shallowly reticulate, cells multi- sporopollenin (as reported in Ericaceae and Onalateral<br />
without particular orientation; interspaces graceae; see Hesse, 1984, 1986; Skvarla et al., 1978;<br />
smooth (Fig. 12F). Seeds of Simira are included in the Waha, 1984). Pollen-connecting threads have usually<br />
present study because of their similarity to those of Para- been correlated (Hesse, 1986) with zoophilous pollichimarrhis,<br />
from which they differ by being larger.] nation. Several species of <strong>Rustia</strong> tend to be dispersed
Introduction to the Neotropical Genera Studied<br />
!-_ wM<br />
B<br />
A ~A<br />
dp ci.or.o ( .. .Cardenas<br />
.<br />
A?.: Seeds. B. Ce of seed cot C, D K<br />
D, 0 .01~- ? '~<br />
.~<br />
imE 1 ; 2.. ,<br />
?IG 1,~. ,aesila SMpooirgah ~,:.: Paa.~.rri. 'L..~.~"o<br />
fseso~lcoep,<br />
.fia<br />
A,<br />
'~.. ~lrcae '.~ B-~:;T-u ?ee c?'i. (aie ~:,: '~!siel &Crea IO, CL. .ed.<br />
o<br />
D,<br />
rihn<br />
(aes 16 A.i :~' ~'~:'~ i' D. seu ofse ~:',FSiialnioi SeynaR&Epnz 169,F.E<br />
F.Clseupofsed oa. . M),sed.Sc"'bas:A'"'!'!.<br />
archmarhs nn;B,0'5 reilba(Rda ia. 28<br />
m<br />
(Kajewshz 1667, A). C. Seed. D. Close-up of seed coat. E, F. Simira longifolia (Steyermark & Espinoza 106898, F). E. Seed.<br />
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;<br />
D, 0.01 mm; E, 10 mm; G, 2 mm.<br />
43
44 Flora Neotropica<br />
in clusters due to connecting threads, some species of cies the exine is foveolate (non-columellate, and os<br />
this genus being pollinated by bees (buzz-pollination without annular thickening). The reticulate exine in<br />
and lap-pollination) and others by hummingbirds (see the above three species may be related to their spe-<br />
Pollination and Other Plant-Animal Interactions). cialized pollination syndrome, buzz-pollination by bees<br />
In all genera pollen is released as single grains, ex- (pers. obs.). <strong>Rustia</strong> schunkeana (Fig. 13E), R. thibaudicept<br />
for a single collection of one species of <strong>Rustia</strong> oides (Fig. 13F), and R. simpsonii (Fig. 14G,H) have<br />
(R. costaricensis, Gomez-Laurito 11698), which con- minute echinate processes. The presence of minutely<br />
sistently releases its pollen in tetrads (Fig. 13B). The echinate exine in these species maybe related to their<br />
pollen of this collection was collected from herbarium pollination syndrome, probably hummingbird pollispecimens<br />
and I believe that it is teratological, and nation (pers. obs.).<br />
should be considered so until its viability has been tested. Two photographs of pollen grains of R. rubra (Fig.<br />
13G,H) were taken to show the different shapes of the<br />
Materials and Methods<br />
pollen grains when freshly released from the anthers<br />
Mature anthers were removed from herbarium (Fig. 13H) and expanded when in contact with moisspecimens,<br />
and acetolyzed according to a modified ture (Fig. 13G).<br />
version of Adams and Morton, 1972. Mature anthers In <strong>Rustia</strong> the pollen is released as single grains,<br />
were boiled in distilled water (in a water bath) for 10 except for a single collection (Gomez-Laurito 11698)<br />
minutes, and macerated through a fine filter. The fil- of R. costaricensis for which the pollen grains were<br />
tered water was then centrifuged and decanted. Pol- constantly released in tetrads (Fig. 13B); in other collen<br />
and debris were treated with glacial acetic acid for lections of the same species the pollen is released as<br />
five minutes, centrifuged, and decanted. Pollen was single grains (Fig. 13A). The species directly observed<br />
then acetolyzed with a freshly prepared mixture of are listed below, with the measurements of the dimen-<br />
90% acetic anhydride and 10% sulfuric acid (adding<br />
sional ranges: R. costaricensis [Skutch 4777, US] (Fig.<br />
acid to anhydride), and boiled in a water bath for 10 13A) 18-19 x 20-21 gpm; R. costaricensis tetrads<br />
minutes. The acetolyzed pollen was then centrifuged [Gomez-Laurito 11698, F] (Fig, 13B) 34-38 x 41-43 pm<br />
and decanted in distilled water three times. The pol- (each grain 19-22 x 20-21 lm); R. occidentalis (Fig.<br />
len suspended in distilled water was then evenly dis- 13C) 19-20 x 20-21 mr; R. dressleri 19-20 x 20-21 jlm;<br />
persed in petri dishes, and left to evaporate on a warm R. alba (Fig. 13D) 19-21 x 20-21 gim; R. schunkeana<br />
surface for three days. The petri dishes were left dry- (Fig. 13E) 19-21 x 20-22 gim; R. thibaudioides (Fig.<br />
ing for this period in order to omit the laborious pro- 13F) 19-21 x 20-22 gm; R. viridiflora (Fig. 14A,B)<br />
cess of critical-point drying of the specimens. Pollen 19-21 x 20-22 grm; R. gracilis (Fig. 14C,D) 19-21 x<br />
grains were transferred by pressing the SEM stubs, 20-22 gm; R. angustifolia (Fig. 14F) 19-21 x 20-22 gLm;<br />
with a small portion of double-stick tape, into the petri<br />
R. formosa (Fig. 14E) 19-21 x 20-22 tm; R. rubra<br />
dishes. The mounted samples were then gold-coated (Fig. 13G-H) 19-20 x 20-25 gm; R. simpsonii (Fig.<br />
in a Ladd sputter-coater, and studied, measured, and 14G,H) 25-26 x 25-26 pim.<br />
photographed using a Phillips 515 SEM at the Cell<br />
Research Center of the University of Texas at Austin. <strong>Tresanthera</strong> (Fig. 15A,B). Pollen grains tricolpor-<br />
In the descriptions and dimensions reported below ate, 23-27 x 24-28 im; colpi acute at ends, membrane<br />
(which were obtained by studying at least 50 pollen granular, pores with regular margin with annular<br />
grains of each species), the first range refers to equa- thickening; exine foveolate. In the variety condaminetorial<br />
diameters, and the second range to polar lengths.<br />
oides (Fig. 15B) the exine is regularly foveolate, while<br />
in the variety thyrsiflora (Fig. 13A) the exine is fo-<br />
<strong>Rustia</strong>. Pollen grains tricolporate, 18-26 x 20-26 veolate with irregular pattern and with holes variably<br />
gm; colpi acute at ends (except in R. simpsonii, Fig. small to minute (Fig. 15A,B). The varieties of this<br />
14G,H), membrane smooth to minutely granular, monotypic genus have been measured as follows: T.<br />
pores with irregular margin with (in reticulate exine) condamineoides var. condamineoides (Fig. 15B) 24or<br />
without (in foveolate exine) annular thickening; 27 x 26-28 im; T. condamineoides var. thyrsiflora<br />
exine reticulate to foveolate; in some species with (Fig. 15A) 23-25 x 24-26 jum.<br />
minute echinate processes (in foveolate exine). In R.<br />
costaricensis (Fig. 13A,B), R. occidentalis (Fig. 13C), Condaminea (Fig. 15C-F). Pollen grains<br />
and R. dressleri the exine is reticulate (and columel- tricolporate, 24-26 x 26-28 jlm; colpi acute at ends,<br />
late) both in the apocolpium and mesocolpium (os membrane granular, pores with regular margin with<br />
with annular thickening), while in the rest of the spe- small annular thickening; exine reticulate (and col-
Introduction to the Neotropical Genera Studied 45<br />
''~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~,.<br />
:'x , ~,,',, r.?e ...~.<br />
. . .~.<br />
~~~~~~~,.....~7,-<br />
?<br />
,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?cF<br />
?......<br />
.~~~~~~~~~~~45<br />
~..?i"~ ~" "~.:, ~ ~ ~ ~ ~ ~ ~ ~ r*'.'<br />
~,:.~,~~~~~~~~~~~~~~~~~~~~~~~~<br />
',:'r?.<br />
r~~~~~~~~~~~<br />
,~~~~~~~~~~~~~~~~~~~~~~~~<br />
.x,.%~ ;";~.~~<br />
'?~:'~~~.i i"'..... .rr:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~r<br />
?c.l ~~s<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1%?<br />
',~;:~"'<br />
, ,r ' {{..,<br />
panded grain. Scale bars: IO pm. ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ bh ' 4in<br />
panded grain. Scale bars' 10 ~m.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
46<br />
..~r<br />
?'?i *. :. !::. ..<br />
~" I . . 1 F - __N<br />
..~~~~~~~~. ^ 1,.<br />
_ _ _ __<br />
Flora Neotropica<br />
.?.. '~ . . . . . ~,; i; .: . ' " '<br />
?~~~~~~~~~~~~~~~~<br />
__' __<br />
?i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.<br />
_. .-ia?Yl--r~ _: l~~ " *<br />
_ sO ;.&r~*a~~~~r~:f?<br />
_ sw _ % ; X I~~~~~~~~~~~~~, __ _ _<br />
f s._ .o'a' ~~~.<br />
_ as 1- ll ''<br />
V W.<br />
s s N ,, | j _ ' t .~~~~~~~~~~~1,<br />
_?-*<br />
~;i ; ;?r .: l ~~?~i _ _<br />
.. .<br />
*s*''.+<br />
'l~~~~~~~~~~~~~~~~ii<br />
_i~~~ E ~ ~ ~~~~ 9 ~~~~~~~~~<br />
r,. *, I 4'<br />
, . . - -.s i.3 -- v (. - _<br />
1?i ?<br />
FI. 4.SE potmirorahsofpole gais f usia , . . irdiloa Luo 67, B) A Euaoralviw<br />
B. Poa iw ,D .gaii Ocin 65 S.C oa ve.D ls-p .R oms Hrne ta.65,U)<br />
eqaoilvew .R nusioi Gaio 43,K, oa iw .~ H.R impoi (Gaiu66,P).Eutra<br />
view. H. Polar view. Scale bars: AX, E-H, 10 1lm; D, 2 pm<br />
FIG. 14. SEM photomicrographs of pollen grains of <strong>Rustia</strong>. A, B. R. viridiflora (Lugo 3674, GB). A. Equatorial view.<br />
B. Polar view. C, D. R. gracilis (O??hioni 655, US). C. Polar view. !. Close-up. E. R. formosa (Heringer et al. 6157, US),<br />
equatorial view. F. R. angustifolia (Glaziou 14937, K), polar view. G, H. R. simpsonii (Glaziou 6564, P). G. Equatorial<br />
view. Hi. Polar view. Scale bars: A-C, E-H, IO gm; D, 2 gm.
Introduction to the Neotropical Genera Studied 47<br />
~ ' . ?.41,<br />
%;!w" '<br />
-. ?<br />
I<br />
"i'r" ~~~~~;E~~~T~~i~~C~~~~1?;~~~'~~r?S<br />
I ~i<br />
V1.~~~~~~~~~~~~~~~1<br />
P<br />
.<br />
i_-Fw --<br />
_. . .- v<br />
Mi~~~~~~<br />
. '<br />
~~" - 'j'<br />
FIG. 15. SEM photomicrographs of' pollen grains of' Tresnthera, Condaminea, and/Picardaea. A. T7resanthera<br />
NY). G. Polar view. H. Equatorial view. Scale bars: l10 !pm.<br />
4V~~~~~~~~~~~~~~~~~~~F~L(~~<br />
?* IC?- ~ It f*PM<br />
FIG 1. SM hotmirogaps o plle gain o Tesathra,Codamne, ad icadaa. . resnter<br />
conamieodesvar tyrsfloa Berard 529,K),poar ie. B T onamieoiesvar codaineide (roawa<br />
306 G) eutoil iw.CF Cnamna oymoa C , eprt &Wbse 60,TE;E,F Bee 51,N)<br />
C,E oa iw . qaoilve.F.Eutra iwofuepne ri. ,H ade ubni Sae 80<br />
NY) G.Polr vew.H. quaoral iew Scle ars IOpm<br />
,<br />
_
48 Flora Neotropica<br />
umellate) in the mesocolpi and foveolate in the<br />
apocolpi. Many formal names have been proposed for<br />
C. corymbosa or its segregates, because of its vestiture<br />
and leaf morphology variation; nevertheless its flower<br />
and pollen (and seed) morphology is constant through-<br />
out its range. Condaminea measurements are as fol-<br />
low: C. corymbosa (Fig. 15C,D) 24-26 x 26-28 gum.<br />
The dubious C. elegans, tentatively placed in this<br />
genus, did not have pollen material available to as-<br />
certain its generic placement.<br />
Picardaea (Fig. 15G,H). Pollen grains tricolporate,<br />
23-25 x 23-24 um; colpi rounded at ends, membrane<br />
granular, pores with irregular margin with marked<br />
annular thickening (Fig. 15H); exine reticulate (and<br />
columellate) in the mesocolpi and foveolate in the<br />
apocolpi. The two previously recognized species are<br />
here reduced to synonymy under one species (P.<br />
cubensis).<br />
Pogonopus (Fig. 16A-D). Pollen grains tricolporate,<br />
16-18 x 17-18 jm; colpi obtusely rounded at<br />
ends, membrane granular, pores with regular margin<br />
with marked annular thickening; exine reticulate (and<br />
columellate) in the mesocolpi and foveolate in the<br />
apocolpi. The three species and two varieties have<br />
practically identical pollen grains, and their measurements<br />
are as follows: P. exsertus (Fig. 16A,B) 17-18 x<br />
17-18 jm; P. speciosus var. sandwithianus (Fig. 16C)<br />
16-18 x 17-18 um; P. tubulosus (Fig. 16D) 16-17 x<br />
17-18 uim.<br />
[Pinckneya (Fig. 16E,F). Pollen grains tricolporate,<br />
24-25 x 26-27 gm; colpi acutely rounded at ends,<br />
membrane granular, pores with irregular margin with<br />
thin annular thickening; exine reticulate (and columellate)<br />
in the mesocolpi and foveolate in the apocolpi.<br />
This genus is closely related to Pogonopus, which is<br />
confirmed by their similarity in pollen morphology].<br />
[Kerianthera<br />
(Fig. 16G,H). Pollen grains pentacol-<br />
porate (Fig. 16G), 28-30 x 25-26 gm; colpi small and<br />
widely acute at ends, membrane densely granular,<br />
pores very small with irregular margin without annu-<br />
lar thickening; exine minutely foveolate throughout].<br />
Chimarrhis (Figs. 17A-H, 18A-E). Pollen grains<br />
tricolporate, 14-17 x 15-18 im; colpi irregularly<br />
acute at ends, membrane granular, pores with regular<br />
margin with annular thickening; exine thickly foveo-<br />
late-reticulate (and columellate) throughout. All the<br />
species have similar pollen grains, and differences<br />
among species are minimal. Two photographs of<br />
grains of C. parviflora were taken to show the different<br />
shapes when released from anthers (Fig. 17B) and<br />
as expanded during germination (Fig. 17A). Pollen<br />
grain measurements were the following: C. parviflora<br />
(Fig. 17A,B) 14-17 x 17-18 jm; C. latifolia (Fig.<br />
17C,D) 16-17 x 17-18 Lgm; C. cubensis (Fig. 17E)<br />
14-16 x 16-17 gum; C. jamaicensis (Fig. 17F) 14-15 x<br />
15-16 gim; C. cymosa (Fig. 17G-H) 15-16 x 16-17 gm;<br />
C. speciosa (Fig. 18A,B) 14-15 x 15-16 gm; C microcarpa<br />
14-15 x 15-16 gm; C. hookeri (Fig. 18C) 16-<br />
17 x 17-18 gm; C. turbinata (Fig. 18D) 16-17 x 17-<br />
18 gm; C. barbata 16-17 x 16-17 pm.<br />
Parachimarrhis (Fig. 18F). Pollen grains tricolporate,<br />
23-25 x 29-31 jm; colpi acute at ends, membrane<br />
granular, pores with regular margin and annular<br />
thickening; exine irregularly reticulate (and<br />
columellate) throughout (Fig. 18F). The pollen of this<br />
monotypic genus (P. breviloba) is easily distinguishable<br />
from other genera because of its irregular reticulation<br />
with randomly missing links and its slightly<br />
larger dimensions.<br />
Dioicodendron (Fig. 18G,H). Pollen grains tricolporate,<br />
13-15 x 13-15 gm; colpi acute at ends<br />
(markedly evident), membrane sparsely minutegranular,<br />
pores with regular margin with thick annular<br />
thickening; exine thinly reticulate (and columellate)<br />
throughout, the reticulation decreasing in size<br />
toward the colpi. This monotypic genus (D. dioicum)<br />
is related to Chimarrhis, to which it is palynologically<br />
similar in its small grains, but differs in having thinly<br />
reticulate exine and well-demarcated colpi.<br />
Dolichodelphys (Fig. 19A,B). Pollen grains tricolporate,<br />
21-23 x 22-23 gm; colpi acute at ends (Fig.<br />
19B), membrane smooth around the pores and<br />
sparsely minute-granular at ends, pores with regular<br />
margin with thick annular thickening; exine irregularly<br />
reticulate (and columellate) in mesocolpi and<br />
foveolate at apocolpi. The pollen surface of this dubiously<br />
placed monotypic genus (D. chlorocrater) is<br />
ornamented with a widely irregular reticulation (Fig.<br />
19A,B).<br />
[Kajewskiella (Fig. 19C,D). Pollen grains<br />
tricolporate, 17-18 x 19-20 gm (K. polyantha); colpi<br />
round-truncate at ends (Fig. 19D), membrane densely<br />
and evidently granular, pores with irregular margin<br />
with irregular and granular thickening; exine regularly<br />
foveolate (and columellate) lumina decreasing in size<br />
toward the apocolpi].
Introduction to the Neotropical Genera Studied 49<br />
4;<br />
lo.<br />
rs:<br />
H. qutoiali Scl Ft. b w<br />
~~~~~~~~~~~~~~~~~~~~~~,H.<br />
Equatorial view. Scale bars: 10 gm."<br />
P~~,<br />
"?<br />
,rlra<br />
~r-.-lrr '<br />
*i<br />
11<br />
~ '<br />
lW<br />
',<br />
?<br />
,N~>. D~....<br />
~fi ._....1<br />
,, .<br />
I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
10<br />
FIG. 16. SEM photomicrographs ofpollen grains of Pogonopus, Pinckneya, and Kerianthera. A, B. Pogonopus exsertus<br />
(Croat 12895, MO). A. Equatorial view. B. Polar view. C. Pogonopus speciosus var. sandwithianus (Deiprete & Apreza<br />
6359), equatorial view. D. Pogonopus tubulosus ( Vargas 1719, GH), equatorial view. E, F. Pinckneya bracteata (Osborn<br />
s.n., LL). E. Polar view. F. Equatorial view. G, H. Kerianthera preclara (Santos & Co~iho 759, K). G. Polar view.<br />
~
50 Flora Neotropica<br />
..~~~~~~~~~~~~~~~~~~~~:<br />
.~ ..~ :~.!.~....<br />
_n<br />
-iS v .......................... 9<br />
- . :A _. B<br />
- ' _<br />
?S<br />
* i~ .. ~<br />
-@.<br />
.E . ..."...M., B<br />
~,<br />
't.<br />
_<br />
' "<br />
ee 1'S_~'??<br />
* ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~<br />
_~~~a E96 4i m G_<br />
Tilted view. B. E r o ......<br />
. . T<br />
^ . ' ' _ f f i 1<br />
ds. ' * .~~~~~~~~~~~~~~~~$u ...<br />
I g;$-):<br />
CI.1? cuesis (Wihotomi26o2aphso polae ran vf?iewrhi. FI. C. jaaicnisf(Pocto<br />
view (Bena 1066, P). H. Equatorial view (E1cnan H338-16, K). Scale bars: IO gm.<br />
.;]I' :..~..<br />
lii<br />
AS]~~~~~~~~~i<br />
2 * a _ _ 2 . % * * - _ y JP};...Z>~~~~~~~~~~~~~~~~~~<br />
_7 *W RJ_ @ _ #<br />
_~~~~~~~~~~~~~~~~~~~~~~~~~j. ;w*\w<br />
l~~lB~~I~<br />
; _s_* ,*s><br />
FIG. 17. SEMphotomicrogrphs of polle grains ofC imarris A, B.C priloa(are &Blo 87 F.A<br />
Tite viw .Eutra<br />
C.cbni Wih 22<br />
viw(Bn 16, ) H<br />
iw fuepne ri.C,D .Itfla(len60,F.C<br />
) oa iw.F .jmiess(rco 91,TX)plrve.GH<br />
qutria view, (E1unani~: H381,K.Sal as 01<br />
itdviw .Plrve.E<br />
.cms.G oa<br />
r<br />
19818 TEX Bepola view G, F). CyoA. Goa
Introduction to the Neotropical Genera Studied 51<br />
nor?<br />
JiA m...<br />
_ - x..,,:... :_<br />
4,",<br />
A<br />
.
ass<br />
52<br />
Flora Neotropica<br />
I<br />
*:J ~Zl:, ,Mt ... A ............ .. +_ B<br />
..9,<br />
I _ '<br />
FI.1.SM htmcorah a Ka n eskiela foln A, riso olcoepy<br />
oihdlhsclrcae<br />
896 qao S.A ial~ temsolium viwo<br />
theclpu.C, .Eqaoiliwo I.Kewila<br />
polyanth (Lvrk&Rddl 1346,: A. polrviwD,eutria cl iw as - C, 0Im<br />
FIG. 19. SEM photomicrographs ofvpollen grains ofDolichodelphys and Kajewskiella. A, B.Dolichodelphys chiorocrater<br />
(Cuatrecasas 8926, US). A. Equatorial view of the mesocolpium. B. Equatorial view of the colpium. C, D. Kajewskiella<br />
polyantha (Lavarak & Ridsdale 31346, A). C, polar view. D, equatorial view. Scale bars: A-F = 10 gtm.<br />
WOOD ANATOMY<br />
There is little information regarding the wood<br />
anatomy of the <strong>Rubiaceae</strong>. Solereder (1908), in his<br />
Systematic Anatomy ofthe Dicotyledons, reported that<br />
the members of the Condamineeae s.l. (among many<br />
other tribes) have sclerenchyma [as "prosenchyma"]<br />
with bordered pitting. Metcalfe and Chalk (1950), in<br />
their extensive Anatomy of the Dicotyledons, merely<br />
listed the genera under general categories and described<br />
the wood of the Condamineeae (sensu Hooker)<br />
without very distinctly bordered pits in Pinckneya.<br />
I found scanty information about the development<br />
of the cork cambium, which Solereder (1908) observed<br />
in Pinckneya to originate from a subepidermal<br />
layer of cells. In an addenda Solereder (1908) reported<br />
that Pitard [?] observed that in Chimarrhis (among<br />
many other genera) "the pericycle contains isolated<br />
bundles of bast-fibers."<br />
Accurate work on the wood anatomy of the family<br />
was presented by Koek-Noorman and colleagues<br />
as follows: with vessels relatively small (
Introduction to the Neotropical Genera Studied 53<br />
Koek-Normann (1970, 1972), Koek-Noorman and transferred to the Condamineeae by Andersson and<br />
Hogeweg (1974), and Koek-Noorman and Puff(1983) Persson (1991). Coutarea has recently been transreported<br />
that the tribes Gardenieae, Ixoreae, Vangueri- ferred to the Catesbaeeae (Delprete, 1995a, 1996d), and<br />
eae, Isertieae, Naucleae, Mussaendeae, Hedyotideae, Exostema was treated as a closely related taxon (Delprete,<br />
Anthospermeae, and Rubieae are all homogeneous in 1995a, 1996d; McDowell, 1996, and pers. comm.).<br />
having only fiber-tracheids, while Psychotrieae, Pae- Koek-Noorman and Hogeweg (1974) performed<br />
derieae, Morindeae, Guettardeae, Hillieae, and Hame- a cluster analysis using 125 wood anatomical characlieae<br />
are homogeneous in having only libriform fibers. ters, but a complete data matrix of such characters was<br />
Koek-Noorman (1969b), in her review of the Ru- not published. Three dendrograms for the Cinchoneae,<br />
biaceae of Suriname, discussed the wood anatomy and <strong>Rondeletieae</strong>, and Condamineeae were produced, and<br />
taxonomy of Chimarrhis as follows: "Schumann was they all displayed a basal "partition in two groups, one<br />
not sure whether C. turbinata was rightly placed in of which shows libriform fibers, the other one fiber<br />
this genus. Ducke (1922) referred it to a new genus, tracheids." In the Condamineeae they reported that<br />
viz. Pseudochimarrhis. According to Bremekamp "the parenchyma pattern seems to be less connected<br />
(1934) and Steyermark (1965) Chimarrhis and with the fiber type, the arrangement of the vessels, and<br />
Pseudochimarrhis are not separable. Record and Hess the presence of crystal sand." Bikkia, Morierina, and<br />
(1943) report that the wood.of C. turbinata greatly Portlandia [which were transferred to the Catesbaeeae<br />
resembles C. longistipulata [= C. microcarpa], the by Delprete (1996d)] formed one cluster. Bikkia camplace<br />
of which is not in doubt. They differ only in panulata is reported as "dissimilar in the frequency<br />
minor features." In the same article Koek-Noorman of radial vessel multiples, relatively large intervascu-<br />
(1969b) described the wood of Chimarrhis as having lar pits, uniseriate rays, relatively large ray-vessel<br />
septate fibers, narrowly vasicentric parenchyma, with pitting, presence of crystal sand, and vague growth<br />
6-12 vessels per mm2 and up to 150 pm wide, upright rings." But this set of characters could be influenced<br />
ray cells up to 50 um, without sheath cells nor broad by environmental factors.<br />
rays. As a conclusion she states that the wood of The monotypic Pinckneya was pointed out by<br />
Chimarrhis resembles that of the Cinchoneae (of Koek-Noorman and Hogeweg (1974) as exceptional<br />
which she studied Capirona, Ferdinandusa, and Cin- to the Condamineeae (sensu Hooker) in having semichona)<br />
more than that of the <strong>Rondeletieae</strong> (of which ring-porous wood, with tangential pore chains and<br />
she studied only Elaeagia and Warszewiczia). concentric parenchyma bands. They did not mention<br />
Koek-Noorman and Hogeweg (1974) presented a that this is the only non-tropical genus analyzed (ensurvey<br />
of some representative taxa of the tribes Van- demic to Georgia and South Carolina, North America)<br />
guerieae, Cinchoneae, Condamineeae, and <strong>Rondeletieae</strong> and that its wood anatomy is certainly influenced by<br />
(all sensu Bremekamp, 1966), which is the only study the seasonal climate to which this genus is exposed.<br />
in which representative genera of the Condamineeae Considering the dry-seasonal forests to which Pogonwere<br />
analyzed. The genera of the Condamineeae opus is endemic, I would also expect its wood to be<br />
(sensu Bremekamp, 1966) analyzed by Koek-Noorman ring-porous (not reported by Koek-Noorman).<br />
and Hogeweg (1974) were Condaminea (C. corym- Pogonopus and <strong>Rustia</strong> (with Calycophyllum and<br />
bosa), <strong>Rustia</strong> (R. formosa), Chimarrhis (C. cymosa, Mussaendopsis) were reported to have fibers clearly<br />
C. microcarpa [as C. longistipulata], C. turbinata, C. septate and with "scanty" pits on their tangential walls.<br />
hookeri), Pogonopus (P. speciosus), Pinckneya (P. bracteata<br />
[as P. pubens]), Bikkia (B. campanulata, B. palauensis),<br />
Portlandia (P. grandiflora), and Morierina (M.<br />
montana). They reported that the Cinchoneae, <strong>Rondeletieae</strong>,<br />
and Condamineeae were rather heterogeneous<br />
Finally, Koek-Noorman and Hogeweg (1974) performed<br />
two cluster analyses of the Vanguerieae,<br />
Cinchoneae, <strong>Rondeletieae</strong>, and Condamineeae combined,<br />
one with equal character weighting and the<br />
other with preferential character weighting. Both<br />
in having both libriform fibers and fiber-tracheids. analyses also showed a basal bipartition so that gen-<br />
This finding is probably due to the poor delimitation era of various tribes were scattered on the dendroof<br />
the three tribes at that time. Unfortunately, the au- grams, and sometimes species of the same genus were<br />
thors discussed only the characteristics of the genera even placed on remote branches. According to these<br />
exceptional to the Condamineeae, and the descriptions two analyses, the authors concluded that none of the<br />
of wood anatomical characters for most genera stud- characters studied has reliable diagnostic value, and<br />
ied (<strong>Rustia</strong>, Condaminea, Pogonopus) are only frag- they were persuaded "not to pursue the attempt to<br />
mentary. Koek-Noormann and Hogeweg (1974) re- define the taxonomic tribes under discussion [Ronported<br />
Exostema and Coutarea to be exceptional for deletieae, Condamineeae, and Cinchoneae as defined<br />
the tribe Cinchoneae. The two genera were subsequently at that time] on the base of wood anatomy."
54 Flora Neotropica<br />
My opinion is that wood anatomy can certainly<br />
constitute an important set oftaxonomical characters<br />
(mainly when tempered with environmental considerations<br />
and geographical distribution) in attempting<br />
to reconstruct a sound evolutionary study of the <strong>Rubiaceae</strong>.<br />
No anatomical studies of the Cinchoneae-<br />
<strong>Rondeletieae</strong>-Condamineeae complex has been performed<br />
since the works ofKoek-Noorman, and I strongly<br />
encourage future workers to undertake such studies.<br />
PRESENCE OF CALCIUM OXALATE CRYSTALS<br />
Solereder (1908), in his anatomical overview of<br />
dicotyledons, listed the presence of calcium oxalate<br />
crystals in many rubiaceous genera. He reported the<br />
Condamineeae (sensu Hooker) as having "Crystalsand,<br />
occasionally together with clustered crystals,"<br />
which he observed in Bikkia, Chimarrhis, Condaminea,<br />
Pinckneya, Pogonopus, Portlandia, and <strong>Rustia</strong>. Of the<br />
buds were collected and preserved in 4:3:1 (chloro-<br />
form:95% ethanol:acetic acid). Young anthers were<br />
extracted from the preserved floral buds, squashed,<br />
and stained with aceto-orcein. Only three successful<br />
counts were obtained: <strong>Rustia</strong> occidentalis (Delprete<br />
6329, TEX) n = 11, Condaminea corymbosa (Delprete<br />
6049, TEX) n = 11, and Pogonopus speciosus var. sand-<br />
withianus (Delprete & Apreza 6359, TEX) n = 11. In<br />
the three genera observed, the chromosomes were very<br />
small and stained darkly.<br />
The counts of Condaminea, <strong>Rustia</strong>, and Pogon-<br />
opus support the transfer of the subtribes Condami-<br />
neeinae and Pinckneyinae (the genera of the group<br />
here studied) to the <strong>Rondeletieae</strong>, since this tribe has<br />
chromosome numbers on a base of x = 9, 10, 11, and<br />
of small size (Kiehn, 1995, and pers. comm.).<br />
genera of the Condamineeae (sensu Hooker), Metcalfe<br />
POLLINATION AND OTHER<br />
PLANT-ANIMAL INTERACTIONS<br />
and Chalk (1950) reported the presence of crystalsand,<br />
either alone or accompanied by clustered crystals,<br />
in the same genera previously reported by<br />
Solereder (1908). None of the genera of the Condamineeae<br />
(sensu Hooker) has ever been reported to<br />
have raphides (and they are probably absent).<br />
Bremekamp (1966) was the first worker to consider<br />
the presence of calcium oxalate crystals as an<br />
important taxonomic character in the <strong>Rubiaceae</strong>, using<br />
it to define subfamilies, tribes, and genera.<br />
Verdcourt (1958) examined thousands of specimens<br />
Birds. <strong>Rustia</strong> has attained a high degree of floral<br />
diversity along with a variety of pollination syndromes<br />
(lap- and buzz-pollination by bees, hummingbird pollination,<br />
butterfly-pollination/visited). The pendulousflowered<br />
species of <strong>Rustia</strong> are probably pollinated by<br />
hummingbirds (at least this appears to be so for <strong>Rustia</strong><br />
thibaudioides; pers. obs.); they have tubular pinkwhite<br />
corollas (R. thibaudioides), red with green lobes<br />
(R. rubra), green (R. viridiflora), and yellow-green (R.<br />
schunkeana).<br />
in order to confirm the validity of this character and<br />
declared it to be very important (but not at the<br />
subfamilial level), because of the universal presence<br />
of raphides in some tribes, and their consistent absence<br />
in some others. Verdcourt (1958) reported the Condamineeae<br />
(which he included in the <strong>Rondeletieae</strong>)<br />
as having crystal-sand, as observed by all previous<br />
workers. Koek-Noorman and Hogeweg (1974) used<br />
Bees. <strong>Rustia</strong> occidentalis is buzz-pollinated by<br />
bees of the genera Euglossa and Melipona (see details<br />
under R. occidentalis); this is the first report of<br />
buzz-pollination in the <strong>Rubiaceae</strong>. The fleshy-flowered<br />
R. bilsana, R. dressleri, and <strong>Tresanthera</strong>, also<br />
having anthers only half-exserted and convex toward<br />
the center, are probably also buzz-pollinated by bees,<br />
but this has not been observed. <strong>Rustia</strong> thibaudioides<br />
this character (classified as raphides, rhombic crystals,<br />
elongated crystals, druses, and crystal sand) in<br />
their anatomical analysis of the Cinchoneae-Condais<br />
visited by bees of the genera Melipona, Ceratina,<br />
and Trigona (pers. obs.), but their effectiveness as pollinators<br />
has not been tested. The minute white flowmineeae-<strong>Rondeletieae</strong><br />
(all have crystal-sand but lack<br />
raphides), but did not publish their observations.<br />
I agree with Halle (1961, 1967) and Robbrecht<br />
(1988) that presence and kinds of raphides evolved<br />
ers of R. costaricensis are presumably lap-pollinated<br />
by small bees, which at least are frequent visitors. In<br />
Condaminea, at anthesis, the filaments are deflexed<br />
toward the lower side of the corolla tube and the anseveral<br />
times in the <strong>Rubiaceae</strong>. Nevertheless, the presence,<br />
quality, and quantity of raphides remains an<br />
important character for phylogenetic purposes, and in<br />
defining tribal and generic groups in the family.<br />
thers are set parallel to each other and opening upward<br />
(Fig. 46A-F), forming a landing area for large<br />
bees that pollinate (lap-pollination) the flowers. The<br />
minute, sweet-scented flowers of Chimarrhis are visited<br />
and pollinated by small bees. Reports of visita-<br />
KARYOLOGY<br />
For genera included here a few counts were obtained<br />
with the following methodology. Young floral<br />
tion by bees were obtained from herbarium labels of<br />
C. latifolia and personally observed in C. glabriflora<br />
and C. hookeri.
Introduction to the Neotropical Genera Studied 55<br />
Wasps. <strong>Rustia</strong> thibaudioides has been observed to<br />
be visited (but not pollinated), and robbed laterally,<br />
by unidentified wasps, followed by ants entering<br />
through their incisions. Flower buds ofPogonopus speciosus<br />
var. sandwithianus have also been observed to<br />
be robbed primarily by wasps and secondarily by ants.<br />
Butterflies. Flowers of <strong>Rustia</strong> thibaudioides and<br />
Pogonopus speciosus var. sandwithianus are visited<br />
(and probably pollinated) by unidentified butterflies.<br />
Specimen labels further report that the flowers of<br />
Chimarrhis latifolia are visited by butterflies (but<br />
probably not pollinated by them).<br />
Ants. Some individuals of <strong>Rustia</strong> thibaudioides are<br />
guarded by unidentified small ants; the same ants were<br />
also observed to feed on the floral nectaries after en-<br />
tering the floral buds through lateral incisions previ-<br />
ously made by wasps. Several individuals of Con-<br />
daminea corymbosa were observed to be guarded by<br />
ants, but I consider them to be only casual visitors.<br />
Ants were seen robbing closed flower buds ofR. thi-<br />
baudioides and P. speciosus var. sandwithianus<br />
through lateral incisions made by wasps.<br />
Mites. Leaf domatia are usually associated with<br />
mites. All of the species of <strong>Rustia</strong>, Picardaea,<br />
Chimarrhis, and Pogonopus [and Pinckneya] with<br />
domatia have been found to harbor several unidenti-<br />
fied taxa of mites, but their possible mutualism is yet<br />
to be shown.<br />
Fruitgalls. The ovaries of Chimarrhis and <strong>Rustia</strong><br />
can be infested by insects that cause fruit galls. Spe-<br />
cifically, fruit galls have been observed in Chimarrhis<br />
barbata, C. turbinata, C. glabriflora, C. hookeri, and<br />
<strong>Rustia</strong> schunkeana. The insects that cause these galls<br />
have not been identified, but they probably belong to<br />
the Coleoptera.<br />
HABITAT AND ECOLOGY<br />
Most genera studied in the present treatment occur<br />
in a broad range of habitats and are represented in<br />
several forest types at several levels, often occupying<br />
specific ecological niches. <strong>Rustia</strong> occurs in several<br />
kinds of habitats and is mostly represented by<br />
medium-tall shrubs that occur in evergreen-seasonal<br />
forests, at low elevations (300-600 m), and usually<br />
in proximity to running water. Several species, however,<br />
are medium-size trees occurring in tropical cloud<br />
forests and rain forests, at medium elevations (500-<br />
2000 m). <strong>Rustia</strong> occidentalis is exceptional, occurring<br />
as a small shrub in swamplands and slow-moving<br />
fresh water at low coastal elevations (near sea level).<br />
<strong>Tresanthera</strong> is the sister genus of <strong>Rustia</strong>, and is represented<br />
by medium-sized trees endemic to coastal<br />
Caribbean rain forests.<br />
Condaminea is most commonly represented by<br />
shrubs. It is very widespread and easily adaptable to<br />
diverse environments (in Central and South America),<br />
occupying dry-seasonal forests, cloud forests, rain<br />
forests, and being especially common in disturbed<br />
secondary forests and on roadsides of recently built<br />
Andean roads (see ecology of C. corymbosa). Its tolerance<br />
to elevational gradients is also remarkable,<br />
ranging from coastal forests (near sea level), to<br />
Andean slopes (up to 2000 m), and down to Amazonian<br />
rain forests.<br />
Picardaea is a rare shrub restricted to limestone<br />
cliffs of seasonal coastal forests of the Caribbean islands<br />
(Cuba and Hispaniola).<br />
Pogonopus is mostly represented by tall shrubs<br />
occurring on slopes in dry-seasonal, deciduous, transitional<br />
tropical forests at medium elevations (500-<br />
1400 m), and rarely in cloud forests (P. tubulosus) of<br />
Central and South America, at medium-high elevations<br />
(1600-2800 m). It is usually encountered in<br />
proximity to creeks and running water.<br />
Chimarrhis is represented by large trees with mature<br />
individuals present at the canopy levels of Caribbean<br />
seasonal forests, riverine forests, rain forests<br />
and cloud forests, to lowland non-inundated Amazon<br />
forests (terra firme), from low to medium elevations<br />
(300-2000 m).<br />
Dioicodendron is a rare tall shrub (to small tree)<br />
present in the understory of seasonal-dry Andean forests<br />
and cloud forests at relatively high elevations<br />
(900-3500 m).<br />
Molopanthera is a tree occurring in seasonal Atlantic<br />
forests of southern Brazil, at medium elevations<br />
(300-700 m).<br />
Dolichodelphys is another rare shrub (to small<br />
tree), but generally widespread, occurring in Andean<br />
cloud forests and rain forests, at medium elevations<br />
(400-1400 m).<br />
Parachimarrhis is a rare tree known from a few,<br />
isolated collections from lowland, seasonally inundated<br />
Amazonian basin.<br />
USES<br />
Members of this group have medicinal, economic,<br />
and ornamental uses. Pogonopus has been used as a<br />
secondary source of quinine to cure malaria and intermittent<br />
fevers. Pogonopus exsertus is used as a local<br />
antimalarial remedy in Guatemala (Standley 79549,<br />
label data). The Colombian local names "quina" and
56 Flora Neotropi<br />
"quina morada" (Delprete & Apreza 6359) for P. speciosus<br />
are suggestive of the local medicinal use of this<br />
species. The single species most often used to cure<br />
malaria is P. tubulosus (as "quina" or "sacha quina"),<br />
which was extensively used by Peruvian, Bolivian,<br />
and Argentinean indigenous tribes as a remedy for<br />
malaria and was later adopted by European colonists<br />
(see details under P. tubulosus). Weddell (1854) reported<br />
that the bark of P. tubulosus [named by him<br />
Howardiafebrifuga (Wedd.) Wedd.] was found "on<br />
the English market" by Elliot Howard, a contemporary<br />
quinologist; Howard isolated an alkaloid from its<br />
bark, which Weddell (1854) named "Howardine" and<br />
described it to have a bitter taste with a styptic aftertaste.<br />
The alkaloids that were later isolated from the<br />
bark of P. tubulosus are pogonopamine, pogonopeine,<br />
pogonopidine, pogonopine (Willaman & Schubert,<br />
1966), and tubulosine (Raffauf, 1970). Tubulosine is<br />
currently being studied for its antitumor properties in<br />
brine shrimp, crown gall tumors of potato, and human<br />
tumor cells (Ma, 1991; Ma et al., 1990). The bioactive<br />
properties of tubulosine have also been studied for<br />
effectively inhibiting the process of peptide chain elongation<br />
in eukaryotic polysomes (Carrasco et al., 1976).<br />
Because of its very attractive calycophylls,<br />
Pogonopus has been cultivated as an ornamental shrub<br />
throughout its natural range and elsewhere. Cultivated<br />
individuals of Pogonopus exsertus are reported from<br />
botanical gardens in the United States (Florida and<br />
Hawaii), Honduras, Cuba, Puerto Rico, and Brazil<br />
(see Cultivated Specimens Examined of this species).<br />
Pogonopus tubulosus is the species with the largest<br />
calycophylls (to 15 cm long and 13 cm wide!), and is<br />
widely cultivated in botanical and home gardens in<br />
Peru, Bolivia, and Argentina.<br />
The large trees of Chimarrhis offer a much-appreciated<br />
pale yellow hardwood used for construction<br />
purposes. The woods of this genus are reported as<br />
follows: Chimarrhis parviflora and C. latifolia as<br />
valuable hardwoods in Costa Rica and Panama, C.<br />
cubensis as dark yellow hardwood in Cuba, C.<br />
jamaicensis as good construction wood in Jamaica,<br />
and C. cymosa as a much sought-after construction<br />
wood in the Lesser Antilles. Chimarrhis glabriflora<br />
and C. hookeri offer excellent timbers used by Ecuadorian<br />
and Peruvian indigenous tribes for construction<br />
of houses and boats; these two species are also<br />
reported (Duke & Vasquez, 1994) to be used for establishing<br />
living fences and as sources of good firewood.<br />
In Venezuela the wood of C. microcarpa is<br />
used to make wheel-spokes, beams, and posts. The<br />
pale yellow hardwood of C. turbinata and C. barbata<br />
are used by indigenous tribes of French Guiana and<br />
Brazil for construction of houses. Plank-wood from<br />
the large buttresses of these two Amazonian fort<br />
giants is commonly used by indigenous tribes to ma<br />
canoe paddles, and is locally called "pau de remo"<br />
Brazil and "bois pagaie" in French Guiana. The<br />
brous bark of C. glabriflora, C. hookeri, and C. turi<br />
nata is used by indigenous people of Amazonian E<br />
uador, Peru, French Guiana, and Brazil to make rope<br />
Ducke (1922) reported that the wood of Par<br />
chimarrhis is brownish yellow and of good quality<br />
SPECIES CONCEPTS<br />
Throughout this study I have used as a worki<br />
hypothesis the taxonomic species concept of Dan<br />
and Heywood (1963): "assemblages of individu;<br />
with morphological features in common and separal<br />
from other such assemblages by correlated morph<br />
logical discontinuities in a number of features." I ha<br />
tried to reevaluate with a critical approach many ml<br />
phological characters that have been historically treat<br />
as fundamental to the macroclassification of the R<br />
biaceae (e.g., valvate corolla aestivation, fruit flesi<br />
ness, placentation, vestiture, leaf shapes, stipul<br />
etc.). At the same time, I tried to find new charactc<br />
that have not previously been used in generic a<br />
specific delimitations (e.g., leaf domatia, lea<br />
branchlet succulence, stipule persistence and morph<br />
ogy, stipular colleters, inflorescence architectu]<br />
anther morphology, etc.). Leaf shape and gene]<br />
vestiture have been treated as useful secondary ch;<br />
acters for the delimitation of species, but I conclud<br />
that they are better used at most at the varietal lev<br />
Throughout my treatment only one infraspeci<br />
rank has been recognized, the variety. Varietal ra<br />
was assigned when morphogeographic population<br />
elements were discerned, especially when the ch;<br />
acters that marked them tended to intergrade in regic<br />
of contiguity. This follows the practice of many cc<br />
temporary practitioners, including Cronquist (pe<br />
comm.), Fryxell (pers. comm.), Gentry (pers. comrr<br />
Takhtajan (pers. comm.), Turner (pers. comm.), a<br />
Webster (pers. comm.), among numerous others.<br />
SYSTEMATIC TREATMENT 01<br />
THE GENERA STUDIED<br />
RUSTIA<br />
1. <strong>Rustia</strong> Klotzsch in Hayne, Getreue Darstell. Ge<br />
14: tabs. 14, 15, 555. 1846; Chamisso<br />
Schlechtendal, Linnaea 4: 179-181. 1829;<br />
Candolle, Prodr. 4: 360-361. Sep 1830; Karste<br />
Linnaea 30: 150-151. 1859; J. D. Hooker<br />
Bentham & Hooker, Gen. PI. 2: 45. 187<br />
Bentham, Bot. Voy. Sulph. 104. 1844; Baill
Systematic Treatment of the Genera Studied 57<br />
Hist. PI. 7: 471. 1880; Hemsley, Biol. Centr.-<br />
Amer. Bot. 2: 14. 1881; Humboldt & Bonpland,<br />
PI. Aequin. 1: 138. 1808; Karsten in Engler, Bot.<br />
Jahrb. Syst. 8: 354-360. 1887; Schumanm in<br />
Martius, Fl. Bras. 6(6): 260-264. 1889; Schumanm<br />
in Engler & Prantl, Nat. Pflanzenfam. 4(4): 18.<br />
1891; Standley, N. Amer. Fl. 32: 7. 1918; Standley,<br />
in each locule, horizontally inserted; immature fruits<br />
green to red, carnose. Capsules coriaceous to woody,<br />
globular, turbinate, or obovate; dehiscing<br />
loculicidally; disk septicidal dehiscence present in old<br />
capsules of some species. Seeds small, pointed at the<br />
ends, angular or with minute marginal wings.<br />
Publ. Field Columbian Mus., Bot. Ser. 7: 26. 1930;<br />
Steyermark in Lasser, Fl. Venez. 9: 261. 1974;<br />
Dwyer, Ann. Missouri Bot. Gard. 67: 482-485.<br />
1980; D. Simpson, Phytologia 33: 4-8. 1976;<br />
Delprete, Novon 5: 133-139. 1995; Delprete,<br />
Brittonia 48: 29-34. 1996; Delprete, Novon 8:<br />
<strong>Rustia</strong> is unique in the <strong>Rubiaceae</strong> in having poricidal<br />
anthers opening by two distinct apical pores. The<br />
closely related <strong>Tresanthera</strong> also has anthers pointed<br />
at apex, but its dehiscence is by one lateral triangular<br />
pore (below the apex); it also has larger fruits, and<br />
flattened and truncate seeds. Both <strong>Rustia</strong> and<br />
231-234. 1998; Delprete in Harling & Andersson, <strong>Tresanthera</strong> are also unique in the <strong>Rubiaceae</strong> in hav-<br />
Fl. Ecuador (in press). 1999. Type species. <strong>Rustia</strong> ing leaves with "pellucid glands." These two genera<br />
formosa (Chamisso & Schlechtendal ex de are in turn closely related to Condaminea, with which<br />
Candolle) Klotzsch.<br />
they share filaments attached at base of corolla tube<br />
and multiseeded loculicidal<br />
Henlea H. Karsten, Linnaea 30: 151. 1859. capsules with minute<br />
Type spe- seeds<br />
cies. Henlea thibaudioides H. Karsten [= <strong>Rustia</strong> thi- horizontally inserted. Condaminea differs from<br />
baudioides (H. Karsten) Delprete] non Henlea Grise- <strong>Rustia</strong> and <strong>Tresanthera</strong> in having four foliose stipules<br />
bach, nom. illeg., Abh. K6nigl. Ges. Wiss. G6ttingen per node (vs. two per node), anthers that open by lon-<br />
9: 37. 1860 [Apr 1861] (Malpighiaceae). gitudinal slits, deciduous calyx, sessile to subpetiolate<br />
Stomandra Standley, Publ. Field Mus. Nat. Hist., Bot. leaves, and leaf blades without pellucid glands (as in<br />
Ser. 23: 247. 1947. Type species. Stomandra costa- the vast majority of <strong>Rubiaceae</strong>).<br />
ricensis Standley [= <strong>Rustia</strong> costaricensis (Standley) The flowers of <strong>Rustia</strong> consistently have valvate<br />
D. H. Lorence].<br />
aestivation (sometimes valvate-reduplicate), are<br />
protandrous (the stigma is receptive only after the<br />
Shrubs to trees, sometimes with small buttresses; anthers are dried up), and vary considerably in size,<br />
bark grayish-pale brown, smooth. Stipules interpetiolar,<br />
below petiole attachments, free at base, entire,<br />
narrowly triangular, readily caducous. Leaves petiolate,<br />
lanceolate, elliptic to ovate, usually acuminate<br />
shape, and habit. In R. occidentalis, R. dressleri, and<br />
R. bilsana the flowers are fleshy and usually erect,<br />
with anthers convex toward the center and closely<br />
united, and buzz-pollinated by bees. In R. thibaudiat<br />
apex, pellucid-punctate; petioles thickened at base,<br />
rarely pulvinate; domatia absent or a tuft of sparse hairs.<br />
Inflorescences terminal, solitary, paniculate, racemoid<br />
secundiflorous or thyrsoid secundiflorous; lateral<br />
branches (when present) opposite, decussate, usually<br />
oides, R. rubra, R. viridiflora, and R. schunkeana the<br />
flowers are non-fleshy, pendulous, brightly colored<br />
(red-green), with anthers included or partially exserted<br />
(consistent within the species), and commonly visited<br />
by hummingbirds, butterflies, and bees. In R. alba,<br />
monochasial. Flowers protandrous; hypanthium narrowly<br />
obconical, turbinate to cupulate. Calyx cupular<br />
or extremely reduced, with barely visible lobes or truncate,<br />
persistent. Corolla tubular, cupular to campanulate,<br />
with more or less spreading lobes, thin to fleshy,<br />
white, red, green, or yellow-green; tube short to long<br />
tubular, glabrous to puberulent inside; lobes (4-)5<br />
(-6), deltoid to narrowly triangular; aestivation valvate<br />
to valvate-reduplicate with contact zone. Stamens<br />
(4-)5, partially or completely exserted, attached<br />
near the base of the corolla tube; anthers dorsifixed<br />
near the base, opening by 2 apical pores. Pollen<br />
tricolporate, exine reticulate to foveolate (sometimes<br />
minutely echinate). Style exserted; style branches<br />
short, thick, ovate to oblong. Ovary 2-celled,<br />
obconical to turbinate; placentation axile; ovules many<br />
R. formosa, R. simpsonii, and R. gracilis the flowers<br />
are non-fleshy (thin), white, either erect or nodding,<br />
with anthers exserted well beyond the corolla, and<br />
commonly visited by bees. None of the species has<br />
been reported to have flowers with any particular fragrance,<br />
except for R. formosa and R. alba (pers. obs.).<br />
The seeds of <strong>Rustia</strong> are minute, horizontally inserted,<br />
non-winged, and irregularly shaped, varying<br />
from apically truncate to uncinate, and 3-4-sided.<br />
<strong>Rustia</strong> is encountered as small shrubs, tall shrubs, to<br />
medium-size trees. At least one species (R. alba, pers.<br />
obs.) has been found with small buttresses. The wood<br />
is usually soft and of no particular use, and the bark<br />
is usually smooth to rugose, and pale brown to grayish.<br />
<strong>Rustia</strong> has 14 species, ranging from Nicaragua to<br />
southern coastal Brazil. The center of diversity of
58 Flora Neotropica<br />
<strong>Rustia</strong> is in South America. Ecuador has the largest species (R. secundiflora [= R. thibaudioides], R.<br />
number of species (six), three of which are endemic. angustifolia, and R. gracilis) from South America, and<br />
Four other species are endemic to southern Brazil, synonymized R. pohliana and R. sellowiana under R.<br />
mostly in the states of Slo Paulo and Rio de Janeiro. formosa. He correctly included Henlea under <strong>Rustia</strong><br />
but erroneously transferred Henlea rosea [= Laden-<br />
Taxonomic History<br />
bergia muzonensis (Goudot) Standl.] to <strong>Rustia</strong><br />
The type species of <strong>Rustia</strong> was first presented by (Schumann, 1891).<br />
Chamisso and Schlechtendal (1829) as Exostema Urban (1931) described <strong>Rustia</strong> haitiensis from<br />
formosum but without description, therefore as nomen specimens without flowers and with old capsules withnudum<br />
(for detailed information about the nomencla- out seeds. This species remains dubious because only<br />
ture of this species see <strong>Rustia</strong>formosa). De Candolle sterile material was available, and it almost certainly<br />
(1830) validated Exostemaformosum Cham. & Schltdl. does not belong to <strong>Rustia</strong> (see doubtful species).<br />
ex DC., as part of Exostemma sect. Pseudostemma; Standley (1930a) transferred Henlea splendens to<br />
after a brief description, he stated, "Species Brasili- <strong>Rustia</strong>, apparently without noticing the obvious simianae.<br />
An Genus proprium?" Under sect. Pseudo- larity to R. secundiflora [= R. thibaudioides] and to<br />
stemma he included E. cuspidatum St.-Hil. [= Bathysa H. thibaudioides [= R. thibaudioides]. He also maincuspidata<br />
(St.-Hil.) J. D. Hook.], E. australe St.-Hil. tained R. rosea [= Ladenbergia muzonensis (Goudot)<br />
[= Bathysa autralis (St.-Hil.) J. D. Hook], E. for- Standl.; see excluded species] and described R. ferrumosum<br />
[= <strong>Rustia</strong>formosa], and E. souzanum Martius ginea [=Bathysa sp.] (Standley, 1916), R. iserii [=Aliex<br />
DC. [Bathysa sp.?].<br />
bertia isernii] (Standley, 1940), R. longifolia [=Dolicho-<br />
<strong>Rustia</strong> was founded by Johann Friedrich Klotzsch delphys chlorocrater] (Standley, 1930b) and R.<br />
(1846), who dedicated its name to Dr. Rust ("Dem venezuelensis (Standley & Steyermark, 1953).<br />
Andeken des verbosteren Prasidenten Dr. Rust Standley (1947) established the genus Stomandra<br />
gewidmet"). Under the genus <strong>Rustia</strong> Klotzsch (1846) [= <strong>Rustia</strong>] which he described as different from <strong>Rustia</strong><br />
placed three species: R. formosa, R. sellowiana (vali- in having smaller flowers and corolla tube shorter than<br />
dated by direct reference to Exostemmaformosum var. the lobes. Both of these characters are not sufficient to<br />
leprosum Cham. & Schltdl.), and R. pohliana (which separate the two genera, and Lorence (in Burger & Tayhe<br />
described).<br />
lor, 1993) recently transferred S. costaricensis to <strong>Rustia</strong>.<br />
Karsten (1859) established the genus Henlea with Donald Simpson (1976) published a short revision<br />
H. thibaudioides, later adding (Karsten, 1861) H. of the <strong>Rustia</strong> species occurring in western South<br />
splendens [= R. thibaudioides]. Henlea has been America, and validated Standley's unpublished R.<br />
treated as a dubious taxon (Robbrecht, 1988, 1993). rubra. In addition, he correctly transferred R. isernii<br />
In the present treatment all the species of Henlea have to Alibertia, erroneously treated R. longifolia [= Dolibeen<br />
treated as synonymous with either Ladenbergia chodelphys chlorocrater] as synonymous with Tresor<br />
<strong>Rustia</strong>. Henlea thibaudioides (the type species of anthera condamineoides, and declared that the short<br />
the genus) is here transferred to <strong>Rustia</strong>, and Henlea description of Henlea thibaudioides [= R. thibaudiis<br />
treated as synonymous with <strong>Rustia</strong> (see discussion oides] "is insufficient to permit any taxonomic deciunder<br />
R. thibaudioides, and rejected species below). sion on its status" and that "it may be something other<br />
Baillon (1880: 471-472) included both Tres- than <strong>Rustia</strong>."<br />
anthera and Henlea in <strong>Rustia</strong>, stating that <strong>Rustia</strong> has Dwyer (1980) described <strong>Rustia</strong> panamensis [= R.<br />
"poricidal or shortly rimose [anthers], enclosed or costaricensis (Standl.) Lorence], without noticing its<br />
exserted" and that it is closely related to Condaminea. close resemblance to Stomandra costaricensis.<br />
Hemsley (1881) transferred Exostemma (sect. Delprete (1995, 1996, 1998) recently described<br />
Pseudostemma) occidentale Benth. to <strong>Rustia</strong> (see dis- five new species of <strong>Rustia</strong> from Panama and western<br />
cussion under R. occidentalis) without comment. South America (R. dressleri, R. alba, R. viridiflora,<br />
Schumann (1889) described three additional <strong>Rustia</strong> R. schunckeana, and R. bilsana).<br />
Key to the species of <strong>Rustia</strong><br />
1. Corollas 1 cm long (assuming that R. venezuelensis has flowers >1 cm long).
Systematic Treatment of the Genera Studied 59<br />
3. Flower buds clavate, rounded at apex; corollas 14 mm long (South America).<br />
5. Leaf blades broadly elliptic, to broadly obovate; width >/2 the length.<br />
6. Corolla lobes /3 of corolla length; calyces cupular and membranous; leaves with 16-21<br />
secondary veins each side (mountains of N Ecuador) .................................................................... 4. R. alba<br />
6. Corolla lobes '/3-/2 of corolla length; calyces much reduced; leaves with 12-16 secondary<br />
veins each side (central and coastal Brazil) ........................................ .......................... 11. R. formosa<br />
5. Leaves narrowly elliptic to oblanceolate to lanceolate; width
60<br />
,~~~<br />
?? , .<br />
? c~~~~~~~~~~~~~?<br />
Systematic Treatment of the Genera Studied 61<br />
FIG. 21. Distribution of <strong>Rustia</strong> costaricensis (closed circles) and R. dressleri (closed triangles).<br />
below; pellucid punctate; primary and secondary veins puberulent hairs (ca. 0.05 mm long) 0.5-0.6 mm from<br />
glabrous, prominent below; secondary veins 12-20 the base, at the same point as filament attachments;<br />
each side; tertiary veins starting subparallel and openly lobes 4-6, ca. /3 of corolla length, 1-1.5 mm x ca. 1.5<br />
reticulate toward the center; petioles 2.5-4.5(-6) cm mm, ovate to triangular, glabrous, minutely papillose<br />
long, 2-3 mm thick, terete or adaxially flattened; pul- at tip of margins inside. Stamens (4-)5, partly exvinus<br />
absent but the petiole thickened, becoming serted because of spreading lobes, subequal, attached<br />
corky at base; domatia absent. Inflorescences pyra- 0.5-0.6 mm from the base of the tube; filaments 1-<br />
midal, densely to laxly paniculate with opposite or 1.2 mm long, barbate at base, with a minute tuft of<br />
subopposite decussate branches, departing almost white-puberulent hairs (hairs 0.1-0.3 mm long); anperpendicularly<br />
from the rachis; 20-33 x 16-29 cm, thers convex toward center, flattened adaxially, yellateral<br />
branches 8-11 pairs, basal portion of axis not low, banana-shaped, 2-3 x 1-2 mm, dorsifixed near<br />
branched 3-7.5 cm long; rachis terete or decussately the base, base rounded, microscopically (40x) papilcompressed,<br />
rachis and branches glabrous to minute- lose throughout. Pollen exine reticulate. Style expuberulent;<br />
flowers alternate on distal branches; dis- serted, 5-6 mm long, glabrous; style branches ovate,<br />
tal bracts subtending primary branches 0.8-2.5 x 0.8- 0.6-0.7 mm long, stigmatic surface smooth. Capsules<br />
1.2 mm, narrowly triangular to deltoid; bracteoles globular to widely obovate, apex hemispherical, 4-<br />
subtending flowers 0.5-1.3 x ca. 1 mm, deltoid. Flow- 5 x 4-5 mm, dark brown with small lenticels, glaers<br />
sessile to pedicellate, pedicels to 2.5 mm long, brous throughout; disk black. Seeds 0.75-1 x 0.25glabrous<br />
to puberulent; hypanthium turbinate, 1-1.5 0.4 mm wide.<br />
x ca. 1 mm, glabrous. Calyx reduced to an undulate<br />
margin, truncate or with barely distinguishable lobes, Distribution (Fig. 21) and ecology. Evergreen-<br />
0.5 mm x ca. 1 mm; lobes (when evident) shallowly seasonal forests of Pacific slopes of Costa Rica, and<br />
deltoid, to 0.5 mm long. Corolla cupular with spread- Pacific and Atlantic slopes of Panama, at 300-900 m.<br />
ing lobes, 3-4 mm long, color white to cream-white; Flowering specimens were collected in February,<br />
tube narrow-cupular, 1-1.5 mm x ca. 1.5 mm, gla- March, and August. Fruiting specimens were collected<br />
brous outside, glabrous inside, with a ring of white- in February and November.
62 Flora Neotropica<br />
Reproductive biology. The collections of this bark grayish. Leafy branchlets glabrous, dark green,<br />
species, from coastal seasonal forests, suggest that it terete; older branches rugose, grayish; lenticels many,<br />
might have two flowering periods (cf. Lorence in punctiform to linear, 0.5-4 mm long. Stipules nar-<br />
Wilbur & Taylor, 1993): one after the dry season and rowly triangular, acuminate, glabrous outside, glaone<br />
after the rainy season.<br />
brous with basal colleters inside, 10-15(-30) x 25-<br />
30 mm, dark green, readily caducous, leaving a<br />
Specimens examined. COSTA RICA. PUNTARENAS:<br />
white-grayish linear scar. Leaves 17-28(-40)<br />
Hilly slopes above Rio Naranjo, near Londres and Villa<br />
Nueva, 300 m, 84028'W, 09?28'N, 10-12 Feb 1988 (fl),<br />
Burger et al. 12314 (CR, F). SAN Jost: Tarrazu, slopes of<br />
Cerro Nara, between Esquipulas and Villa Nueva, 84?03'W,<br />
09?29'N, 12 Feb 1988 (fl), G6mez-Laurito 11698 (F).<br />
PANAMA. CHIRIQUI: Burica Peninsula, San Bartolo<br />
Limite, 19-21 km W of Puerto Armelles, 400 m, 83?00'W,<br />
08?17'N, 19 Feb 1973 (fr), Busey 464 (MO[3], US), 24 Feb<br />
1973 (fr), Croat 22191 (MO[2], US). VERAGUAS: Valley of<br />
Rio Dos Bocas, 15.6 km NW of Santa Fe, 450-550 m, 31<br />
Aug 1974 (fr), Croat 27709 (MO); Valley of Rio Dos Bocas,<br />
11 km from Escuela Agricola Alto de Piedra, on rd. to<br />
Calovebora, 450 m, 30 Aug 1974 (fl), Croat 27551 (MO),<br />
14 Nov 1974 (fr), Mori & Kallunki 3086 (MO).<br />
<strong>Rustia</strong> costaricensis is unique in the genus in having<br />
minute flowers (3-5 mm long), small usually<br />
globular capsules (4-5 mm long), and inflorescences<br />
openly paniculate with delicate, sometimes scandent,<br />
secondary branches.<br />
Dwyer (1980) described <strong>Rustia</strong> panamensis on the<br />
basis of fruiting material, but without referring to<br />
Stomandra costaricensis. The two taxa are obviously<br />
synonymous and were treated accordingly by Lorence<br />
(cf. Burger & Taylor, 1993).<br />
Two herbarium specimens of <strong>Rustia</strong> costaricensis<br />
(Busey 464, MO) were annotated by Dwyer as "R.<br />
veraguensis," but this epithet remained unpublished.<br />
The type of <strong>Rustia</strong> costaricensis is atypical in having<br />
inflorescences puberulent throughout, with<br />
branches slightly geniculate and widely zig-zagging,<br />
while the other collections have inflorescences openly<br />
paniculate, glabrous, with secondary branches thin and<br />
delicate and not geniculate. In combination this set of<br />
characters suggests that these plants deserve formal<br />
recognition, but more collections are needed.<br />
2. <strong>Rustia</strong> occidentalis (Bentham) Hemsley, Biol.<br />
Centr.-Amer., Bot. 2: 14. 1881. Exostemma (sect.<br />
Pseudostemma) occidentale Bentham, Bot. Voy.<br />
Sulph. 104. 1844. Type. Colombia. Nariiio: Isla<br />
Gorgona, 1841 (fl), Hinds 354 (lectotype, K, selected<br />
by Delprete, 1999b; isolectotype, BM).<br />
Figs. 3A, 4A, 5E, 13C, 22, 23<br />
Shrub 2-5 m tall, exceptionally a tree to 8 m tall,<br />
to 30 cm dbh, much-branched, rarely single-stemmed;<br />
x 4-12<br />
cm, L/W 2.5:1 to 4:1; narrowly elliptic, lanceolate to<br />
oblanceolate, cuneate to acute (BA = 18-25?) at base,<br />
acute-caudate at apex, tapering to a long acuminate<br />
apex, the acumen often falcate, 1.5-3 cm long; dark<br />
green above, pale green below, foliaceous; drying<br />
grayish olive-green, stiff-chartaceous; glabrous above<br />
and below; pellucid punctate; primary and secondary<br />
veins glabrous, slightly depressed above, prominent<br />
below, secondary veins 11-16 each side; tertiary veins<br />
starting subparallel and reticulate in the center; petioles<br />
20-40 mm long, 1-2.5 mm thick, adaxially concave<br />
to flattened; pulvinus evident in young leaves,<br />
becoming corky in mature leaves; domatia absent.<br />
Inflorescences reduced-paniculate to racemoid, opposite<br />
to subopposite decussate lateral branches; 4-<br />
1(-15) x 2-10(-13) cm, lateral branches to 4 pairs,<br />
basal portion of axis not branched to 3 cm long; rachis<br />
decussately compressed to terete, rachis and<br />
branches glabrous; flowers alternate on distal branches<br />
or in sparse cymules; distal bracts 1-2 x ca. 1 mm,<br />
deltoid; bracteoles subtending flowers ca. 1 x 0.7-1<br />
mm, deltoid, glabrous. Flowers pedicellate, pedicels<br />
4-15 mm long, glabrous; hypanthium narrowly turbinate,<br />
3-5 x 2-3 mm, glabrous; flower buds clavate.<br />
Calyx reduced to an undulate margin, to 3 mm long,<br />
sometimes ciliolate. Corolla tubular, medially faintly<br />
constricted with perpendicularly spreading lobes, 1.1-<br />
1.4 cm long, purple to flesh-red outside, white to pearlwhite<br />
inside, carnose to coriaceous when fresh; tube<br />
subcylindrical, 7-9 mm long, 2.5-3.5 mm wide at base<br />
and 4-5 mm wide at the orifice, glabrous outside, glabrous<br />
inside, without ring of hairs; lobes 5 (rarely 4),<br />
/2 of corolla length, 5-6 x 1.7-2 mm, triangular, glabrous<br />
outside and inside. Stamens 5, exserted because<br />
of spreading lobes, subequal, attached 4-5 mm from<br />
the base of the tube; filaments ca. 2 mm long, glabrous,<br />
distally terete, basally flattened-adnate to tube,<br />
ca. 0.8 mm at base; anthers convex toward the center,<br />
yellow, banana-shaped, 5-7 x 1.8-2 mm, dorsifixed<br />
near the base, base rounded, smooth throughout. Pollen<br />
exine reticulate. Style exserted, 10-14 mm long,<br />
glabrous; style branches ovate, 0.8-1.2 x ca. 0.9 mm,<br />
stigmatic surface smooth. Capsules obovoid to widely<br />
obovoid (rarely globular), obtuse to rounded at base,<br />
apex hemispherical, 7-12 x 7-9 mm, dark brown to<br />
black at maturity, with minute lenticels, glabrous<br />
throughout; disk septicidal dehiscence present of old<br />
capsules. Seeds 1.06-1.26 x 0.67-0.83 mm.
Systematic Treatment of the Genera Studied 63<br />
?<br />
l: 6mm .":, .<br />
=*' ??, ,I ~ ~ ~<br />
?"?i?r?~<br />
FIG 22 Rutiaoccdenali (rofrsmaeiladDlet63,TE.A.Hbtfinoesnewthaue<br />
FIG. 22. <strong>Rustia</strong> occidentalis (from fresh material and Delprete 6329, TEX). A. Habit of inflorescence with mature<br />
leaves. B. Mature capsule. C. Flower bud. D. Longitudinal section of flower bud. E. Flower in later stage of anthesis (note<br />
style protruded and receptive). F. Detail of receptive style branches. G. Stamen, dorsal view. H. Stamen, lateral view.<br />
rr~:?'<br />
D,-
64<br />
Flora Neotropica<br />
thesis. C. Flower buzz-pollinated by Euglossa<br />
7z<br />
sp. D. Flower buzz-pollinated by Melipona sp.<br />
I~. 3._.: ?:<br />
.......<br />
..<br />
.. F<br />
:"'<br />
... . .'.. '..<br />
.... ........<br />
?:'?? :~~ :'. .:<br />
F:<br />
FIG. 23. <strong>Rustia</strong> occidentalis. A. Coastal environment typical of R. occidentalis (Choc6, Colombia). B. Flower in an-<br />
Distribution (see Fig. 29) and ecology. Mostly in<br />
swampy coastal areas (Fig. 23A), near the coast but<br />
not influenced by salt water, in proximity of streams<br />
in lowland costal forests, and in coastal areas of some<br />
islands, 0-300 m, in Nicaragua, Guatemala, Costa<br />
Rica, Panama, Colombia, and N Ecuador. Flowering<br />
specimens were collected in all months of the year<br />
except January. Fruiting specimens were collected in<br />
February, March, April, July, August, and December.<br />
Pollination biology. I observed <strong>Rustia</strong> occiden-<br />
talis in coastal swampy sites of Choc6, Colombia,<br />
where in some areas this species is in almost pure<br />
stands. The flowers of this species are fleshy and<br />
purple-red to flesh-red, and never pendulous. <strong>Rustia</strong><br />
occidentalis is mainly visited by bees (Euglossa sp.<br />
and Melipona sp.; see Fig. 23C,D), which perform a<br />
rather characteristic buzz-pollination. During late-<br />
morning hours, if one listens carefully in a large popu-<br />
lation of this species, it is possible to hear the scat-<br />
tered buzzing of bees visiting the flowers. This is the<br />
first species of the entire <strong>Rubiaceae</strong> that has been<br />
observed and reported to be buzz-pollinated.<br />
Representative specimens examined. GUATEMALA.<br />
IZABAL: San Juan de Nicaragua, 1841, Friedrichsthal s.n. (F)<br />
[dubious label data-specimen probably collected in Guatemala].<br />
*
Systematic Treatment of the Genera Studied 65<br />
NICARAGUA. ZELAYA: Area of Bahia de Bluefields,<br />
Rio Escondido, near Rio Yanten, 0-30 m, 2 Apr 1949 (fr),<br />
Molina 2085 (F).<br />
COSTA RICA. Cocos ISLAND: Valley behind Bahia<br />
Iglesias, 9-11 Apr 1979 (fr), Foster 4133 (CAS, F, US), Mar<br />
1970 (fl), G6mez 3307 (F); Wafer Bay, 13 Apr 1965 (fl),<br />
Jimenez 3173 (F, NY); along brook flowing into Wafer Bay,<br />
18 Apr 1930 (fl), Svenson 331 (F, UC), 3-13 Sep (fl), Stewart<br />
315 (CAS, GH, US), 5 Dec 1959 (fl), Klawe 1492 (US).<br />
LIM6N: Near edge of waterway, 3 km N of Moin, 5 m, 13<br />
Feb 1965 (fl), Lent 359 (F); Cafio de Moin, 13 Feb 1965 (fl),<br />
Jimenez 2898 (F). PUNTARENAS: Peninsula de Osa, Parque<br />
Nacional Corcovado, Estaci6n San Pedrillo, 10-200 m, 23<br />
Jul 1991 (fr), Delprete et al. 5267 (CR, MO, TEX).<br />
PANAMA. BOCAS DEL TORO: Old Bank Island, 17 Feb<br />
1941 (fl-fr), Wedel 2114 (F, MO); Laguna of Chiriqui, Nov-<br />
Dec 1885 (fl), Hart 109 (US). COL6N: Corr. Crist6bal, near<br />
mouth of Rio Chagres, 11 Mar 1935 (fr), Allen 860 (BR, G,<br />
MO, P); Corr. Crist6bal, Chagres, Feb-Mar 1850 (fl-fr),<br />
Fendler 289 (F, K[2], MO[3], US); Corr. Crist6bal, Ft.<br />
Sherman, 1.5 km SW of Ft. San Lorenzo, near sea level, 20<br />
Sep 1979 (fl), Antonio 1935 (MO); Corr. Crist6bal, W of<br />
Limon Bay, Gatun lake, 30 Mar 1956 (fr), Johnston 1745<br />
(MO); Miguel de la Borda, 20 Apr 1970 (fr), Croat 9828<br />
(F, MO, NY); along Cafio Rey, near Cocle del Norte, 18 Aug<br />
1972 (fl), Dressier 4207 (MO); along Rio Culebra, above<br />
Santa Isabel, 10 Aug 1911 (fl), Pittier 4166 (NY, US).<br />
DARIEN: Puerto St. Dorothea, 21 Jul 1962 (fl), Dwyer 2290<br />
(F[2], MO, US); Punta Guayabo Grande, 0-50 m, 20 Apr<br />
1980 (fr), Antonio & Hahn 4240 (MO); vic. of Pifias, 2 Mar<br />
1967 (fr), Duke 10567 (ECON), 10605 (MO).<br />
COLOMBIA. CHoc6: Upstream of Pizzarro, Rio<br />
Baud6, 2 Aug 1973 (fr), White & Warner 82 (GH, MO); near<br />
Baud6, right bank of Rio Baud6, ca. 5 m, 2 Feb 1967 (fr),<br />
Fuchs & Zanella 21703 (US); Mecana, N of Bahia Solano,<br />
Rio San Juan, 77?21'W, 06?16'N, 31 Dec 1983 (fr), Juncosa<br />
1589 (MO); Rio San Juan, 80 m, May 1853 (fl), Triana 1784<br />
(3262.3) (BR, F); Corr. Pangui, trail toward swamps, 17 Dec<br />
1992 (fl-fr), Delprete 6319 (CHOCO, COL, F, MO, NY,<br />
TEX), 6329 (NY [4]); Utria bay, trail toward mtns., 18 Dec<br />
1992 (fl-fr), Delprete 6330 (CHOCO, COL, F, MO, NY,<br />
TEX); Mun. Nuqui, Corr. Termales, between quebrada Piedra<br />
Piedra and Rio Terco, 0-25 m, 5 Sep 1994 (fr) Acevedo-<br />
Rodriguez et al. 6774 (US), 9 Sep 1994 (fr) Acevedo-<br />
Rodriguez et al. 6900 (US). NARIFIO: Isla de Gallos, near<br />
Tumaco, Feb 1943 (fr), Dryander 2583 (US); Gorgona Island,<br />
NE side, 5-20 m, 17 Mar 1975 (fl-fr), Cabrera 3264<br />
(MO, NY, QCA); Gorgona Island, 18 Oct 1924 (fl), St.<br />
George Expedition 624 (US). VALLE: La Cuarantena, Buenaventura<br />
Bay, 9 Jun 1944 (fl), Killip & Cuatrecasas 38970<br />
(US); Pacific Coast, Rio Naya, Puerto Merizalde, 1 Mar 1943<br />
(fl), Cuatrecasas 14317 (US); Malaga Bay, mouth of Quebrada<br />
Sierpe, 17 Feb 1983 (fl-fr), Gentry et al. 40418A (MO).<br />
ECUADOR. ESMERALDAS: Estero Batia, 22 Jul 1967 (fr),<br />
Jdtiva & Epling 771 (NY[3], S[2], US[3], VEN); near<br />
Borb6n, 1967 (fr), Jdtiva & Epling 2201 (NY[2], S[2],<br />
US[2]); Limones-Borb6n, 5 km before Borb6n, 10 m,<br />
79?00'W, 01?07'N, 5 Sep 1980 (fl-fr), Holm-Nielsen et al.<br />
26039 (AAU); Borb6n, between Punto de Piedra and<br />
Empalme, 78?58'W, 0104'N, 11 Feb 1993(fl-fr), Jaramillo<br />
14990 (QCA); San Lorenzo, Rio San Antonio, Finca El Horizonte,<br />
12 Aug 1967 (fl), Sparre 18300 (S); San Lorenzo, Rio<br />
San Antonio, El Chorro, 22 Aug 1967 (fl), Sparre 18340(S).<br />
Bentham (1844) first described this species as<br />
Exostemma (sect. Pseudostemma) occidentale from<br />
specimens collected from the Gorgona Island, near the<br />
Pacific coast of Colombia. In his protologue Bentham<br />
did not refer to specific collections, but nevertheless<br />
one herbarium specimen preserved at Kew bears the<br />
label: "Exostemma occidentale Benth., Voy. Sulph.<br />
104, a large shrub, Isle Gorgona, Hinds 1841." This<br />
is probably one of the specimens that Bentham saw<br />
when he described the species, and I selected this as<br />
lectotype (Delprete, 1999b). Hemsley (1881) later<br />
transferred it to <strong>Rustia</strong>, citing three specimens: "Gua-<br />
temala (Friedrichstal); Panama (Fendler 289); Co-<br />
lombia Hb. Kew." The last specimen is probably the<br />
one seen by Bentham when he described this species.<br />
<strong>Rustia</strong> occidentalis is the most widespread species<br />
of the genus, usually encountered as much-branched<br />
shrubs 2-4 m tall, often with roots submerged in wa-<br />
ter. This species is easily recognized by its carnose<br />
flesh-red to purple flowers, deep-yellow convex ("ba-<br />
nana-like") anthers, and clavate flower buds.<br />
<strong>Rustia</strong> occidentalis is most similar to R. dressleri<br />
(from Panama), but the latter has white flowers with<br />
yellow lobes, young leafy branchlets semi-succulent,<br />
and leaf blades 30-45 cm long with 24-30 secondary<br />
veins (exceptionally to 30 cm long with 11-16 sec-<br />
ondary veins in R. occidentalis).<br />
3. <strong>Rustia</strong> dressleri Delprete, Novon 5(2): 137, fig.<br />
3. 1995. Type. Panama. Panama: El Llano-Carti<br />
Hwy., 8-10 km N of El Llano, 31 Aug 1974 (fl),<br />
Dressier 4703 (holotype, MO). Fig. 24<br />
Shrub to tree to 7 m tall. Leafy branchlets 7-9 mm<br />
wide, semi-succulent, grayish, glabrous, terete; older<br />
branches rugose, pale brown-grayish; lenticels not<br />
found. Stipules not seen (readily caducous), leaving<br />
a linear scar ca. 1 mm wide, of the same color as the<br />
stem. Leaves 30-54 x 9.5-13 cm, L/W 3:1 to 4:1;<br />
narrowly elliptic to oblanceolate, cuneate (BA = 20-<br />
25?) at base, acute or acuminate at apex, the acumen<br />
ca. 1 cm long; shiny grass-green above, pale grayish<br />
green below, subcoriaceous; drying olive-green, semi-<br />
coriaceous; glabrous above and below; evidently pel-<br />
lucid punctate; primary and secondary veins glabrous,<br />
prominent below, secondary veins 24-30 each side,<br />
subparallel; tertiary veins starting subparallel and<br />
openly reticulate in the center, very evident above,
66 Flora Neotropica<br />
Sn~<br />
.r ," AS ^p .\ . .. . . . ,'<br />
'<br />
',"<br />
A<br />
FIG. 24. <strong>Rustia</strong> dressleri (A-D from Dressier 4703, MO, holotype; E, F from Herrera 1663, TEX). A. Habit of inflorescence<br />
with mature leaf. B. Abaxial view of blade showing pellucid punctation. C. Detail of open corolla from flower<br />
bud, showing callous base and descending pilose tufts. D. Flower bud (clavate). E. Detail of corolla lobe tip, abaxial view.<br />
F. Stamen with open anther. (Reproduced with permission from Novon 5, fig. 3. 1995.)
Systematic Treatment of the Genera Studied 67<br />
faintly evident below; petioles 28-35 mm long, ca. 3 mm Jul 1994 (fl), Herrera 1663 (CR, GB, MEXU, MO, PMA,<br />
thick, adaxially concave to flattened, glabrous; pul- STRI, TEX).<br />
vinus not evident but thickened, becoming corky at<br />
base; domatia absent or a tuft of few hairs. Inflores- <strong>Rustia</strong> dressleri is similar to R. occidentalis, from<br />
cences reduced panicles with opposite decussate which it differs in having white flowers with brownbranches;<br />
(12-)18-30 cm long, basal branches 6.5- ish yellow lobes (flesh-red in R. occidentalis), fila-<br />
13 cm long, lateral branches 4-7 pairs, basal ments with a basal tuft of retrorse<br />
portion<br />
white-pilose hairs<br />
of axis not branched 0.9-2 cm long; rachis decussately (puberulent in R. occidentalis), larger inflorescences<br />
compressed, terete at base, rachis and branches semi- (to 11 cm long in R. occidentalis), young branchlets<br />
succulent,<br />
thick and<br />
glabrous throughout, moderately puberu- semi-succulent, the whole plant glabrous,<br />
lent at nodes; flowers in opposite cymules on rachis and leaf blades 30-45 cm long (exceptionally to 30<br />
and lateral branches; distal bracts subtending lateral cm in R. occidentalis) with 24-30 secondary veins<br />
branches 2.5-3 x 3-4 mm, deltoid, glabrous; along each side (11-16 in R. occidentalis).<br />
bracteoles subtending flowers 1-1.5 x 1-2 mm, deltoid,<br />
glabrous. Flowers subsessile to pedicellate, 4. <strong>Rustia</strong> alba Delprete, Novon 5(2): 133, fig. 1. 1995.<br />
pedicels 0.5-4 mm long, glabrous; hypanthium<br />
obconical, 2.5-3.5 x Type. Ecuador. Carchi: Environs of Maldonado,<br />
2-2.5 mm, glabrous; flower buds<br />
wet montane forest, 1450-1650 m, 31 May 1978<br />
clavate, 2.5 mm at base and 3.5-4 mm at bulge. Ca-<br />
(fl), Madison, Plowmann & Besse 4808 (holotype,<br />
lyx extremely reduced, with a wavy margin or with<br />
AAU; isotypes, F, QCA, SEL-n.v., US).<br />
barely distinguishable lobes, 0.5 mm long, sometimes<br />
Figs. 3B, 6F, 13D, 25A-E, 26A,B<br />
ciliolate. Corolla tubular with spreading lobes, 1.2-<br />
1.4 cm long, white with yellow lobes, semi-fleshy<br />
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<br />
3.5-4 mm wide at the orifice, glabrous outside, branched tree, with straight trunk, with small butglabrous<br />
inside, with a ring of white-pilose descend- tresses; bark grayish. Leafy branchlets glabrous, tering<br />
to erect hairs, 5-6 mm from the base, at the same ete; older branches rugose, grayish to pale brown;<br />
point of the filament attachments at base of lobes; lenticels sparse. Stipules narrowly triangular, glalobes<br />
5, ca. /2 of corolla length, 5-6 mm x ca. 3 mm, brous outside, glabrous and with few colleters inside,<br />
triangular, semi-fleshy, glabrous outside, white-stri- 28-37 x 7-9 mm, reddish green, readily caducous,<br />
gose at base, margins microscopically (40x) papillose leaving a linear scar, ca. 0.5 mm wide, of the same<br />
inside. Stamens 5, partially exserted because of nar- color of the stem. Leaves 25-31 x 13-16 cm, L/W<br />
rowly spreading lobes, subequal, attached 5-6 mm 1.6:1 to 1.7:1; broadly elliptic to broadly obovate,<br />
from the base of the tube; filaments ca. 2 mm long, acute to rounded (BA<br />
distally terete, basally flattened-adnate to tube, barbate<br />
at base, with a tuft ofretrorse white-pilose hairs (0.1-<br />
0.2 mm long); anthers convex toward the center, yellow,<br />
banana-shaped, 4-5 x 1.8-2 mm, dorsifixed near<br />
the base, bases of thecae rounded, subequal, microscopically<br />
(40x) papillose-echinate throughout. Pollen<br />
exine reticulate. Style exserted, 13-16 mm long,<br />
glabrous; style branches ovate to narrowly ovate, ca.<br />
1.5 mm long, stigmatic surface smooth. Immature<br />
capsules green and semi-fleshy when fresh; mature<br />
capsules not seen. Seeds not seen.<br />
Distribution (Fig. 21) and ecology. Known only<br />
from three collections made in the provinces of Panama<br />
and San Bias (Panama). The specimens were collected<br />
in flowering stage in July, August, and September.<br />
Additional specimens examined. PANAMA. PANAMA:<br />
El Llano-Carti Hwy., 10-12 kmN of El Llano, 12 Sep 1974<br />
(fl), Dressier 4749 (MO). SAN BLAS: Rio Playon, Chico,<br />
Campamento Neba Dummat, 09 15'N, 78?15'W, 50 m, 10<br />
= 30-45?) at base, obtuse and<br />
short-acuminate at apex; the acumen to 1 cm long;<br />
blade pale green above, yellowish green below, semicoriaceous;<br />
drying pale green, semi-leathery; glabrous<br />
above and below; pellucid punctate; primary and secondary<br />
veins glabrous, prominent below; secondary<br />
veins 16-21 each side; tertiary veins starting subparallel<br />
and openly reticulate in the center, faintly<br />
evident above and below; petioles 2-3 cm long, 3-<br />
4.5 mm thick, adaxially narrow-concave to flattened,<br />
glabrous, basally thickened but not pulvinated; domatia<br />
absent. Inflorescences laxly paniculate, pyramidal,<br />
lateral branches decussate; 30-46 cm long,<br />
basal branches 22-30 cm long, lateral branches 6-8<br />
pairs, basal portion of axis not branched 7-11 cm long;<br />
rachis terete, rachis and branches glabrous to minutely<br />
hirtellous; distal bracts 4-5 x 4-5 mm, deltoid;<br />
bracteoles subtending flowers 2-3 x 2-3 mm, deltoid,<br />
glabrous. Flowers pedicellate, pedicels 4-6 mm long,<br />
glabrous to minutely puberulent; hypanthium narrowly<br />
obconical, 4-6 x 2-3 mm, glabrous; flower buds<br />
cylindrical, pointed at apex. Calyx cupular with small
68<br />
..............p.. U<br />
E(;I F<br />
'<br />
.<br />
.....1 .<br />
Flora Neotropica<br />
view of blade showing pellucid punctation. C. Flower bud (acute at apex) showing cupular calyx. D. Stamen. E. Detail of<br />
anther external surface. F. Receptive style. (Reproduced with permission from Novon 5, fig. 1. 1995.)<br />
..<br />
i
Systematic Treatment of the Genera Studied 69<br />
4 ,~~~~~~~~6<br />
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tct<br />
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r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
~? ' ::!~::'~.:".:';!:.d :~. .......~,.,:.'":~:~Z~ 2..:,: g.<br />
.:.,..<br />
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t?3~~~~~~~~~~~~~~~~~~~~~~~?<br />
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?_: ~e~ ,e '??. i~~~~~~~~~~~~~~~~~~~~~~~~~~....<br />
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i![[l-;<br />
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FIG. 26. <strong>Rustia</strong> alba. A. Mature tree, ca. 15 m tall, within dense vegetation (arrows indicate tree canopy). B. Inflores-<br />
cence with immature fruits.<br />
lobes, membranous, glabrous, greenish white, 1.5-3<br />
mm long; lobes 5, 1-2.5 x 1.5-2 mm, ciliolate. Corolla<br />
tubular with a basal constriction and reversed<br />
lobes, 1.5-1.9 cm long, white, semi-fleshy when fresh;<br />
tube subcylindrical with basal constriction, 5-6 x 2 mm,<br />
3.5-4 mm wide at the orifice; glabrous, minutely verrucate<br />
outside; microscopically (40x) granulate inside,<br />
without a ring of hairs; lobes 5, /3 of corolla length,<br />
9-11 x ca. 3 mm, narrowly triangular, reversed, glabrous,<br />
minutely verrucose outside, densely ascending<br />
short-strigose at base, minutely papillose throughout,<br />
Pollen exine densely foveolate (non-echinate). Style<br />
exserted, 16-20 mm long, glabrous; style branches<br />
ovate, 3-3.5 x 1-1.3 mm, stigmatic surface papillose.<br />
Capsules obovoid, apex obtusely hemispherical, 8-<br />
12 x 7-9 mm, without lenticels, glabrous throughout,<br />
dark-brown; dehiscing loculicidally, disk septicidal<br />
dehiscence absent even in old capsules. Seeds 2.25-<br />
2.9 x 0.65-0.75 mm.<br />
Distribution (see Fig. 29) and ecology. Known<br />
only from the cloud forest of Carchi province, NW<br />
microscopically (40x) papillose at margins inside. Ecuador, at 1500-1700 m. An extremely rare and<br />
Stamens 5, fully exserted, subequal, attached 4-5 mm threatened species, because the area where it occurs<br />
from the base of the tube; filaments ca. 3 mm is<br />
long, highly disturbed and primarily dedicated to cow<br />
flattened, widened at base, with a tuft of ascending pastures. Flowering specimens were collected in Febstrigose<br />
hairs (0.3-0.5 mm long); anthers cylindrical, ruary, May, and August.<br />
subsagittate at base, rectilinear to slightly curved (not Specimens examined. ECUADOR. CARCHI: El<br />
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<br />
the thecae subequal at base, papillose throughout. 1992 (fl-fr), Palacios et al. 9663 (MO, QCNE, TEX); Mun.
70 Flora Neotropica<br />
Maldonado, rd. Maldonado-Chical, 1500-1700 m, 14 Aug<br />
1994 (fl-fr), Delprete & Verduga 6414 (AAU, MO, NY,<br />
QCA, QCNE, TEX, UPS), 6419 (NY, TEX), 6420 (NY,<br />
QCA, TEX).<br />
<strong>Rustia</strong> alba is similar to R. formosa but differs in<br />
having a larger inflorescence (exceptionally to 36 cm<br />
long in R. formosa); calyx cupular, more expanded,<br />
membranous, and greenish white; smaller corollas;<br />
and shorter styles with longer, membranous style<br />
branches (ca. 1 mm long in R. formosa and ca. 3 mm<br />
long in R. alba). <strong>Rustia</strong> alba and R. formosa are trees<br />
commonly reaching 12 m tall and with coriaceous to<br />
semi-coriaceous leaves, but R. formosa is encountered<br />
only in southern Brazil.<br />
<strong>Rustia</strong> alba differs from R. bilsana in having larger<br />
flowers, inflorescences, and capsules, expanded mem-<br />
branous (vs. much-reduced) calices, widely obovate<br />
(vs. oblanceolate) leaves, and white (vs. purple) co-<br />
rollas (see discussion under R. bilsana).<br />
The type material of <strong>Rustia</strong> alba was collected<br />
from an individual with several stump sprouts ca. 6 m<br />
tall. Near the type locality, among a few young trees<br />
(see Fig. 26A) I encountered one living stump, with<br />
sprouts 4-5 m long, that was ca. 40 cm dbh.<br />
5. <strong>Rustia</strong> bilsana Delprete, Novon 8: 234, figs. 1, 2.<br />
1998. Type. Ecuador. Esmeraldas: Quinind6, Bilsa<br />
Biological Station, Mache Mtns., 35 km W of Qui-<br />
ninde, 5 km W of Santa Isabel, along Cube River,<br />
premontane wet forest, 00?21'N, 79044'W, 500 m,<br />
2 Oct 1996, Clark 2979 (holotype, QCNE; iso-<br />
types, AAU, CAS, COL, DAV, F, G, GB, GH, K,<br />
MO, NY[2], P, QCA, QCNE, SEL, TEX, US,<br />
VEN). Figs. 27A-E, 28A-D<br />
Tree to 19 m tall, to 47.5 cm diam. Leafy branchlets<br />
glabrous. Stipules narrowly triangular, glabrous<br />
outside, glabrous and with a few colleters inside, 23-<br />
27 x 8-10 mm, readily caducous, leaving a linear scar<br />
ca. 0.5 mm wide. Leaves 33-55 x 13-21 cm, oblanceolate,<br />
L/W 2.5, acute-decurrent at base, rounded and<br />
short-acuminate at apex, thick-chartaceous, glabrous<br />
above and below; secondary veins 15-18 pairs, faintly<br />
evident above and below; petioles 2-2.5 cm long;<br />
domatia absent. Inflorescences laxly paniculate, pyramidal,<br />
16-26 cm long, basal branches 9-12 cm long,<br />
lateral branches 4-5 pairs. Flower pedicillate, pedicels<br />
1-2 mm long, glabrous; hypanthium obconical, 3-5<br />
x ca. 3 mm, glabrous; flower buds cylindrical, pointed<br />
at apex. Calyx reduced to a wavy margin with barely<br />
distinguishable lobes, 0.5-1 mm long. Corolla tubu-<br />
lar with a basal constriction and reversed lobes, 0.8-<br />
0.9 cm long, purple, semi-fleshy; tube 3.5-4 mm long,<br />
glabrous outside and inside; lobes 5, 2/3 of corolla<br />
length, 5-5.5 mm long, narrowly triangular, reflexed,<br />
glabrous outside, microscopically (40x) papillose at<br />
margins inside. Stamens 5, fully exserted, subequal,<br />
attached ca. 4 mm from the base of the tube; filaments<br />
2 mm long, flattened, widened at base, with a basal<br />
tuft of antrorse, retrorse strigose hairs; anthers convex<br />
toward the center and rounded at base, 4.1-4.4 x<br />
1.2-1.3 mm, dorsifixed near the base, the thecae equal<br />
at base. Pollen not seen. Style exserted, 9-11 mm<br />
long, glabrous; style branches ca. 1.5 mm long, stigmatic<br />
surface papillose. Capsules widely obovoid,<br />
apex obtusely hemispherical, 7-8 x 6-6.5 mm, without<br />
lenticels, glabrous throughout, dark brown; disk<br />
septicidal dehiscence absent even in old capsules.<br />
Seeds 0.6-1.2 x 0.3-0.5 mm.<br />
Distribution (Fig. 29) and ecology. Known only<br />
from two collections from the premontane wet forest<br />
of Esmeralda province, in the Mache Mountains, Bilsa<br />
Biological Station, in N Ecuador, at 400-600 m. The<br />
specimens have been collected in late flowering-early<br />
fruiting stage in October.<br />
Specimens examined. ECUADOR. ESMERALDAS: Quininde,<br />
Bilsa Biological Station, Mache Mtns., 35 km W of<br />
Quininde, 5 km W of Santa Isabel, premontane wet forest,<br />
00?21'N, 79?44'W, 400-600 m, 3 Oct 1994, Clark, Bass,<br />
Pitman 139 (MO, QCA, QCNE, TEX).<br />
<strong>Rustia</strong> bilsana is similar to R. alba, from the cloud<br />
forests of northwestern Ecuador, in having floral buds<br />
pointed at apex and widely obovoid capsules. The two<br />
differ in the former having leaves oblanceolate (vs.<br />
widely obovate), acute-decurrent [BA = 20-25?] at base<br />
(vs. acute-rounded [30-45?]); inflorescence ca. 16 cm<br />
long (vs. 30-46 cm long); calyx reduced, with a wavy<br />
margin (vs. cupular and small lobes); corolla purple<br />
and 8-9 mm long (vs. white and 15-19 mm long);<br />
corolla lobes 5-5.5 mm long (vs. 9-11 mm long),<br />
glabrous inside (vs. densely ascending short-strigose<br />
at base); anthers half-exserted (vs. fully exserted and<br />
pendulous); style 9-11 mm long (vs. 16-20 mm long),<br />
style branches 1.5 mm long (vs. 3-3.5 mm long); capsules<br />
7-8 mm long (vs. 8-12 mm long); and seeds 0.6-<br />
1.2 x 0.3-0.5 mm (vs. 2.25-2.9 x 0.65-0.95 mm).<br />
This species differs from <strong>Rustia</strong> occidentalis in<br />
being tall trees growing at medium elevations (vs.<br />
shrubs growing in swampy or wet areas), having<br />
smaller flowers, flower buds pointed (vs. clavate),<br />
bigger and more branched inflorescences, and smaller<br />
capsules.
Systematic Treatment of the Genera Studied 71<br />
FIG. 27. <strong>Rustia</strong> bilsana (Clark et al. 139, TEX, holotype). A. Habit of inflorescence with mature leaf. B. Flower bud.<br />
C. Open flower. D. Stamen, dorso-lateral<br />
view. E. Receptive style. (Reproduced with permission from Novon 8, fig. 1. 1998.)<br />
.'<br />
.
72 Flora Neotropica<br />
? N<br />
i A: ....<br />
stage of anthesis (top) and one in later stage (bottom).<br />
6. <strong>Rustia</strong> rubra Standley ex D. Simpson, Phytologia<br />
33: 4. 1976. Type. Ecuador. Pastaza: in clearing<br />
at La Esperanza, Rio Chingual, near jct. with Rio<br />
Las Ollas, ca. 1700 m, 7 Aug 1943 (fl), Steere<br />
8021 (holotype, F; isotypes, US, NY).<br />
Figs. 5F, 13G-H, 30, 31<br />
Shrub to tree 15-20 m tall, to 30 cm dbh, much<br />
branched tree, with straight trunk, sometimes with<br />
small buttresses; bark grayish. Leafy branchlets gla-<br />
brous, terete (rarely obtusely compressed); older<br />
branches rugulose, grayish to pale brown; lenticels<br />
sparse, elongate to linear. Stipules narrowly triangu-
Systematic Treatment of the Genera Studied 73<br />
lar, glabrous outside, glabrous and with few colleters<br />
inside, 20-35 x 4-6 mm, reddish green, readily caducous,<br />
leaving a linear scar ca. 0.5 mm wide, of the<br />
same color of the stem. Leaves 22-37(-46) x 8.5-<br />
14(-17.5) cm, L/W 2:1 to 3:1, elliptic, oblanceolate<br />
to obovate, acute to rounded (BA = 23-32 mm long, glabrousstyle branches ovate, 0.5-<br />
0.7 mm long, stigmatic surface smooth; immature<br />
fruits red and semi-carose. Capsules obovoid to<br />
oblong-elliptic, apex subtruncate to obtusely hemispherical,<br />
(9-)11-20<br />
25-45?), acute to<br />
obtuse-acuminate at apex; the acumen (when present)<br />
deltoid, to 1 cm long; blade dark green above, yellowish<br />
below, chartaceous to semi-coriaceous; drying<br />
olive-green or pale brown, stiff-chartaceous to<br />
semi-leathery; glabrous above, glabrous to minutely<br />
puberulent below; pellucid punctate; primary and secondary<br />
veins glabrous to minutely puberulent, prominent<br />
below; secondary veins 14-20 each side; tertiary<br />
veins starting subparallel and openly reticulate in the<br />
center, faintly evident above, very evident below;<br />
petioles 2-5.5(-7) cm long, 2-3(-4) mm thick, terete<br />
to adaxially flattened, glabrous; basally thickened but<br />
not pulvinate; domatia absent. Inflorescences densely<br />
paniculate, pyramidal, twice to thrice compound, lateral<br />
branches decussate, often secundiflorous, sometimes<br />
racemoid; rachis, branches, pedicels, calyx,<br />
corollas, and juvenile fruits deep red; 20-90 x 12-50<br />
cm, lateral branches 4-12 pairs, basal portion of axis<br />
not branched 3-9 cm long; rachis terete to decussately<br />
compressed, rachis and branches glabrous to minutely<br />
puberulent; flowers alternate on distal branches; distal<br />
bracts subtending lateral branches 1-4 x 2-3 mm,<br />
deltoid; bracteoles subtending flowers 0.5-0.8 x 0.5-<br />
0.8 mm, deltoid, glabrous. Flowers pedicellate,<br />
pedicels 3-15 mm long, glabrous to minutely puberulent;<br />
hypanthium narrowly obconical, 3-4 x 1.5-2 mm,<br />
glabrous to puberulent; flower buds faintly clavate.<br />
Calyx reduced to a wavy margin or with minute lobes,<br />
ca. 0.5 mm long; lobes 5 (when distinguishable), 0.1-<br />
0.5 x 1.5-2 mm. Corolla tubular, 1.8-2.2 cm long,<br />
deep red with green to yellow-green lobes outside,<br />
deep red inside, semi-fleshy when fresh; tube subcylindrical,<br />
striate, 1.6-1.9 cm long, 2-2.5 mm wide at<br />
base and 2-3 mm wide at the orifice, glabrous outside;<br />
ascending long-strigose at base inside, without<br />
a ring of hairs, the remaining medial and superior<br />
zones glabrous; lobes 5, ca. 1/10 of corolla length, 2-<br />
3.5 x 1.5-2 mm, deltoid, glabrous outside, glabrous,<br />
minutely papillose (40x) at margins inside. Stamens<br />
5, included or partially exserted, subequal, attached<br />
5-6 mm from the base of the tube; filaments 10-15<br />
mm long, sparsely pilose at base; anthers subcylindrical,<br />
sagittate to caudate at base, rectilinear (never<br />
convex), (4-)5-8 mm long, 1-2.5 mm wide at apex,<br />
dorsifixed near the base, base sagittate to caudate, the<br />
thecae sometimes very unequal (3 and 7 mm long on<br />
same filament), smooth throughout. Pollen exine<br />
densely foveolate, minutely echinate. Style exserted,<br />
x 5-8 mm, black, without (rarely<br />
with few) lenticels, glabrous to puberulent below the<br />
disk; disk black, glabrous; disk septicidal dehiscence<br />
absent even in old capsules. Seeds 1.16-1.5 x 0.33-<br />
0.53 mm wide.<br />
Distribution (Fig. 29) and ecology. Primary and<br />
secondary rain forests and cloud forests, and relative<br />
remnants, in low to medium elevations, 450-1900 m,<br />
of Amazonian Ecuador and Peru. Flowering specimens<br />
were collected in February, March, June, and<br />
July. Fruiting specimens were collected in January,<br />
May, and June.<br />
Specimens examined. ECUADOR. NAPO: Rd. Baeza-<br />
Tena, S slope of Cordillera de Guacamayos, 1800 m, 9 Feb<br />
1980 (fl), Harling & Andersson 16423 (GB[3], TEX[3]); zone<br />
of Sofia, valley of Rio Chingal, 15 Jun 1944 (fr), Steere 8339<br />
(F, NY); Cant6n Archidona, S slope of Vulcan Sumaco, rd.<br />
Hollin-Loreto, km 31, 00?43'S, 77?36'W, 1200 m, 8-17 Jan<br />
1989 (fr), Alvarado 233 (AAU, NY, QCNE); Parque Nacional<br />
Sumaco Napo-Galeras, block 19, 00?53'S, 77?31'W, 1200 m,<br />
22 Apr 1996 (fr), Freire & Cerda 461 (QCNE); Parque<br />
Nacional Galeras, 1 km SW of Comunidad Santa Rosa de Arpino,<br />
block 19, 00?51'S, 77?31'W, 1050 m, 3 Apr 1996 (fr),<br />
Vargas & Grefa 908 (QCNE); rd. El Chaco-El Reventador,<br />
San Rafael waterfall, 1500 m, 77?35'W, 00?06'S, 16 Dec 1993<br />
(fl), Freire-Fierro & Yanez 2697 (NY, QCNE). PASTAZA:<br />
Mera, 25 Mar 1940 (fl-fr), Lugo 115 (S[2]); Mera, 1100 m,<br />
27 Feb 1972 (fl), Harling 11076 (GB[2], TEX[2], US); 1-5 km<br />
NW ofMera, 7 Mar 1980 (fl),Harling & Andersson 17003 (GB);<br />
Mera, Rio Pastaza, Hac. La Gloria above El Topo, 1900 m,<br />
2-9 May 1944 (fr), Steere & Camp 8228 (F[2], NY, US); confluence<br />
of Rio Quijos with Rio Salado, below Baeza, 1930 m, 2<br />
Mar 1944 (fl), Ownbey 2682 (F, IAN, US). SucuMBios: San<br />
Rafael, on rd. El Chaco-Lumbaqui, 1300-1400 m, 77?35'W,<br />
00?06'S, 20 Jun 1996 (fr), StAhl et al. 2800 (AAU, QCA).<br />
TUNGURAHUA: Forest of Volcan Tungurahua, Sodiro s.n. (P).<br />
PERU. HUANUCO: Prov. Pachitea, Pucallpa, W part of<br />
Sira Mtns., 24 km SE of Puerto Inca, 09?28'S, 74?47'W, 29<br />
Jul 1988 (fl), WallnOfer 113-29788 (K[2], LZ, NY, USM,<br />
W). PAsco: Prov. Oxapampa, W side of Cordillera de San<br />
Matias, between Iscozacin and summit, 75?12'W, 10?1 'S,<br />
450 m, 22 Jun 1982 (fl), Smith 2075 (MO).<br />
Dubious collection: Peru. Huanuco: W of Tingo<br />
Maria, 685 m, 19 Sep 1962 (fl), J. Schunke Vigo 6162<br />
(F, MO, US). The label data report these specimens<br />
as being collected from a tree 7-8 m tall, with "moderate<br />
purplish-pink (2.5 RP 7/8)" flowers. These specimens<br />
have semi-leathery leaves, 28-32 x 7.5-10.5 cm<br />
wide; inflorescences sparsely and thinly branched, 36-<br />
41 cm long; capsules 4-5 mm long and 2-3 mm wide.
74 Flora Neotropica<br />
101<br />
Al?<br />
o~~~~~~~~~~~b<br />
AI<br />
10~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
i~~~~~~~~~~~~~?<br />
I e ' ' i- i i i ' 7 a 1<br />
FIG. 29. Distribution of<strong>Rustia</strong> occidentalis (solid circles), R. venezuelensis (open star), R. alba (solid star), R. thibaudi-<br />
oides (solid squares), R. schunkeana (solid triangles), R. rubra (open circles), R. bilsana (open square) and R. viridiflora<br />
(open triangles).<br />
I
Systematic Treatment of the Genera Studied 75<br />
S cmn<br />
.I f<br />
C'<br />
t ~ ~ ~ : Flt<br />
FIG. 30. <strong>Rustia</strong> rubra (A-D from Harling 11076, US; E from Steere 8021, F, holotype). A. Habit of inflorescence<br />
(frondose). B. Flower at anthesis. C. Open anther. D. Detail of style. E. Mature capsule.
76 Flora Neotropica<br />
'':?" ~<br />
...... . .. ~~~~..... I 1 _ _ I .<br />
- JI :,...........: .......... ....<br />
...': : ."... .:.... ?..................................... ....<br />
:.:: . : i<br />
.. :<br />
*::: _ . '.i; i U l |l ll<br />
~ ~ ~ ~ ~ ~ ~ ~ ~ ,Pj<br />
',. :. : :<br />
FIG.31Rstiarbra inforscec.. wit. foeS i ateI and _mtr .? e,, * fut(poobBeil 'J.,=ns . ' : . ..".. ! ' .ig;'; ....:... .Nx '. i Shl. | I | I I I |<br />
, .................<br />
_<br />
_ ~~~~~~~~~~~~~~?:C^',#^?' i '<br />
. . .. i _:i : : > 3 -. ! .. . .. . ........................................<br />
- . :: :~~~~~~~~~~~~~~~~~~~~~~~<br />
?<br />
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ':i?i :?.<br />
* . .. .: ...:'. i. :~ ~ ~ ~ ~~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~.?i~<br />
??. i::<br />
. .. ..~ .....~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
~~ ~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
FIG. 31.?? Ru rubrairsee s t ia~I~<br />
fru it .h<br />
* i ..i.-s.:'....o.X.'.. }...: .. ii. ; .i' > . ^'.,: i .-,xF }-..::;- . ~<br />
FIG. 31. <strong>Rustia</strong> rubra inflorescence, with flowers in anthesis and StChl).i?ie immature fiuits (photo by Bertil<br />
The capsules are much smaller than the normal size<br />
of R. rubra, and probably undersized because infested<br />
by insects, and with circular exit hole on the<br />
capsule wall. Because of its semi-leathery leaves,<br />
sparsely and thinly branched inflorescences, and its<br />
collection locality, I tentatively attribute these specimens<br />
to R. rubra. Nevertheless, future collections might<br />
prove that this collection represents an undescribed<br />
species of <strong>Rustia</strong>.<br />
Simpson (1976) used for this species the name that<br />
was handwritten by Standley on herbarium specimens.<br />
It has pendulous flowers and occurs in the Ecuadorian<br />
and Peruvian Amazon. At the end of the description<br />
of R. rubra, he stated: "Both R. rubra and R.<br />
splendens [= R. thibaudioides] have short deltoid corolla<br />
lobes that are about as long as wide. <strong>Rustia</strong> splendens<br />
[= R. thibaudioides] has typically short petioles<br />
(mostly 1-2 cm long) while in R. rubra they generally<br />
exceed 2.5 cm, and the corolla tube of the former<br />
is usually longitudinally striate, longer, and with larger<br />
diameter than the smaller, non-striate tube of the latter.<br />
The most obvious character separating these two<br />
species is the markedly obtuse leaf base of R. rubra<br />
which contrasts with the broadly acute base of R.<br />
splendens [= R. thibaudioides]." This set of charac-<br />
ters is in agreement with my definition of these two<br />
species, and reliably distinguishes them.<br />
<strong>Rustia</strong> rubra differs from R. schunkeana in hav-<br />
ing the rachis of the inflorescences, flowers, and im-<br />
mature capsules deep red, corolla lobes '/lo of the co-<br />
rolla length, inflorescences densely paniculate with<br />
5-10 lateral branches (vs. racemoid-thyrsoid, un-<br />
branched to 1-3 branches in R. schunkeana).<br />
7. <strong>Rustia</strong> viridiflora Delprete, Novon 5(2): 135, fig.<br />
2. 1995. Type. Ecuador. Morona-Santiago: Achut-<br />
za, Jibaro settlement in the vicinity of Macuma,<br />
ca. 50 km N of Macas, 23 Mar 1973 (fl), Lugo<br />
3674 (holotype, GB). Figs. 6E, 14A,B, 32A-F<br />
Tree to 20 m tall, to 20 cm at dbh. Leafy branch-<br />
lets glabrous to minutely puberulent, terete; older<br />
branches glabrate, rugulose, grayish to pale brown;<br />
lenticels sparse, punctiform to linear, to 1 cm long.<br />
Stipules not seen (readily caducous), leaving a linear
Systematic Treatment of the Genera Studied 77<br />
pA<br />
C<br />
:^\ .A,<br />
:1I<br />
't/.-a --. ; i '<br />
'<br />
C<br />
FIG. 32. <strong>Rustia</strong> viridiflora (A-E from Lugo 3674, GB, holotype; E from Prieto CHUP-20). A. Habit of inflorescence with<br />
mature leaf. B. Abaxial view of blade showing pellucid punctation. C. Flower bud showing bulbose base. D. Open anther.<br />
E. Detail of receptive style. F. Mature capsule showing conical disk. (Reproduced with permission from Novon 5, fig. 2. 1995.)<br />
I
78 Flora Neotropica<br />
scar 1-1.5 mm wide, of the same color of stem. Leaves<br />
38-69x-9.5-20.5 cm, L/W 3:1 to 3.5:1; lanceolate to<br />
elliptic, acute to obtuse (35-65?) at base, acute to attenuate<br />
at apex, sometimes tapering to an acumen 2-<br />
3 cm long; blade dark green above, dull green below,<br />
stiff-foliaceous; drying pale brown, stiff-chartaceous;<br />
glabrous above and below, pellucid punctate; primary<br />
and secondary veins glabrous, prominent below; secondary<br />
veins 19-25 each side; tertiary veins starting<br />
subparallel and openly reticulate in the center, faintly<br />
evident above, fairly evident below; petioles 4.5-7 cm<br />
long, 3-5 mm thick, subterete, glabrous; basally thickened<br />
but not pulvinate; domatia absent. Inflorescences<br />
laxly paniculate, pyramidal, with decussate<br />
racemoid lateral branches; 22-54 x 9-22 cm, lateral<br />
branches 3-7 pairs, basal portion of axis not branched<br />
10-20 cm long; rachis terete to decussately obtusecompressed,<br />
rachis and branches glabrous to minutely<br />
puberulent; flowers on distal racemoid branches or on<br />
racemoid-cymules; bracts subtending lateral branches<br />
1-3 x 2-3 mm, deltoid; bracteoles subtending flowers<br />
0.5-1 x 0.7-1 mm, deltoid, glabrous. Flowers<br />
pedicellate, pedicels 6-9 mm long, glabrous to minutely<br />
puberulent; hypanthium narrowly obconical, 4-<br />
6 x 2-4 mm, glabrous to minutely golden-hirtellous;<br />
flower buds cylindrical. Calyx reduced to a wavy<br />
margin with barely distinguishable lobes, 0.3-0.5 mm<br />
long. Corolla tubular with bulbous base and small<br />
constriction just above, 1.9-2.2 cm long, green<br />
throughout (rarely reddish with green lobes), semifleshy<br />
when fresh; tube subcylindrical, striate, 1.4-<br />
1.8 cm long, 4-5 mm wide at base and 2-2.5 mm wide<br />
at the orifice, glabrous or golden-hirtellous outside;<br />
antrorse-strigose at base inside, without a ring of hairs,<br />
the remaining medial and superior zones glabrous or<br />
with sparse hairs; lobes 5, ?-4/5 of corolla length, 3-<br />
5 x ca. 2.5 mm, triangular to ovate, glabrous outside,<br />
glabrous, with minute glandular hairs (hairs to 0.5 mm<br />
long) at margins inside. Stamens 5, included (only<br />
the very tip exserted), subequal, attached 3.5-5.5 mm<br />
from the base of the tube; filaments 6-7 mm long,<br />
sparsely ascending-pilose, with a tuft of golden-pilose<br />
hairs at base; anthers elongated, rectilinear (never<br />
convex), 11-12 x 1.2-1.3 mm, dorsifixed near base,<br />
base subcaudate, with sharp ridges, papillose-echinate<br />
to ascending-strigulose, descending glandular-pilose<br />
at base. Pollen exine densely foveolate, sparsely minutely<br />
echinate. Style exserted, 22-28 mm long, glabrous<br />
and minutely verrucose throughout; style<br />
branches ovate, acute at apex, 1-1.3 mm long, stigmatic<br />
surface smooth to minutely papillose; immature<br />
fruits green to reddish. Capsules very narrowly<br />
obconical, apex conical, (14-)18-23 x dehiscence absent even in old capsules. Seeds 0.87-<br />
1.27 x 0.33-0.83 mm.<br />
Distribution (Fig. 29) and ecology. Primary and<br />
disturbed forests, from low to medium elevations,<br />
(300-)800-1200 m, of S Amazonian Ecuador. Flowering<br />
specimens were collected in March, October,<br />
and November. Fruiting specimens were collected in<br />
March and November. Its pollination biology has not<br />
been studied, but its flowers are likely to be pollinated<br />
by hummingbirds.<br />
Specimens examined. ECUADOR. MORONA-<br />
SANTIAGO: Near Lim6n (General Plaza Gutierrez),<br />
roadside in remnant forest, 1200 m, 27 Mar 1981 (fr),<br />
Berg 1227 (QCA); rd. Bella Union-Limon, km 30,<br />
78?17'W, 02?48'S, 8 Nov 1991 (fl), Persson et al. 43<br />
(QCA); low hills W of Rio Upano, along Rio Chupiangas,<br />
800-1050 m, 20-22 Nov 1944 (fr), Prieto<br />
CHUP-20 (NY[2], US). PASTAZA: Cushillo Urco, ca.<br />
8 km N of Puerto Sarayacu, 6 Oct 1974 (fl), Lugo 3946<br />
(GB, US); Chapeton on Rio Bobonaza, ca. 8 km ESE<br />
of Canelos, 400-450 m, 77?40'W, 01?38'S, 2-5 Nov<br />
1995 (fl), St&hl et al. 1822 (AAU, QCA); trail between<br />
Chapet6n and Canelos, 400-450 m, 77?43'W, 01 ?35'S,<br />
6 Nov 1995 (fl-fr), Stahl et al. 1866 (AAU, QCA).<br />
<strong>Rustia</strong> viridiflora is unique in the genus in its long<br />
narrowly obconical capsules with conical disk, corollas<br />
basally bulbose, and long rectilinear anthers. Its<br />
corollas are commonly entirely green (rarely red tube<br />
with green lobes). This is one of the four species of<br />
<strong>Rustia</strong> with pendulous flowers (Delprete, 1996a), all<br />
of which have long-narrow corollas and occur in western<br />
South America. A key to the pendulous-flowered<br />
species of <strong>Rustia</strong> was published by Delprete (1996a).<br />
8. <strong>Rustia</strong> schunkeana Delprete, Brittonia 48: 29, fig.<br />
1. 1996. Type. Peru. San Martin: Prov. Mariscal<br />
Cacheres, Tocache Nuevo, NW of the nursery of<br />
Instituto Agropecuario de Tocache, 20 Apr 1970<br />
(fl), J. Schunke Vigo 3952 (holotype, COL;<br />
isotypes, F, G, NY, US). Figs. 6A, 13E, 33A-D<br />
Shrub 3-5 m tall, much branched, with slender<br />
branches, outer branches often scandent. Leafy branchlets<br />
glabrous, dark green, terete; older branches pale<br />
brown to grayish; lenticels sparse, punctiform to linear.<br />
Stipules present only in bud, narrowly triangular,<br />
glabrous outside, glabrous, with few colleters in-<br />
4-6 mm, black, side, 30-40 x 5-6 mm, dark green, readily caducous,<br />
without lenticels, glabrous throughout; disk septicidal leaving a linear whitish scar, ca. 0.5 mm wide. Leaves
Systematic Treatment of the Genera Studied 79<br />
n t<br />
E<br />
FIG. 33. <strong>Rustia</strong><br />
schunkeana (A-C from Schunke Vigo 3952, COL, holotype; D from Schunke Vigo 6696, F). A. Habit<br />
of inflorescence with mature leaf. B. Open flower and flower bud. C. Open anther. D. Mature capsule. (Reproduced with<br />
permission from Brittonia<br />
48, fig. 1. 1. 1996.)<br />
30-45(-60) x 8.5-12 cm, L/W 3:1 to 3.5:1; narrowly<br />
lanceolate, oblanceolate to narrowly elliptic, acute<br />
(BA = 20-30?) at base, acute at apex, sometimes tapering<br />
to an acumen 2-3 cm long, dark green above,<br />
pale green below, subcoriaceous; drying brown, stiff-<br />
chartaceous; glabrous above, glabrous to hirsutulous<br />
below, pellucid punctate; primary and secondary veins<br />
glabrous to hirsutulous, prominent below; secondary
80 Flora Neotropica<br />
veins 15-25 each side; tertiary veins starting subpar-<br />
allel and openly reticulate in the center, evident be-<br />
low; petioles 2-7.5 cm long, 2.5-4.5 mm thick,<br />
subterete, glabrous to hirsutulous, basally thickened<br />
but not pulvinate; domatia absent. Inflorescences<br />
racemoid secundiflorous panicles to thyrsoid-<br />
secundiflorous with opposite or subopposite decus-<br />
sate branches, sometimes without lateral branches,<br />
25-60 cm long, to 17 cm wide, basal branches to 20<br />
cm long, lateral branches usually ascending and to 4<br />
pairs, basal portion of axis not branched 2-15 cm long;<br />
rachis terete, rachis and branches glabrous (rarely hir-<br />
sutulous); flowers alternate on distal branches or<br />
rarely in cymules, usually hanging downward, secund<br />
(pedicel attachments opposite or alternate); distal<br />
bracts 1-3 x 2-3 mm, deltoid; bracteoles subtending<br />
flowers 0.5-0.8 x 1-2 mm, deltoid, glabrous. Flow-<br />
ers pedicellate, pedicels 4-25 mm long, glabrous to<br />
hirtellous; hypanthium narrowly obconical, 4-6 x 2-<br />
2.5 mm, glabrous to hirtellous; flower buds cylindri-<br />
cal. Calyx reduced to a wavy margin with barely dis-<br />
tinguishable lobes, ca. 1 mm long. Corolla tubular<br />
with small erect lobes (not spreading), 2.5-4 cm long,<br />
greenish yellow (rarely pinkish yellow), semi-fleshy<br />
when fresh; tube subcylindrical, faintly striate, 1.5-<br />
2.5 cm long, 2.5-4 mm wide at base and 3.5-5 mm<br />
wide at the orifice, glabrous outside; antrorse-strigose<br />
at base inside, without a ring of hairs, the remaining<br />
medial and superior zones glabrous; lobes 5, 4-'/3 of<br />
corolla length, 9-13 x 1.5-3 mm, triangular to ovate,<br />
glabrous outside, glabrous, with minute conical-bullate<br />
hairs (hairs 0.1-0.3 mm long) inside. Stamens 5,<br />
included (only the very tip exserted), subequal, attached<br />
4.5-5.5 mm from the base of the tube; filaments<br />
6-7 mm long, flattened at base, microscopically<br />
(40x) papillose throughout, minutely barbate at base<br />
with white puberulent hairs; anthers cylindrical, rectilinear<br />
(never convex), 9-12 x 1.5-2 mm, dorsifixed<br />
near the base, base elliptic, smooth throughout. Pollen<br />
exine densely foveolate (non-echinate). Style exserted,<br />
30-40 mm long, glabrous, microscopically<br />
(40x) papillose; style branches ovate, 1-1.5 mm long,<br />
stigmatic surface smooth; immature fruits reddish<br />
green, semi-fleshy. Capsules obovoid to oblong, apex<br />
shallowly hemispherical, (10-)15-20 x 7-9 mm, without<br />
lenticels, dark brown, glabrous throughout; disk<br />
black, obviously exceeding the calyx; disk septicidal<br />
dehiscence absent even in old capsules. Seeds 1.17-<br />
1.67 x 1-1.33 mm.<br />
Distribution (Fig. 29) and ecology. Primary and<br />
secondary lowland rain forests (200-350 m) to pre-<br />
montane wet forests (to 2000 m), often on river banks,<br />
on the W Andean slopes of Amazonian Ecuador and<br />
Peru. Found in association with Grias, Couepia,<br />
Clusia, Ficus, Guarea, Geonoma, Mabea, and Ardisia<br />
(Jaramillo 14167,14290). Flowering specimens were<br />
collected in March, May, and June. Fruiting specimens<br />
were collected in February, June, August, September,<br />
and December.<br />
Specimens examined. ECUADOR. NAPO: Nor-Oriente,<br />
Nuevo Rocafuerte, SW of village, forest toward Rio Braga,<br />
200 m, 27 Feb 1981 (fr), Jaramillo & Coello 4322 (QCA);<br />
Archidona-Tena region, W of Tena, along creek between Rio<br />
Lupi-yacu and Rio Pano, near foot of cordillera de<br />
Guacamayos, 800 m, 25 Mar 1944 (fl), Ownbey 2735 (F[2],<br />
IAN, MO[2], US[2]); Cerro Antisana, 2 km SW of Tena,<br />
00?30'S, 78?W, 700 m, 27 Aug 1960 (fl-fr), Grubb et al. 1485<br />
(K, NY[2]). ZAMORA-CHINCHIPE: Canton Nangaritza, Rio<br />
Nangaritza, between Pachicutza and Miazi, 78?38'W,<br />
04?12'S, 1000 m, 12 Dec 1990 (fr), Neill & Palacios 9718<br />
(MO, QCNE, TEX[2]); 19 Dec 1990 (fr), Palacios 6616<br />
(MO); left margin of Rio Nangaritza, near Miasi military<br />
camp, 04?20'S, 78?40'W, 900 m, 20 Oct 1991 (fr), Jaramillo<br />
14167 (QCA), 950 m, 22 Oct 1991 (fr), Jaramillo 14290<br />
(NY, QCA).<br />
PERU. HUANUCO: Villa Isabel, Rio Cuchara, 21 Sep<br />
1961 (fr), Schunke Vigo 5683 (F[2]). SAN MARTIN: Prov.<br />
Mariscal Caceres, Dtto. Tocache Nuevo, trail to Santa Rosa,<br />
Rio Mishollo, 350 m, 4 Aug 1973 (fr), Schunke Vigo 6696<br />
(F, MO, NY[2]); Prov. Mariscal Caceres, Dtto. Tocache<br />
Nuevo, Prov. Mariscal Cacheres, Santa Rosa de Mishollo,<br />
4 km from Puerto Pizana, 8 May 1971 (fl), Schunke Vigo<br />
4877 (COL, F, G, NY, US); Prov. Mariscal Caceres, Dtto.<br />
Tocache Nuevo, Bambamarca, 20 Jun 1978 (fr), Schunke<br />
Vigo 10270 (F, MO, NY); Prov. Lamas, 2-4 km N of San<br />
Antonio, along Rio Cumbasa, 400 m, 2 Oct-4 Nov 1937 (flfr),<br />
Belshaw 3514 (F, GH, LL, MO, NY, UC, US); Dtto.<br />
Uchiza, 500 m, 23 Jun 1969 (fl), Schunke Vigo 3205 (COL,<br />
F, G, NY, US, VEN).<br />
Local names. Peru: cascarilla masha (San Martin,<br />
Schunke 3952), sacha cascarilla (Huanuco, Schunke<br />
Vigo 5683), cascarillita (San Martin, Schunke Vigo<br />
6696, 10270).<br />
<strong>Rustia</strong> schunkeana is similar to the recently described<br />
R. viridiflora (Delprete, 1995) in having greenish<br />
corollas, the former differing from the latter in<br />
having thyrsoid (vs. paniculate) inflorescences, cylindrical<br />
flowers 25-40 mm long (vs. basally bulbose,<br />
19-22 mm long), and capsules ovoid to oblong (vs.<br />
narrowly obconical).<br />
<strong>Rustia</strong> schunkeana has yellow-green flowers<br />
(rarely with pinkish base), while R. thibaudioides has<br />
pink-red flowers with white apices, and R. rubra has<br />
deep-red flowers sometimes with green apices. <strong>Rustia</strong><br />
schunkeana also differs from R. thibaudioides in hav-<br />
ing corolla lobes l/4--13 of the corolla length (vs. '/o-'/8),<br />
oblong-obovoid capsules 15-20 mm long (vs. obo-
Systematic Treatment of the Genera Studied 81<br />
void, 8-12 mm long), and disk exceeding the calyx<br />
(vs. not exceeding the calyx).<br />
The specimens of Jaramillo 14290 (NY, QCNE)<br />
have been observed to have large, fleshy fruit galls.<br />
The galls develop from the base of the ovary, and<br />
enlarge laterally into an irregular-obovoid shape, 1.5-<br />
2.2 x 0.9-1.5 cm. When sectioned, the galls have longitudinal<br />
burrows of minute larvae of unknown identity<br />
(Coleoptera?).<br />
9. <strong>Rustia</strong> thibaudioides (H. Karsten) Delprete, comb.<br />
nov. Henlea thibaudioides H. Karsten, Linnaea 30:<br />
151. 1859. Type. Colombia. Meta: Mun. San Juan<br />
de Arama, Reserva Nacional Serra de la Macarena,<br />
2 km before La Danta research station, 6 Jan 1993<br />
(fl-fr) Delprete & Fagua 6378 (neotype, NY, selected<br />
here; isoneotypes, COL, F, MO, NY[2],<br />
TEX). Figs. 4C, 6C-D, 13F, 34A-E, 35A-C<br />
<strong>Rustia</strong> splendens (H. Karsten) Standley, Publ. Field.<br />
Columbian Mus., Bot. Ser. 7: 26. 1930. Henlea<br />
splendens H. Karsten, Fl. Columb. 1: 157, pl. 78.<br />
1861. Type. Colombia. Meta: Llano de San Martin,<br />
Jiramene, vic. of Rio Meta, Karsten s.n. (lectotype,<br />
W, selected by Delprete, 1996a; isolectotype, LE?).<br />
According to Tryon (1963), the first set of Karsten's<br />
collections is preserved at LE, but no material has<br />
been seen from this institution. Also, plate N. 78 in<br />
Flora Columbiae is a beautiful color illustration of<br />
the type.<br />
<strong>Rustia</strong> secundiflora K. Schumann in Martius, Fl. Bras.<br />
6(6): 261. 1889. Type. Colombia. Caqueta: Upper<br />
Amazonas, at base of Mt. Araracoara, Jan-Feb (fl),<br />
Martius 3139 (B*; lectotype, M, selected by<br />
Delprete, 1996a; photo-M at F, MO, VEN).<br />
Shrub 5-6 m tall or tree 8-10(-12) m tall, to 25 cm<br />
dbh, much-branched shrub to much-branched singlestemmed<br />
tree; bark yellowish-grayish. Leafy branchlets<br />
glabrous to golden-pubescent, dark green, terete;<br />
older branches glabrate, rugose, grayish to pale brown;<br />
lenticels sparse, small. Stipules narrowly triangular,<br />
glabrous to minutely puberulent to hirtellous outside,<br />
glabrous with few colleters inside, 20-30 x 4-5 mm,<br />
dark green, readily caducous, leaving a grayish scar<br />
encircling the stem, evident in older branchlets.<br />
Leaves 25-50(-65) x (6-)10-17 cm, L/W 2:1 to 3.5:1;<br />
lanceolate to oblanceolate to obovate, acute to rounded<br />
(BA = tellous (rarely hirsute) below; pellucid punctate; primary<br />
and secondary veins glabrous to hirtellous<br />
(rarely hirsute), prominent below; secondary veins<br />
11-18 each side; tertiary veins starting subparallel and<br />
openly reticulate in the center, faintly evident below;<br />
petioles 1.5-6 cm long, 1.5-4.5 mm thick, terete to<br />
adaxially concave to flattened, pulvinus present in<br />
young leaves, becoming corky in older leaves;<br />
domatia absent. Inflorescences openly to densely<br />
paniculate to thyrsoid-secundiflorous with opposite or<br />
subopposite decussate branches; (12-)25-70(-153!)<br />
cm long, 5-25(-45) cm wide at base, lateral branches<br />
to 15 pairs, basal portion of axis not branched 5-25<br />
cm long; rachis terete to decussately obtuse-compressed,<br />
rachis and branches glabrous or puberulent<br />
to hirtellous; flowers alternate on distal branches or<br />
rarely in cymules, usually pointing downward, giving<br />
the false appearance of secundiflorous position<br />
(pedicel attachment opposite and not secundiflorous);<br />
distal bracts 1-4 x 2-4 mm, deltoid; bracteoles subtending<br />
flowers 0.5-1 x 0.5-1 mm, deltoid, glabrous<br />
to minutely puberulent. Flowers pedicellate, pedicels<br />
3-15 mm long, glabrous, puberulent to hirtellous,<br />
often curved downward; hypanthium narrowly<br />
obconical, 2-5 x 1.5-2.5 mm, glabrous to puberulent;<br />
flower buds cylindrical. Calyx extremely reduced,<br />
truncate or with minutely wavy margin, 0.3-1 x 2-4 mm,<br />
glabrous to puberulent. Corolla tubular, 2-2.5(-3) cm<br />
long, white apically and pink-reddish at base outside,<br />
white inside, semi-fleshy when fresh; tube subcylindrical,<br />
striate, 1.9-2.2 cm long, 2.5-3.5 mm wide at<br />
base and 2.5-3.5 mm wide at the orifice, glabrous to<br />
puberulent to hirtellous outside; antrorse-strigose to<br />
strigulose at basal 2-3/4 of the tube inside, without a<br />
ring of hairs; lobes 5, '/,o-'/ of corolla length, 1.5-4.5<br />
x 1.5-2.5 mm, deltoid, glabrous to minutely puberulent<br />
outside, glabrous, minutely papillose (40x) at<br />
margins inside. Stamens 5, included or only the tips<br />
exserted, subequal, attached 5-7.5 mm from the base<br />
of the tube; filaments 7-8.5 mm long, minutely (40x)<br />
ascending strigulose to papillose at lower half portion;<br />
anthers cylindrical, narrowly oblong, rectilinear or<br />
slightly convex, 5-6.5 x 0.9-1.4 mm, dorsifixed near<br />
the base, base rounded, microscopically (40x) papillose-echinate<br />
(rarely smooth) throughout, dehiscing<br />
by two pores at apex. Pollen exine foveolate, minutely<br />
echinate. Style exserted, 30-35 mm long, glabrous,<br />
subtetragonous; style branches ovate, 0.4-1 mm long,<br />
stigmatic surface smooth. Capsules obovoid to widely<br />
20-40?) at base, acute to obtuse-acuminate at obovoid, acute at base, apex subtruncate to obtusely<br />
apex, the acumen, when present, 1-2 cm long, dark hemispherical, 8-12 x 6-7(-10) mm, dark brown<br />
green above, pale olive-green below, foliaceous to without lenticels, glabrous to puberulent below the<br />
semi-coriaceous; drying green-brownish, stiff-chart- disk; disk glabrous; disk septicidal dehiscence absent<br />
aceous; glabrous above; glabrous, puberulent to hir- even in old capsules. Seeds 1.4-1.5 x 0.75-1.25 mm.
82 Flora Neotropica<br />
l I l<br />
'4i ~A<br />
C CA<br />
FIG. 34. <strong>Rustia</strong> thibaudioides (from fresh material and Delprete & Fagua 6378, TEX). A. Habit of inflorescence with<br />
mature leaves. B. Flower, lateral view and inside view. C. Flower bud. D. Open anther. E. Mature capsule. (Reproduced<br />
with permission from Brittonia 48, fig. 2. 1996 [as R. splendens].)
Systematic Treatment of the Genera Studied 83<br />
:., ...... .,,,,~! 1<br />
F -? 6..o,2'?C~<br />
.<br />
...' ....~<br />
.I~~~~~~~~~~~~~~~~~~~~~~~~~N<br />
? .:<br />
? , '": .. ...<br />
_ :.f<br />
'' ,-' ;'<br />
.<br />
'<br />
; ,............. ..<br />
''~~~~~~~~~~~~~:;:,? i.? :<br />
8 ...'"....~., .. ,, ~:'* :';, t.';..~....: ... :?:, ~<br />
.. ~!._ , ,7 . ~,..>...:i, ......:...,.<br />
.~ ~ = ,~ ,'. 4 , ................ .....<br />
~~??Xl. _ d? _ .:~~?~ . ,<br />
~.~~~~~~~~~~~~~~~~~g :i:.r':<br />
|~~~~~~~~~~~~~~~~~~~~~~~~~::? I~<br />
??<br />
. 35.: Rd . A. Sal (o_, a mI .U..': c,i<br />
'' .;r r iBsFI ~<br />
I 11 '&Q i'<br />
anhsi, C.' Flwri"nhsssenfo otmuwad(oeicue ope'n anthers..<br />
: -:<br />
i?-.<br />
r<br />
;' "<br />
.<br />
. ....<br />
,<br />
I"' .. . ..:_ ii .. @ " . _: __.<br />
'~~~l ,'~' :_i;,,, ,?-_4-_'"".<br />
,._~~~ o -.<br />
fiF: .--<br />
. _<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?...L|.<br />
.. '1:"~~' ,<br />
:..~ie<br />
FIG. 3S. <strong>Rustia</strong> thibaudioides. A. Sh--b F5 m tall (arrow indicates main trunk). B. Inflorescen.c ca. I m long, in<br />
anthesis. C. Flower in anthesis seen f.om bottom upward (note included open anthers).<br />
r<br />
.
84 Flora Neotropica<br />
Distribution (Fig. 29) and ecology. Gallery forests<br />
of central (states of Boyaca, Cundinamarca, Santander,<br />
Tolima) and Amazonian Colombia, to 1400<br />
m, in lateritic soils. A single specimen was collected<br />
in Venezuela (state of Tachira), and a few specimens<br />
were reportedly collected in Brazil and their labels<br />
read: "Martius Iter Brasiliense, Rio Negro, Brazil."<br />
This species has long inflorescences with up to a few<br />
hundred flowers, and the blooming period of a single<br />
individual could last up to two months. Flowering<br />
material was collected in January, February, March,<br />
July, October, November, and December. Specimens<br />
in early fruiting stage have been collected in, January<br />
and December.<br />
Near Rio Negro, E of Prado, 10 Feb 1944 (fl), Little 7186<br />
(COL, US). VAUPtS: Environs of Mitfi, trail to Cerro Miti,<br />
29 Oct 1976 (fl), Davis 184 (COL, F, GH); cabeceras of Rio<br />
Cuduyary, 10 Dec 1943 (fl), Alien 3217 (COL, G, MO, US);<br />
Rio Vaupes, Miti, 9 Oct 1966 (fl), Schultes 24356 (COL).<br />
VENEZUELA. TACHIRA: Ca. 35 km SSE of San Crist6bal,<br />
La Buenafia, 6-12 km W of Quebrada Colorado, 600-<br />
1200 m, 07?28'N, 72?09'W, 20-21 Mar 1981 (fr), Liesner<br />
& Gonzdles 10940 (VEN).<br />
BRAZIL. AMAZONAS: Rio Negro, Martius Iter Brasiliense<br />
2607 (M), 2608 (M), s.n. (M).<br />
Local name. Colombia: doncello (Meta, Fosberg<br />
19440).<br />
Pollination biology. This species was observed in<br />
<strong>Rustia</strong> thibaudioides is unique in the genus in having<br />
apically white and basally pink corollas, included<br />
bloom in the Serrania de La Macarena (Department anthers, deltoid corolla lobes 1/lo-'/8 of the corolla<br />
of Meta). The flowers of R. thibaudioides are mainly length, and obovoid capsules 8-12 mm long with disk<br />
pendulous, sometimes as many as several hundred per not exceeding the calyx (Fig. 34B-E); the closely reinflorescence,<br />
pinkish red at base and white at the lated R. schunkeana has deltoid corolla lobes ?4-'/3<br />
medio-distal portion, and the anthers remain included of the corolla, anthers excluded at anthesis, and obin<br />
the corolla tube at anthesis. They are frequently long-obovoid capsules 15-20 mm long (Fig. 33B-D).<br />
visited by hummingbirds, bees, wasps, and butterflies.<br />
The bees most frequently visiting the flowers are of<br />
the genera Melipona, Ceratina, and Trigona (2 spp.)<br />
(pers. obs.). No detailed pollination biology studies<br />
were performed, but I suspect that this species is pollinated<br />
primarily by hummingbirds and secondarily<br />
by bees. Some flowers were observed to be robbed<br />
by wasps through lateral incisions. Many individuals<br />
were also guarded by ants, which are probably feeding<br />
on the easily accessible floral nectaries.<br />
Karsten (1859) cited Henlea thibaudioides, the<br />
type species of Henlea, as "Crescit in planitie ad orientalem<br />
Cordillerae Bogotensis ad littora fluminis<br />
Meta" [Colombia, Cordillera Oriental, Rio Meta]. No<br />
types of this species were found at W, and I had no<br />
access to the LE specimens. Because of this, I am<br />
selecting as the neotype for H. thibaudioides a specimen<br />
ofDelprete & Fagua 6378 (NY) that corresponds<br />
to its original description and was collected close to<br />
the locality cited by Karsten (Serrania de la Macarena,<br />
near the riverwash of Rio Meta). Henlea thibaudioides<br />
Representative specimens examined. COLOMBIA. is here transferred to <strong>Rustia</strong>, and <strong>Rustia</strong> and Henlea<br />
ANTIOQUIA: San Rafael, 29 Nov 1958, Daniel 5533 (US). are congeneric.<br />
BOYACA: Llano de San Martin, Bogoti, 300 m, [1851-1857], A discussion of the lectotypification and synonymy<br />
Triana 1785 (3262.4) (P); Llano de San Martin, Jiramena, of <strong>Rustia</strong> splendens and R. secundiflora was presented<br />
220 m, Jan 1856 (fl), Triana s.n. (3262) (F, COL). CAQUETA: in<br />
Rd. Puerto Rico-Florencia, left<br />
Delprete, 1996a. The<br />
side of Rio Guayas, 350lectotype<br />
ofHenlea splendens<br />
400 m, 7 Oct 1975 (fl), Cabrera 3701 (F); confluence of Rio (= R. thibaudioides) is a specimen preserved at W,<br />
Mesay with Rio Cufiar6, 1-6 Mar 1980 (fl), Pabdn 960<br />
with embossed "Dr. Karsten," and labeled in his hand-<br />
(COL[2]). CUNDINAMARCA: Near Guayabetal, trail to Manza- writing "Henlea splendens - Llano de St. Martin." The<br />
nares, Valley of Rio Negro, Western Cordillera, 1400 m, 3 lectotype of R. secundiflora is a specimen preserved<br />
Jan 1956 (fl-fr), Vogel 54 (VEN). GUAVIARE: Cafio Grande, at M, with its label stating "Mart. Obs. 3139 - ad M.<br />
between Calamar and San Jose del Guaviare, 240 m, 1 Nov Araracoara - Dr. Martius Iter Brasil: Jan-Feb." On a<br />
1939 (fl), Cuatrecasas 7370 (F). META: Uribe, Cordillera small label glued to the herbarium sheet, Martius an-<br />
Oriental, 770 m, 20 Dec 1942 (fl-fr), Fosberg 19440 (NY[2], notated this specimen as "Hamelia albopurpurea," an<br />
UC, US[2]); Mun. San Juan de Arama, Reserva Nacional<br />
unpublished name.<br />
Serra de la Macarena, 2 km before La Danta research sta-<br />
<strong>Rustia</strong> warscewicziana Klotzsch is another<br />
tion, 6 Jan 1993 (fl-fr) Delprete & Fagua 6381<br />
unpub-<br />
(COL, F, MO,<br />
lished name for this taxon. The<br />
TEX), 6382 (COL, NY); Caflo Lozada, riverside of Rio<br />
corresponding her-<br />
Guayabero, 350 m, 30 Jan 1959 (fl), Pinto et al. 370<br />
barium<br />
(COL, specimens were preserved at Berlin (B) and<br />
P). PUTUMAYO: Rio San Miguel, left affluent Quebrada La were destroyed. Fortunately the specimens were pho-<br />
Hormiga, 290 m, 15 Dec 1940 (fl), Cuatrecasas 11076 tographed by Macbride (negatives N. 10 and 11) and<br />
(COL, F, US). SANTANDER: Region around Landazuri, 70 km the photographs distributed to several herbaria (F,<br />
N of Velez, 1 Jul 1944 (fl), Fassett 25427 (US). TOLIMA: MO, NY, US). These specimens were collected by
Systematic Treatment of the Genera Studied 85<br />
Warscewicz in Colombia ("Neu Grenada"), in the tending primary branches 1-3 x 1-2 mm, deltoid;<br />
southern Cordillera Occidental ("cordillera mer. bracteoles subtending flowers ca. 0.5 mm long, deloccident."),<br />
an area very close to the type locality of toid. Flowers not seen. Pollen not seen. Capsules<br />
Henlea splendens (= R. thibaudioides). Simpson subglobose to obovoid, acute at base, apex hemi-<br />
(1976) studied the two photographs, and referred to spherical, 6-9 x 6-6.5 mm, dark brown to black with-<br />
R. warscewicziana as follows: "It is certainly a <strong>Rustia</strong> out lenticels, glabrous throughout; disk black, exceedbut<br />
the panicle seems much larger and more branched ing the calyx; pedicels below mature capsules 9-15<br />
that any I have seen otherwise." After close exami- mm long. Seeds 0.93-1 x 0.4-0.57 mm.<br />
nation of the photographs I concluded that the destroyed<br />
specimens pertain to R. thibaudioides. Distribution (Fig. 29) and ecology. This species<br />
Extensive field research has been carried out in the is known only from the type specimens, reportedly<br />
area where the three types were obtained, and I have collected from a tree 10-12 m tall growing on dry<br />
collected R. thibaudioides at several localities. This canyon slopes, ca. 1550 m, in the state of Merida,<br />
species is a very beautiful shrub, usually 6-7(-10) m Venezuela. The single fruiting specimen was collected<br />
tall, with very attractive inflorescences that vary in in May. Flowering specimens are still unknown.<br />
size and shape, sometimes to 1.5 m long, most of the<br />
time horizontal and sometimes ascendent. In the in- This is the only species of <strong>Rustia</strong> collected in Venflorescences<br />
of R. thibaudioides the flowers are pen- ezuela. Steyermark (1974) also reported R. longifolia<br />
dulous, which in herbarium specimens have been er- Standl., but this taxon is instead Dolichodelphys<br />
roneously interpreted as secund.<br />
chlorocrater.<br />
<strong>Rustia</strong> venezuelensis is similar to R. formosa, of<br />
southern Brazil; the former is presently kept distinct<br />
from the latter because of its smaller sparsely flow-<br />
10. <strong>Rustia</strong> venezuelensis Standley & Steyermark, ered inflorescence and its very isolated geographical<br />
Fieldiana, Bot. 28: 616. 1953. Type. Venezuela. distribution. Future collections of flowering speci-<br />
Merida: Between Mucuchachi and Canagua, be- mens will be crucial in establishing the relationships<br />
low El Bao, 1570 m, 6 May 1944 (fr), Steyermark of this species.<br />
56333 (holotype, F; isotype, VEN). Fig. 6G,H<br />
Tree 10-12 m tall, much branched; bark grayish.<br />
Leafy branchlets glabrous, pale brown-grayish, terete;<br />
older branches rugose, white-grayish. Stipules not<br />
seen (readily caducous), leaving a linear scar ca. 0.5<br />
mm wide, of the same color as stem. Leaves 20-37 x<br />
8-14 cm, L/W 2:1 to 2.5:1; widely obovate, acute (BA<br />
?30?) at base, narrowly obtuse, acuminate at apex; the<br />
acumen deltoid, 1-1.5 cm long; olive-green above and<br />
pale green below, subcoriaceous; drying olive-green,<br />
semi-coriaceous; glabrous above and below; pellucid<br />
punctate; primary and secondary veins glabrous,<br />
prominent below, secondary veins 12-15 each side;<br />
tertiary veins starting subparallel and openly reticulate<br />
toward the center, evident only above; petioles<br />
15-35 mm long, 2-3 mm thick, subterete or adaxially<br />
flattened, pulvinus absent but thickened becoming<br />
corky at base, glabrous; domatia absent. Inflorescences<br />
reduced paniculate, with opposite decussate<br />
ascending branches; 17-28 x 13-24 cm, basal primary<br />
branches 8-15 cm long, lateral branches 3-6 pairs,<br />
basal portion of axis not branched ca. 7 cm long; rachis<br />
terete or decussately obtuse-compressed, rachis<br />
and branches glabrous; flowers alternate on distal<br />
branches or in terminal lax cymules; distal bracts sub-<br />
11. <strong>Rustia</strong> formosa (Chamisso & Schlechtendal ex<br />
de Candolle) Klotzsch in Hayne, Getr. Darstell.<br />
Gew. 14: tab. 15. 1846. Exostema formosum<br />
Chamisso & Schlechtendal ex de Candolle, Prodr.<br />
4(4): 361. 1830. Exostema formosum Chamisso<br />
& Schlechtendal var. fi laeve Chamisso &<br />
Schlechtendal ex de Candolle, nom. inval., Prodr.<br />
4(4): 361. 1830. Exostema formosum Chamisso<br />
& Schlechtendal, nom. nud., Linnaea 4:179. 1829.<br />
Exostema formosum var. fi laeve Chamisso &<br />
Schlechtendal, nom. inval., Linnaea 4:179. 1829.<br />
Type. Brazil. Sao Paulo: Estagao Biologica Alto<br />
da Serra, 19 Oct 1931 (fl-fr), C. Lemos 28678 (neotype,<br />
NY, selected here; isoneotypes, F, GH, US).<br />
Figs. 3C, 5C, 14E, 36A-E<br />
<strong>Rustia</strong> sellowiana Klotzsch in Hayne, Getr. Darstell.<br />
Gew. 14: tab. 15. 1846 (validated by direct refer-<br />
ence to description of Exostema formosum var.<br />
leprosum Chamisso & Schlechtendal). Exostema<br />
formosum Chamisso & Schlechtendal var. leprosum<br />
Chamisso & Schlechtendal ex de Candolle, nom.<br />
inval., Prodr. 4(4): 361. 1830. Exostemaformosum<br />
var. leprosum Chamisso & Schlechtendal, nom.
86 Flora Neotropica<br />
inval., Linnaea 4: 179. 1829. Type. Brazil. Rio de<br />
Janeiro: Corcovado, Caminho de Larangeiros, Jan<br />
1834 (fl), Luschnath 179 [Martius H. Fl. Bras. 111]<br />
(neotype, BR, here selected). Martius H. Fl. Bras.<br />
111 (B*, BR, frag-F, G, K, M, NY, P) are dubious<br />
isoneotype specimens because their labels report<br />
various collectors, localities, and dates. In Herbarium<br />
Florae Brasiliensis, Martius (1837) reports N. 111<br />
as "Exostemma formosum Cham. et Schlecht.<br />
Linnaea IV. p. 179. Quina do Rio de Janeiro:<br />
Brasiliens. Arbor 25-pedalis. - Est forma leprosum<br />
Cham. et Schlecht. 1. c. In monte Corcovado et in<br />
Caminho de Larangeiras: fl. Jan. Dryas."<br />
<strong>Rustia</strong> pohliana Klotzsch in Hayne, Getr. Darstell. Gew.<br />
14: tab. 15. 1846. Type. Brazil. Goias: Ponzo Alto a<br />
Caldas Nuovas, 1837 (fl), Pohl 809 (lectotype, W,<br />
here selected; isolectotypes, F, GH).<br />
<strong>Rustia</strong> mosenii K. Schumann ex Glaziou, nom. nud., Bull.<br />
Soc. Bot. France 56, mem. III: 337. 1909. The fourword<br />
description does not distinguish this from other<br />
species (Glaziou, 1909). Cited specimens: Glaziou<br />
21548a (B*, C-n.v., K).<br />
Shrubs 4-6 m tall, exceptionally trees to 12 m tall,<br />
to 30 cm dbh, much-branched, sometimes with small<br />
buttresses; bark yellowish grayish. Leafy branchlets<br />
grayish-tomentulous, tetragonous-terete; older<br />
branches glabrate, rugose, terete, grayish; lenticels<br />
punctiform, 0.5 mm wide. Stipules narrowly triangular,<br />
acuminate, tomentulous outside, glabrous with<br />
basal colleters inside, (15-)25-35 x 13-16 mm, pale<br />
green, readily caducous, leaving a white-grayish linear<br />
scar. Leaves (22-)25-38(-54) x (7-)9-17(-23)<br />
cm, L/W 1.6:1 to 2:1; elliptic to oblanceolate to obovate,<br />
acute to obtuse (BA = x 1-1.5 mm, deltoid, glabrous to minutely puberulent,<br />
often ciliolate. Flowers pedicellate, pedicels 3-5 mm<br />
long, glabrous (rarely minute puberulent), fragrant;<br />
hypanthium narrowly obconical, 3-5 x 1.5-2.5 mm,<br />
glabrous; flower buds cylindrical. Calyx smallcupular,<br />
truncate or with barely distinguishable lobes,<br />
3-4 mm wide; lobes 5, 0.2-0.4 x 0.9-1.2 mm, shallowly<br />
deltoid, ciliolate. Corolla tubular with spreading-reversed<br />
lobes, 1.5-1.8 cm long, white to creamwhite,<br />
rarely pink-reddish at base outside, white<br />
inside, semi-fleshy when fresh; tube narrowly funnelform,<br />
9-12 mm long, 2-2.2 mm at base and ca. 2.5<br />
mm wide at the orifice, glabrous to minutely puberulent<br />
outside, glabrous inside, with a ring of white-sericeous<br />
hairs (0.5-1 mm long), 4-5 mm from the base,<br />
at the same point of the filament attachments at base<br />
of lobes; lobes 5, '/3-l of corolla length, 5-7 x 0.9-<br />
1.2 mm, narrowly triangular, glabrous to minutely<br />
puberulent outside, white-strigose at base (hairs 0.5-<br />
1 mm long), margins densely papillose inside. Stamens<br />
5, exserted because of spreading lobes, subequal,<br />
attached 4-5 mm from the base of the tube; filaments<br />
4-5 mm long, very thin, pendulous, glabrous, barbate<br />
at base, with a tuft of white sericeous hairs (0.8-1 mm<br />
long); anthers cylindrical, narrowly oblong, 6-8 x 1-<br />
1.5 mm, dorsifixed near the base, base rounded, mi-<br />
30-50?) at base, rounded<br />
at apex; dark green above, pale green-grayish below,<br />
semi-coriaceous; drying grayish olive-green, chartacroscopically<br />
(40x) papillose-echinate throughout.<br />
Pollen exine densely foveolate (non-echinate). Style<br />
exserted, 20-25 mm long, glabrous; style branches<br />
oblong, 0.9-1.1 mm long, stigmatic surface glabrous<br />
to microscopically (40x) papillose. Capsules obovoid<br />
to subglobose, acute at base, apex subtruncate, 5-7<br />
(-8)<br />
ceous; glabrous above, glabrous to golden-pilose below;<br />
pellucid punctate; primary and secondary veins<br />
glabrous to golden-pilose, prominent below, secondary<br />
veins 12-16 each side, subparallel; tertiary veins<br />
openly reticulate, faintly evident; petioles (10-) 13-<br />
18(-25) mm long, 2.5-3 mm thick, adaxially concave<br />
to flattened, glabrous to golden-pilose, pulvinus absent;<br />
domatia absent or a tuft of sparse hairs 1-1.5 mm<br />
wide, hairs white 0.2-0.4 mm long. Inflorescences<br />
openly paniculate, pyramidal, with opposite or subopposite<br />
decussate branches; 24-30(-36) cm long, (8-)<br />
14-20 cm wide at base, lateral branches 4-8 pairs,<br />
basal portion of axis not branched (4-)6-14 cm long;<br />
rachis decussately compressed or quadrangular (rarely<br />
terete) in cross section, rachis and branches glabrous<br />
(rarely minute- puberulent); flowers alternate on distal<br />
branches; distal bracts decreasing distally from 12<br />
to 3 mm long, 1-1.5 mm wide, linear, narrowly triangular<br />
to deltoid, bracteoles subtending flowers 1-1.5<br />
x ca. 5 mm, dark brown with small lenticels, glabrous<br />
throughout. Seeds 1.4-1.6 x 0.48 mm.<br />
Distribution (see Fig. 38A) and ecology. Margins<br />
and gaps in mesophytic gallery forests and Cerrado,<br />
usually in proximity of creeks and fast rivers of SE<br />
Brazil in the states of Goias, Minas Gerais, Distrito<br />
Federal, Rio de Janeiro, and SAo Paulo, at 100-1000<br />
m. Flowering specimens were collected in all months<br />
of the year except December and January. Fruiting<br />
specimens were collected in June, August, October,<br />
and November.<br />
Specimens examined. BRAZIL. DISTRITO FEDERAL:<br />
Cachoeirinha, affluent of left margin of Rio Paranoa, 26 May<br />
1982 (fl), Pereira 270 (K); hills ca. 10 km N of Planaltina,<br />
975 m, 2 Oct 1965 (fl-fr), Irwin et al. 8881 (MO, NY, US,<br />
VEN); 50 km E of Brasilia, 700-1000 m, 20 Aug 1964 (flfr),<br />
Irwin & Soderstrom 5369 (F, NY, UC, US, VEN); S slope<br />
ofMorro da Canastra, 920 m, 15?35'S, 47?54'W, 19 Oct 1983<br />
(st), Kirkbride 5431 (F, NY); estrada de Sobradinho, 24 Jan<br />
1965 (fl), Sucre & Heringer 624 (IAN, VEN); Sumare, 23<br />
Feb 1959 (fl), Pereira & Duarte 4484 (RB); between
Systematic Treatment of the Genera Studied 87<br />
t4r<br />
0~~~~~~~~~~~~<br />
tl<br />
..<br />
.~~~~~~~~~ I~~~~~~~~~~~~~<br />
....<br />
,'.~~~~~~~~~~~~'<br />
FIG. 36. <strong>Rustia</strong>formosa (A-D from Pereira 270, MO; E from Irwin et al. 8881, NY). A. Habit of inflorescence with<br />
view of blade showing pellucid punctation. E. Mature capsule.
88 Flora Neotropica<br />
Calcareo and Silva, 1 May 1963 (fl), Pires et al. 9539 (F, ofMartius 111 reports the common name "Quina do<br />
NY); Parque Municipal do Gama, 1000 m, 21 Mar 1966 (fl), Rio de Janeiro," probably because its bark was used<br />
Irwin et al. 14160 (NY); Rio Sao Bartolomeu, 22 Apr 1980 as a secondary source of quinine. Reported as<br />
(fl), Heringer et al.<br />
very<br />
4448 (NY, US), 12 Jun 1980 (fr), 5082<br />
ornamental tree with<br />
(MO, UB, US), 9 Feb 1981 (fl), 6157(MG, NY, US); Escola<br />
fragrant flowers.<br />
Agro-T6cnica de Brasilia, Beira do Rio, 23 Jul 1963 (fl-fr),<br />
Heringer 9156 (VEN); Area de Protegco Ambiental de<br />
Dubious collection: Brazil. "Prov. S. Pauli," 1868,<br />
Cafuringa, Fazenda Palestina, 4 Mar 1993 (fl), Pereira & Burchell 3181 (BR, K[3]). These specimens have very<br />
Mecenas 2442 (US); Area de Protenao Ambiental de large leaves (to 51 cm long and 21 cm wide!) that are<br />
Cafuringa, 26 Mar 1993 (fl), Pereira & Alvarenga 2503 (SP, obviously pellucid-puncated; stipules exceptionally<br />
US). GOIAS: Prov. Alexania, Rio Arelia, 9 Aug 1990 (fl), De large, 3.5-4.5 x 5-7 mm, long-lanceolate, entirely<br />
Paula 3233 (UB); Mun. Caldas Novas, Serra de Caldas [No- glabrous; inflorescences rather stout, woody, 33 cm<br />
vas], 10 Jul 1976(f), Hatschbach 38796 (US); Goias, 1844<br />
long, with opposite-decussate branches old fruits of<br />
(fl), Weddell 2637 (P), 2638 (P); Serra Dourada, ca. 17 km<br />
very small size (7-8<br />
S ofGoiis Velho, 6 km NE of Mossamendes, 750 m, 9 May<br />
1973 (fl), Anderson 9927 (NY, US); Serra Dourada, 27 May<br />
1965 (fl), Heringer 10468 (F, RB); Contraforte Central, NE<br />
of Catallo, 875 m, 23 Jan 1970 (fl), Irwin et al. 25206 (F,<br />
MO, NY, US, VEN). MINAS GERAIS: Mun. Corrego<br />
Barreiras, rd. BR-365, Pato de Minas, 26 Mar 1980 (fl),<br />
Hatschbach 42945 (MG, NY, UB, UC); Mun. Araguari, Rod.<br />
BR-40, Rib. Jordao, 13 Apr 1981 (fl), Hatschbach 43851<br />
(BR, F, INPA, NY, SP, US); Mun. Paracatu, Fazenda<br />
Acangau, 17?12'S, 47?06'W, 5 Mar 1989 (fl), Mendonca et<br />
al. 1292 (UB, US); Teixeira Loares, Mar 1908 (fl), Sampaio<br />
612 (F); Faz. do Brejao, 31 Jul 1960 (fl), Heringer 7581<br />
(UB); 1838, Claussen 669 (F, NY); 1839, Claussen s.n. (G);<br />
1816-1821, Saint-Hilaire 348 (F, P). Rio DE JANEIRO: Rio<br />
de Janeiro, rd. Vista Chinesa, 500 m to Parque Nacional<br />
Tijuca, 20 Jun 1985 (fl), Argeli & Oliveira 691 (HRB, US);<br />
Rio de Janeiro, Vista Chinesa, 1 Mar 1931 (fl), Brade 10639<br />
(F, GH); Rio de Janeiro, Campo Grande, Serra Mendanha,<br />
600-700 m, 4 Apr 1978 (fl), Martinelli et al. 4133 (RB); Rio<br />
de Janeiro, Mt. Corcovado, 1866, Glaziou Ib" (BR, F); Rio<br />
de Janeiro, 1833, Riedel 1236 (BR, US); Mun. Parati,<br />
Laranjeiras, trail to Praia do Sono, 100 m, 10 Jun 1991 (fr),<br />
Giordano et al. 1005 (RB); Mun. Parati, trail to Morro da<br />
Pedra Rolada, 300 m, 28 Jun 1995 (fr), Campos et al. 48<br />
(RB); Gavea, E Guanabara, 4 Mar 1947 (fl), Duarte et al.<br />
s.n. (RB 60854) (NY); Gavea, 4 Mar 1947 (fl), Apparicio<br />
59161 (F[2]); old rd. down serra to Angra dos Reis, 1 Feb<br />
1968 (fl), Smith & McWilliams 15364 (US); Rio de Janeiro,<br />
1833, Vauthier 205 (F, G, K); Rio de Janeiro, Apr 1836 (fl),<br />
Vauthier 15 (F, G); Rio de Janeiro, 1844 (fl), Widgren (BR,<br />
F, US). SAo PAULO: CubatAo, 9 Aug 1899 (fl), Edwall 23938<br />
(F); Cubatao, Serra de Morrao, Rio Quilombo, 5 Jun 1990<br />
(fr), Kirizawa & Chiea 2350 (SP); Cubatao, 15 Mar 1923<br />
(fl), Gehrt s.n. (SP); slope of Serra do Mar, estrada velha,<br />
Cubatlo, 10 Apr 1956 (fl), Kuhlmann 3769 (F, NY, TEX);<br />
Santos, Feb 1875 (fl), Mosen 3408 (F[3], P); Reserva<br />
Ecol6gica de Jur6ia, Peruibe, 11 Nov 1982 (fr), Rodriguez<br />
& Figuereido 15745 (UB). WITHOUr LOCALITY: Brazil, Pohl<br />
2681 (M); Brazil, 1839 (fl), Pohl s.n. (M); Brazil, Feb-Mar<br />
1823, Riedel s.n. (M).<br />
x 4-5 mm), already open. Because<br />
of its small fruits, large leaves, and collection<br />
locality, I tentatively attribute these specimens to R.<br />
formosa, but flowering collections of these individuals<br />
might prove to be an undescribed taxon.<br />
<strong>Rustia</strong> formosa is a beautiful shrub, with white,<br />
fragrant flowers easily distinguishable from the other<br />
species of <strong>Rustia</strong> by their throat and lobes (internally)<br />
densely pubescent. From R. alba (of the Ecuadorian<br />
cloud forests) this species differs in the extremely<br />
reduced calyx and its geographical distribution. <strong>Rustia</strong><br />
simpsonii differs from R. formosa in having much<br />
larger, thicker corollas, short-glandular hair at the<br />
internal margins of the corolla lobes (vs. densely pubescent),<br />
and cupular-membranous calyx with obvious<br />
lobes (vs. extremely reduced).<br />
This species was first presented by Chamisso and<br />
Schlechtendal (1829) as Exostema formosum, with<br />
two varieties (var. a leprosum and var.? laeve). There<br />
was no description for E. formosum, but there were<br />
descriptions for each of the two varieties. Therefore,<br />
E. formosum is a nomen nudum, and the two varieties<br />
are invalid because they were attributed to an invalid<br />
Local names and uses. Brazil: caapeba (Minas<br />
Gerais, Sampaio 612, Hoene 11865), guapeba branca<br />
(Campos et al. 48), quina do Rio de Janeiro (Martius<br />
111), sobrasil (Sao Paulo, Edwall 23938). The label<br />
species.<br />
De Candolle (1830) validated Exostemaformosum<br />
Cham. & Schltdl. ex DC., under which he cited the<br />
two varieties a leprosum andfi laeve. The two variet-<br />
ies remained invalid because de Candolle did not cite<br />
the typical varieties of the species.<br />
Klotzsch (1846) separated this species from<br />
Exostema (as Exostemma) to establish the genus<br />
<strong>Rustia</strong>, under which he recognized three species. He<br />
validated R. pohliana by a description, and R. sel-<br />
lowiana by indirect reference to previously published<br />
description of E. formosum var. laeprosum; E. form-<br />
osum had already been validated by de Candolle<br />
(1830). Schumann (1889), in Flora Brasiliensis,<br />
treated these three species as synonymous, a conclu-<br />
sion with which I concur.<br />
The description of E. formosum var. laeve was<br />
prepared by Chamisso and Schlechtendal (1829) from
Systematic Treatment of the Genera Studied 89<br />
a Sellow collection from tropical Brazil. No Sellow ing-reflexed lobes, 1.8-2.2 cm long, white to creamspecimens<br />
of this species were encountered. There- white inside and outside, thick and fleshy when fresh;<br />
fore, I am here selecting a neotype for <strong>Rustia</strong>formosa, tube faintly funnelform, 8-10 mm long, ca. 4 mm wide<br />
the type species of the genus. Lemos 28678 (NY) was<br />
selected as neotype for R. formosa because it was<br />
at base and ca. 6 mm wide at the orifice, glabrous<br />
outside, with a white glabrous basal zone inside, with<br />
collected from eastern tropical Brazil, and corresponds a ring of ascendent-pilose hairs 6-7 mm from the base,<br />
most closely to the description ofE.formosum var. laeve. at the same point of the filament attachments at base<br />
of lobes; lobes 5,<br />
12. <strong>Rustia</strong> simpsonii Delprete, sp. nov. Type. Brazil.<br />
Rio de Janeiro: Petropolis, 21 Feb 1873 (fl),<br />
Glaziou 6564 (holotype, P; isotypes, BR, K,<br />
frag-F). Figs. 14G,H, 37A-C<br />
Arbores R.formosae similes sed ab ea corollis majoribus<br />
18-22 mm longis, 4-6 mm latis (nec 15-18 mm longis, 2-<br />
2.5 mm latis), corollae lobis secus medium glabris, intus<br />
juxta margines pilis glandulosis biseriatis provisi (nec basi<br />
albo-strigosis intus juxta margines dense papillosis) antheris<br />
?5 mm latis (nec 1-1.5 mm) basi extensioribus sphaeroidesis<br />
papillosis auctis (nec nudis) diversa.<br />
/2-2/3 of corolla length, 12-14 x 4-<br />
5 mm, narrowly triangular, glabrous outside, glabrous<br />
in central zone with two rows of minute (40x) glandular<br />
hairs inside, near the margins. Stamens 5, exserted<br />
because of spreading lobes, subequal, attached<br />
ca. 6 mm from the base of the tube; filaments ca. 7<br />
mm long, with a compact tuft of white ascending hirsute<br />
hairs at base; anthers cylindrical, narrowly oblong,<br />
ca. 9 x ca. 5 mm, dorsifixed near the base, base<br />
rounded, with papillose spheroid extension at base,<br />
smooth throughout. Pollen exine densely foveolate,<br />
minutely echinate. Style exserted, 20-23 mm long,<br />
glabrous; style branches narrowly ovate, ca. 2 mm<br />
long, stigmatic surface microscopically (40x) papillose.<br />
Capsule not seen. Seeds not seen.<br />
Trees; bark grayish. Leafy branchlets glabrous,<br />
terete; older branches rugose, grayish; lenticels many,<br />
punctiform to linear, 0.5-3 mm long. Stipules not seen<br />
(readily caducous), leaving a white linear scar. Leaves<br />
34-38 x 10-11 cm, L/W 3:1; oblanceolate, cuneate<br />
(BA = 20-25?) at base, obtuse at apex, sometimes with<br />
a deltoid acumen ca. 1 cm long; dark green above,<br />
yellowish green below, semi-coriaceous; drying ol-<br />
Distribution (see Fig. 39B) and ecology. Known<br />
only from the surroundings of Rio de Janeiro and<br />
Petropolis, an environment that is now mostly destroyed.<br />
This species is known only from two collections<br />
by Glaziou made in 1867 and 1873, and is possibly<br />
extinct. For notes on conservation biology of this<br />
species, see discussion under R. gracilis.<br />
ive-green, subcoriaceous; glabrous above, glabrous<br />
below; pellucid punctate; primary and secondary veins<br />
glabrous, prominent below, secondary veins 15-17<br />
each side, subparallel; tertiary veins starting sub-<br />
Specimens examined. BRAZIL. Rio DE JANEIRO: Rio<br />
de Janeiro [Corcovado and Serra da Estrella (Glaziou,<br />
1909)], 1867, Glaziou 1098 (BR[2]).<br />
parallel and reticulate in the center; petioles 41-45 mm<br />
long, 2.5-3 mm thick, terete or adaxially narrowly<br />
concave, thickened at base, but not pulvinate; domatia<br />
On the herbarium sheet Glaziou 6465 preserved<br />
at Field (isotype fragment-F), Donald Simpson wrote:<br />
absent. Inflorescences openly paniculate, pyramidal, This collection [was] cited by Schumann in Fl.<br />
with opposite or alternate, decussate branches; ca. 36 Bras. as R. formosa (C. & S.) Kl. but the flowcm<br />
long, basal branches to 20 cm long, in ca. 5 pairs, ers are extraordinarily thick. The anthers debasal<br />
portion of axis not branched ca. 8 cm long; ra- hiscing by 2 pores at the apex confirms its<br />
chis decussately compressed, narrowly rhombic or placement in <strong>Rustia</strong>, and the corolla lobes benarrowly<br />
elliptic in cross section, rachis and branches ing at least as long as the tube and calyx<br />
glabrous; flowers on tertiary branches alternate or in<br />
terminal lax cymules; distal bracts 6-10 x 3-5 mm,<br />
subtruncate places it in or near R.formosa, but<br />
the flower is nearly twice as thick as in most<br />
narrowly triangular to deltoid; bracteoles subtending<br />
flowers 3-5 x 2-4 mm, narrowly deltoid. Flowers<br />
pedicellate, pedicels 9-12 mm long, glabrous; hypanthium<br />
obovate to obconical, 4.5-7 x 4-6 mm, glabrous;<br />
flower buds cylindrical, pointed at apex. Calyx<br />
cupular, semi-membranous, with barely distinguishable<br />
lobes, 2-3.5 x 6-8 mm; lobes 5, ca. 1 x ca. 3 mm,<br />
R. formosa, and the corolla lobes are ca. twice<br />
as long as the tube (i.e. 2/3 the corolla length)<br />
while Schumann describe R. formosa as having<br />
corolla lobes 1/3 to 12 corolla length (i.e. less<br />
than or equalling the tube). <strong>Rustia</strong> gracilis has<br />
corolla lobes 2/3 the corolla length but flowers<br />
are very slender in that species.<br />
shallowly deltoid. Corolla short-tubular with spread-<br />
- D. Simpson,<br />
July 1977.
90 Flora Neotropica<br />
., ~<br />
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~~W<br />
,.~~~~~~~~~~~~~~~~~~~4<br />
.? '~~~~~~~P ~ ~ ~ ~ X<br />
..~~~~~~~~~~~~~~~~!Mq<br />
FI.37 uti ipsni(Gaiu 54 B n P yes.A Hbtofiforsecewt mtr la . pnflwr<br />
shwngodopnanhr adesetd eetiesyl.C.Dtilo nte bs hoigspeoi xenin
Systematic Treatment of the Genera Studied 91<br />
<strong>Rustia</strong> simpsonii is unique in the genus in having<br />
anthers with papillose spheroid extensions at base<br />
(Fig. 38C). It differs from the closely related R.<br />
formosa in having corollas 18-22 x 4-6 mm (vs. 15-<br />
18 x 2-2.5 mm), corolla lobes glabrous in central zone<br />
and with two rows of glandular hairs at margins inside<br />
(vs. white-strigose at base and margins densely<br />
papillose inside), anthers ca. 5 mm wide (vs. 1-1.5 mm)<br />
and with papillose spheroid extensions at base (vs.<br />
without such extensions), and calyx 2-3.5 x 6-8 mm<br />
(vs. 0.2-0.4 x 0.9-1.2 mm) (Fig. 38A-C).<br />
Donald Simpson has been an ephemeral, extremely<br />
accurate botanist, and the author of the only brief study<br />
specifically dedicated to <strong>Rustia</strong>. His acute observations,<br />
penciled on the herbarium specimens (and folders)<br />
at the Field Museum (F), have been very helpful<br />
in my revision of this genus, and I therefore honor him<br />
with the present appellation.<br />
13. <strong>Rustia</strong> angustifolia K. Schumann in Martius, Fl.<br />
Bras. 6(6): 262. 1889. Type. Brazil, Rio de Janeiro,<br />
Serra da Estrella, 1885 (fl), Glaziou 14937 (B*;<br />
photo-B at F, GH, MO, NY, VEN; lectotype, G,<br />
here selected; isolectotypes, BR, K, MA, R). The<br />
specimen Glaziou 14937 preserved at R has the<br />
date (manu Glaziou) of 25 August 1883, but on<br />
other duplicates the collecting date is solely 1885<br />
(manu Glaziou). Figs. 6B, 14F, 38B, 39A-E<br />
Shrubs 4-6 m tall, with slender branches usually<br />
terete; bark grayish-whitish. Leafy branchlets<br />
golden-tomentulose; older branches glabrate; lenticels<br />
absent. Stipules narrowly triangular but not acuminate,<br />
tomentulose outside, glabrous, with short<br />
colleters inside, 18-22 x 1.3-1.5 mm, light green,<br />
readily caducous, leaving a linear scar. Leaves (10-)<br />
11.5-33(-36.5) x section, rachis and branches minutely puberulent;<br />
flowers alternate on distal branches or in cymules;<br />
distal bracts 3-7 x 3-4 mm, narrowly lanceolate;<br />
bracteoles subtending flowers 1-2 x 0.5-0.7 mm,<br />
narrowly lanceolate, tomentulous outside, glabrous<br />
inside. Flowers pedicellate, pedicels 8-20 mm long,<br />
glabrous to minutely puberulent; hypanthium narrowly<br />
obconical, 2.5-5 x 2-3.5 mm, glabrous; flower<br />
buds cylindrical, acute at apex. Calyx small-cupular,<br />
truncate or with barely distinguishable lobes, 3-4 mm<br />
wide; lobes 5, 0.5-0.9 x 0.5-1 mm, shallowly triangular<br />
to deltoid, ciliolate. Corolla tubular with spreading<br />
lobes, 1.4-1.8 cm long, pinkish red to flesh-red<br />
outside, pearl-white inside, thick and fleshy when<br />
fresh; tube cylindrical 1.1-1.4 x 3-4 mm, glabrous<br />
outside, glabrous inside, with a tuft of sparse hairs at<br />
the sides of each filament attachment, 5-6 from the<br />
base (microscopically papillose); lobes 5, 1/5-l/4 of<br />
corolla length, 3-4 x 1-1.2 mm, ovate-triangular, glabrous,<br />
with the margins minutely papillose inside.<br />
Stamens 5, exserted because of spreading lobes, subequal,<br />
attached 2.5-5.5 mm from the base of the tube;<br />
filaments 1.8-2 mm long, barbate at base, with white<br />
to golden-yellow ascending hairs 0.1-0.2 mm long;<br />
anthers cylindrical, narrowly oblong, 5-8 x 0.6-0.9 mm,<br />
dorsifixed near the base, base rounded to subsagittate,<br />
microscopically (40x) papillose at very base. Pollen<br />
exine foveolate (non-echinate). Style exserted, 15-23<br />
mm long, glabrous; style branches narrowly triangular,<br />
0.9-1.1 mm long, glabrous, stigmatic surface<br />
smooth. Capsules obovoid to oblong-obovoid, apex<br />
subtruncate to shallowly hemispherical, 10-16 x 7-<br />
10 mm, sometimes with sparse punctiform lenticels,<br />
dark brown, glabrous throughout. Seeds 1.33-1.5 x<br />
0.4-0.73 mm.<br />
Distribution (Fig. 38B) and ecology. Understory<br />
shrubs in primary and secondary forests in the moun-<br />
2.5-4(-5.5) cm, L/W 3:1 to 5:1, tains surrounding Rio de Janeiro, Brazil. Flowering<br />
narrowly lanceolate, acute at base and at apex, some- specimens were collected from March to May. For<br />
times tapering to an acuminate apex to 1-1.5 cm long, conservation biology of this species, see discussion<br />
with a round tip; dark green above, pale green below, under R. gracilis (below).<br />
semi-coriaceous; drying olive-green, chartaceous;<br />
glabrous above and below, pellucid punctate; primary Specimens examined. BRAZIL. Rio DE JANEIRO: Rio<br />
and secondary veins glabrous, secondary veins (8-) de Janeiro, Serra da Estrella, old rd. Rio de Janeiro-<br />
10-17 each side, tertiary veins openly reticulate; pet- Petropolis, 23 May 1988 (fr), Gomes et al. 323 (HRB,<br />
ioles (10-)15-35 mm long, 1-2.5 mm thick, adaxial- RB[2]); Rio de Janeiro [Corcovado and Serra da Estrella<br />
ly concave to flattened, pulvinus present in young (Glaziou, 1909)], Mar 1872 (fl), Glaziou 2607 (K); Rio de<br />
leaves, 4-6 mm long, 1.5-2.5 mm Janeiro, 1891, Glaziou 18906 (K); Rio de Janeiro, Serra da<br />
thick, rarely absent;<br />
Estrella<br />
domatia absent. Inflorescences<br />
[near Mandioca (Glaziou, 1909)], 2 May 1880 (fl),<br />
pendulous, long-pe- Glaziou 12038 (F, G, P); in primary forest, Macah6, Maydunculate,<br />
paniculate with opposite decussate<br />
Jun 1832 (fl), Riedel & Luschnath 1062 (F, NY); Serra do<br />
branches; 11-20 x 8.5-16 cm, lateral branches 3-5<br />
Orgios, Alto Macah6, 1891, Glaziou 18306 (G); Porto Estrepairs,<br />
basal portion of axis not branched 6-8.5 cm lla, Dec 1823 (st), Riedel 66 (BR); Riedel 68 (BR); vic. of Manlong;<br />
rachis compressed, narrowly rhombic in cross- diocca, Martius 2964 (M), 2965 (M), 2966 (M), 2967 (M).
92 Flora Neotropica<br />
Ao<br />
FIG. 38. Distribution of four species of <strong>Rustia</strong>. A. R. formosa. B. R. simpsonii (stars), R. gracilis (diamonds) and R.<br />
angustifolia (circles).<br />
This species is unique in the genus in having long<br />
pedunculate, pendulous inflorescences; terminal fas-<br />
cicles of leaves at the tip of the vegetative branchlets;<br />
leaves narrowly lanceolate, three to five times as long<br />
as wide; corollas flesh-red outside and pearl-white<br />
inside, and corolla lobes one-fifth of the corolla length.<br />
Until few years ago, the last collection of R.<br />
angustifolia was dated 1891, and because of the in-<br />
creasing destruction of the natural environment in the<br />
surroundings of Rio de Janeiro (Dean, 1995), this<br />
species had been suspected to be extinct. Neverthe-<br />
less, the recent collection (1988) of Gomes et al. 323<br />
at Serra da Estrella, has proven the ongoing existence<br />
of this extremely rare and endangered species.<br />
14. <strong>Rustia</strong> gracilis K. Schumann in Martius, Fl. Bras.<br />
6(6): 263. 1889. Type. Brazil. Rio de Janeiro:<br />
Riverbanks of Rio Mandioca, Dec 1822-Jan 1823<br />
(fl), Riedel 424 (B*; lectotype, M, here selected;<br />
isolectotypes, BR[2], F, GB, GH, K, US).<br />
Figs. 5D, 14C,D, 38B, 40A-E<br />
3, 0<br />
Shrub 4-6 m tall, with slender branches; bark<br />
grayish-whitish. Leafy branchlets glabrous, slender,<br />
terete to tetragonous, grayish; lenticels absent.<br />
Stipules not seen (readily caducous), leaving a white<br />
linear scar. Leaves (11-)14-18 x 3.5-5.5 cm, L/W<br />
3:1 to 3.6:1, lanceolate to oblanceolate, acute (BA =<br />
18-25?) at base, acute at apex, tapering to an acuminate<br />
apex to 1-1.5 cm long, with a round tip; dark<br />
green above, pale green below, membranaceous; drying<br />
olive-green, chartaceous; glabrous above, glabrescent<br />
to sparsely pubescent below, pellucid punctate;<br />
primary and secondary veins glabrescent to sparsely<br />
pubescent, secondary veins 10-13 each side, tertiary<br />
veins openly reticulate; petioles 13-20(-25) mm long,<br />
1-1.5(-2) mm thick, subterete; pulvinus absent;<br />
domatia absent or a tuft of sparse hairs 0.5 mm wide,<br />
hairs white 0.5-0.6 mm long. Inflorescences openly<br />
paniculate with opposite decussate branches; 12-16<br />
x 11-13(-20) cm, lateral branches 3-5 pairs, basal<br />
portion of axis not branched 3.5-6.5 cm long; rachis<br />
compressed, elliptic or narrowly rhombic (rarely terete)<br />
in cross section, rachis and branches glabrous to<br />
minutely puberulent, extremely thin and delicate (in
Systematic Treatment of the Genera Studied 93<br />
FIG 3. Rsta ngutioli 143, ( fom laio B ,ioettp;BDfo id102 Y rmRee6)<br />
E.~~~~~~~~~r M"ba capsur ture l e.I<br />
FIG. 39. <strong>Rustia</strong> angustifolia (A from Glaziou 14937, BR, isolectotype; B-D from Riedel 1062, NY; E from Riedel 68).<br />
A. Habit of inflorescence with young and mature leaves. B. Flower bud. C. Stamen. D. Detail of anthers base, dorsal view.<br />
E. Mature capsule.
94 Flora Neotropica<br />
C cm<br />
[~~~~~~~~~~~~~~~~~~~~~~~~~5<br />
FIG. 40. <strong>Rustia</strong> gracilis (A-D from Riedel 424, BR; E from Glaziou 14893, BR). A. Habit of inflorescence with mature<br />
leaves. B. Abaxial view of blade showing pellucid punctation. C. Flower bud. D. Style. E. Mature capsule. E. Stamen,<br />
showing pilose base and open anther.<br />
w '.<br />
: .....:.: .......... :~ : : - . . . . . 5 :<br />
FI.40uti r rm cils(- R e?44 R rmGaiu183 R.A ai fnlrscc ihmtr<br />
showing pilos? bag~~~~~~ andopnanter
Systematic Treatment of the Genera Studied 95<br />
comparison to all other species); flowers alternate on<br />
distal branches or in cymules; distal bracts 2-3 x 1-<br />
1.5 mm, narrowly triangular; bracteoles subtending<br />
flowers 1-1.5 x 0.5-0.7 mm, narrowly triangular.<br />
Flowers pedicellate, pedicels 5-10 mm long, very thin<br />
and delicate, glabrous; hypanthium narrowly<br />
obconical, 2-2.5 x 1-1.5 mm, glabrous; flower buds<br />
clavate. Calyx narrowly cupular, truncate or with<br />
barely distinguishable lobes, 1.5-2.5 mm wide; lobes<br />
deltoid, 0.8-1 x 0.8-0.9 mm, ciliolate. Corolla tubular<br />
with narrowly spreading lobes, 1.2-1.8 cm long,<br />
white with pink-reddish base outside, white inside,<br />
thin and delicate when fresh; tube narrowly funnelform,<br />
5-7 mm long, ca. 1 mm wide at base and 2-2.5<br />
mm wide at the orifice, glabrous outside, glabrous and<br />
microscopically (40x) papillose inside, with a tuft of<br />
white-pilose hairs (0.8-1 mm long) at the sides of each<br />
filament attachment and base of lobes; lobes 5, 1'3-/2<br />
of corolla length, 5-7 x 0.9-1.1 mm, narrowly triangular,<br />
glabrous outside, with few sparse hairs and margins<br />
minutely papillose inside. Stamens 5, exserted<br />
because of spreading lobes, subequal, attached 5-6<br />
mm from the base of the tube; filaments 3-3.5 mm<br />
long, very thin and pendulous, barbate at base, with a<br />
tuft of white ascending hairs (0.8-1 mm long); anthers<br />
cylindrical, narrowly oblong, (2-)3-3.5 x 0.5-0.6 cially the threatened parks of Serra do Orgaos and<br />
Serra de Estrella.<br />
Specimens examined. BRAZIL. Rio DE JANEIRO: Rio<br />
de Janeiro, Serra de Estrella, Mandioca, 29 Mar 1884 (fl),<br />
Glaziou 14893 (BR, F, G, P); riverside close to Mandioca,<br />
Feb-Mar 1823 (fr), Riedel 367 (BR); E of Rio de Janeiro,<br />
Serra do Orgaos, Caneca, Dec 1974 (fl), Occhioni 6655 (US);<br />
Rio de Janeiro [Serra dos Orgaos, Alto Macahe (Glaziou,<br />
1909)], 1881, Glaziou 12036 (B*, BR, F, G[2], R); Rio de<br />
Janeiro, Glaziou 19437 (BR).<br />
<strong>Rustia</strong> gracilis is unique in the genus in having<br />
inflorescences with extremely thin and delicate<br />
branches (hence the specific epithet), and pedicels and<br />
corollas long and slender, which gives an overall frail<br />
appearance. This species could be confused with R.<br />
angustifolia, which is also an endemic of the mountain<br />
slopes surrounding Rio de Janeiro, but it differs<br />
from R. angustifolia in its openly paniculate inflorescences,<br />
slender whitish corollas (1-2 mm wide), and<br />
its membranous oblanceolate leaves. On the other<br />
hand, the inflorescences of R. angustifolia are rather<br />
stout, long-pedunculate, and densely corymbose distally;<br />
having thicker blood-red to flesh-red corollas<br />
(3-4 mm wide), and semi-coriaceous narrowly lanmm,<br />
ceolate leaves.<br />
dorsifixed at 0.2-0.3 mm from the base, base rounded,<br />
the base of two thecae often asymmetrical, microscopically<br />
(40x) papillose-echinate. Pollen exine<br />
Doubtful Species<br />
densely foveolate (non-echinate). Style exserted, 15-<br />
<strong>Rustia</strong> haitiensis Urban, Ark. Bot. 24A<br />
16(-18) mm long, glabrous, minutely papillose at dis-<br />
(4): 36. 1931.<br />
tal zone; style branches ovate, 0.4-0.5 mm<br />
Type. Haiti. Massif de la Hotte, western group,<br />
long, gla-<br />
Jeremie, left arm of<br />
brous, stigmatic surface glabrous; ovaries often<br />
Voldrogue River, 400 m, very<br />
rare, 22 Jul 1928<br />
parasitized by insects, causing an abnormal<br />
(fl-fr), Ekman H-10408 (B*; lecenlargement<br />
even before anthesis, becoming spherical and to<br />
totype, S, here selected).<br />
5-6 mm diam. but with no seeds inside. Capsules<br />
obovoid to oblong-obovoid, apex subtruncate, 6-8 x Shrub or small tree, glabrous. Leafy branchlets<br />
4-5 mm, dark brown, glabrous throughout. Seeds biconvex; older branches subterete; lenticels sparse.<br />
0.87-1.16 x 0.43-0.67 mm.<br />
Stipules narrowly triangular, acuminate, 6-8 mm<br />
long, readily caducous. Leaves petiolate, petioles 10-<br />
Distribution (Fig. 39B) and ecology. All the 20 mm long, 1.5-2 mm thick, adaxially concave to<br />
specimens examined have been collected either in flattened; blades 17-21 x 5-8 cm, L/W ca. 3:1; obo-<br />
Serra do Orgaos or Serra de Estrella, in the vicinity vate-elliptic, acute (BA<br />
of Rio de Janeiro. Flowering collections have been<br />
collected from December through March. Fruiting<br />
specimens were collected in February and March.<br />
Conservation status. This species is also extremely<br />
rare; most specimens were collected before<br />
1900, with a single collection from 1974 [Occhioni<br />
6655]. <strong>Rustia</strong> gracilis (with R. simpsonii and R.<br />
angustifolia) emphasize the extreme importance in<br />
protecting the remnants of the few patches left of<br />
Brazilian Atlantic forest (cf. Dean, 1995), and espe-<br />
= 27-32?) at base, widely<br />
obtuse to rounded at apex, chartaceous; glabrous<br />
above and below; pellucid punctate; primary and secondary<br />
veins glabrous, slightly depressed above,<br />
prominent below; secondary veins 11-14 each side;<br />
tertiary veins starting subparallel and reticulate in the<br />
center; domatia absent. Flowers, capsules, seeds, and<br />
pollen not seen.<br />
From Urban's (1931) original description: Inflorescences<br />
terminal, paniculate, with opposite to<br />
subopposite decussate lateral branches; ca. 25 cm<br />
long; rachis and branches glabrous, with lenticels;
96 Flora Neotropica<br />
distal bracts 2-3 mm long, lanceolate and acuminate,<br />
bracteoles subtending flowers ca. 1 mm long, deltoid,<br />
glabrous. Flowers pedicellate, pedicels ca. 2 mm long<br />
(in fruits). Capsule globose, to 3.5 mm long.<br />
14 Oct 1863 (fl), Isern 2121 (holotype, F) =<br />
Alibertia isernii (Standley) D. R. Simpson.<br />
Simpson (1976) transferred this species to<br />
Alibertia, a conclusion with which I concur.<br />
Distribution and ecology. Known only from the<br />
type, from the now-destroyed cloud forest on Massif <strong>Rustia</strong> longifolia Standley, Publ. Field Columbian<br />
de La Hotte in southern Haiti. From the description, Mus. Nat. Hist., Bot. Ser. 8: 159. 1930. Type.<br />
the specimens were collected in flowering stage in Colombia. Caqueti: Between Santa Marta and Mar-<br />
July, with some old capsules persisting on branches. sella, Aug 1926 (fr), Woronow & Jusepczuk 6426<br />
In the protologue Urban (1931) stated: "Rarissima (holotype, LE-n.v.; isotypes, F, LE-n.v.). =Dolichoet<br />
parca, Jul cum inflor. valde juvenilibus et capsulis delphys chlorocrater K. Schumann & K. Krause.<br />
vetustis. Obs. An re vera hujus generis, antherae adhuc<br />
ignotae demonstrabunt. Habitus et folia glandulosopunctata<br />
ad genus <strong>Rustia</strong>m spectant" [... Even though <strong>Rustia</strong> rosea (Klotzsch & Karsten ex Walpers) K.<br />
it was not possible to see the anthers, habit and pellu- Schumann in Engler & Prantl, Nat. Pflanzenfam.<br />
cid-punctate leaves show that it belongs to <strong>Rustia</strong>]. 4(4): 18. 1891. Basionym: Henlea rosea Klotzsch<br />
Only one sterile specimen (Ekman H-10408, S) of & Karsten ex Walpers, nomen., Ann. Bot. 2: 788.<br />
this species was found. The above description is based 1852. Type. Pro. syn. = Ladenbergia muzonensis<br />
on personal observations, to which I added the transla- (Goudot) Standley (1930a).<br />
tion of Urban's (1931) original description. The leaves<br />
Schumann (1891) erroneously treated H. rosea as<br />
pellucid punctation is not as obvious as in all the spe- a valid species and as the type species of Henlea. As<br />
cies of this genus, suggesting that this species prob- noted by Standley (1930a), H. rosea was never valably<br />
does not belong to <strong>Rustia</strong>. Until further collecidly<br />
described. Walpers (1852) treated this taxon as<br />
tions of flowering and fruiting material can be made the<br />
valid and listed it as synonymous with Cascarilla<br />
identity of this taxon is destined to remain doubtful.<br />
muzonensis [= Ladenbergia muzonensis], an interpretation<br />
with which I agree.<br />
Excluded Species<br />
Henlea muzonensis (Goudot) Klotzsch & H. Karsten<br />
ex Walpers, Ann. Bot. Syst. 2: 788. 1852. Cinchona<br />
muzonensis Goudot, Phil. Mag. New Ser.<br />
3: 132. 1828. Cascarilla muzonensis (Goudot)<br />
Weddell, Ann. Sci. Nat. 3(10): 11. 1848. Cascarilla<br />
[sect.] muzonia Weddell, Hist. Nat. Quinquinas<br />
90. 1849. Buena muzonensis (Goudot) Weddell,<br />
J. Linn. Soc. 11: 187. 1869. Type. Colombia.<br />
Santander: Near Muzo, Goudot s.n. (lectotype, P,<br />
selected by Andersson, 1994). = Ladenbergia<br />
muzonensis (Goudot) Standley (1930a).<br />
<strong>Rustia</strong>ferruginea Standley, Contr. U.S. Natl. Herb.<br />
18: 142. 1916. Type. Panama. Col6n: Along Rio<br />
Fat6, in forest or thickets, 10-100 m, Aug 1911<br />
(fl), Pittier 4201 (holotype, US). = <strong>Rustia</strong> pauciflora K. Schumann ex Solereder, nomen.,<br />
Ber. Deutsch. Bot. Ges. 8: (99). 1890. <strong>Rustia</strong>pauciflora<br />
K. Schumann, in herb. Type. Tobago. Forest<br />
of Cremorue River, 360 m, Nov 1889 (fr), Eggers<br />
5812 (holotype, M; isotypes, K, US).<br />
Bathysa (?).<br />
This species does not belong to <strong>Rustia</strong>, but because<br />
of the depauperate conditions of the unicate specimen,<br />
it is impossible to attribute this species to a genus with<br />
certainty.<br />
<strong>Rustia</strong> isernii Standley, Publ. Field Mus. Nat. Hist.<br />
Bot. Ser. 22: 212. 1940. Type. Peru. Chachamayo,<br />
= <strong>Tresanthera</strong><br />
condamineoides Karsten var. condamineoides.<br />
For further details on the taxonomic history and<br />
identity of this taxon see discussion under <strong>Tresanthera</strong><br />
condamineoides Karsten var. condamineoides.<br />
TRESANTHERA<br />
2. <strong>Tresanthera</strong> H. Karsten, Fl. Columb. 1: 37. pl 19.<br />
1858; Hooker in Bentham & Hooker, Gen. P1. 2(1):<br />
44. 1873; Baillon, Hist. P1. 7: 471. 1880; Karsten,<br />
Bot. Jahrb. Syst. 8: 356. 1887; Schumann in Engler<br />
& Prantl, Nat. Pflanzenfam. 4(4): 19. 1891;<br />
Solereder, Ber. Deutsch. Bot. Ges. 8: 99. 1890;<br />
Solereder, Bull. Herbier Boissier 1: 280-181.<br />
1893; Standley, Publ. Field Columbian Mus., Bot.<br />
Ser. 7: 371. 1931; Steyermark, Bol. Soc. Venez.<br />
Cienc. Nat. 25: 248. 1964; Steyermark in Lasser &<br />
Steyermark, Fl. Venezuela 9(1): 256-261. 1974. Type<br />
species. <strong>Tresanthera</strong> condamineoides Karsten.
Systematic Treatment of the Genera Studied 97<br />
Shrubs to single-stemmed trees, terminal large pyramidal shrubs (rarely medium-sized trees),<br />
branches with soft pith in the center, with leaves clus- and has never been found to develop buttresses. Its<br />
tered at apex of each branch; bark grayish-pale brown. wood is usually soft and of no particular use, its bark<br />
Stipules interpetiolar, free at base, overlapping in bud, is smooth and grayish, and its seeds are minute, horinarrowly<br />
triangular, readily caducous. Leaves peti- zontally inserted, non-winged, flattened, and apically<br />
olate, elliptic to oblanceolate, obtuse or short-acumi- truncate (as in some species of <strong>Rustia</strong>).<br />
nate at apex; pellucid-punctate; petioles thickened at <strong>Tresanthera</strong> is sometimes confused with <strong>Rustia</strong>,<br />
base, not pulvinate, becoming corky in older leaves; but the latter has terminal poricidal anthers, anther<br />
domatia absent. Inflorescences terminal, solitary, connective tissue not as evident, smaller fruits, and<br />
paniculate or single thyrsoid axis; lateral branches variously shaped seeds.<br />
(when present) cymose or thyrsoid. Flowers<br />
protandrous; hypanthium obconical, acute at base.<br />
Taxonomic History<br />
Calyx reduced to a wavy margin or with minute lobes,<br />
persistent. Corolla campanulate with patent to re- <strong>Tresanthera</strong> was founded by Karsten (1858), deflexed<br />
lobes, fleshy when fresh, yellowish orange to riving the name from the Greek tresis (perforation)<br />
orange-red outside, yellowish white to cream-white and antheros (anther) to refer to its porate anthers.<br />
inside; tube basally ventricose with medial constric- Karsten produced a very detailed description of the<br />
tion, glabrous; lobes 5; aestivation valvate-reduplicate genus and of T. condamineoides, the type species. This<br />
with contact zone. Stamens 5, alternate to corolla genus was included (with Henlea) by Hooker (1873)<br />
lobes, half-exserted, attached near the base of corolla in <strong>Rustia</strong>. Nevertheless, Karsten (1887) maintained<br />
tube; anthers convex toward the center, acute at apex, them as separate and published a brief description on<br />
yellow, dorsifixed near the base, dehiscing by a com- their basic morphological differences. After a series of<br />
mon lateral triangular pore below the apex. Pollen unpublished and invalid names, Solereder (1893)<br />
tricolporate, exine reticulate. Style exserted; style legitimized the taxon T. pauciflora K. Schum. ex Soler.<br />
branches membranous, ovate, reflexed at maturity. Steyermark (1964a) published <strong>Tresanthera</strong><br />
Ovary 2-celled, obconical, glabrous, placentation thyrsiflora, basing its description only on portions of<br />
axile; ovules many in each locule, horizontally in- an inflorescence. Later, Steyermark (1974) stated that<br />
serted; immature fruits oblong-elliptic, reddish green the three species recognized by him could as well be<br />
and semi-carose when fresh. Capsules woody, nar- conspecific, suggesting that subsequent collections<br />
rowly obovoid; disk obviously exceeding the calyx; might better clarify their identity. In the present treatdehiscing<br />
loculicidally, disk septicidal dehiscence ment <strong>Tresanthera</strong> is treated as monotypic, with two<br />
rarely present in old capsules. Seeds compressed, infraspecific categories, and ranging from the coastal<br />
horizontal, scobiform, with reticulate exotesta. forests of Venezuela to the island of Tobago.<br />
<strong>Tresanthera</strong> is unique in the <strong>Rubiaceae</strong> in having<br />
poricidal anthers opening by a common lateral<br />
1. <strong>Tresanthera</strong> condamineoides H. Karsten, Fl.<br />
pore<br />
of triangular shape, just below the apex. The common<br />
Columb. 1: 37, pl. 19. 1869.<br />
lateral pore is the result of the two convex thecae that<br />
Figs. 3E, 4D, 7A-C, 15A,B, 42, 43, 44<br />
converge apically so that the anther becomes pointed. Shrubs to<br />
The connective tissue in the dorsal portion of the antrees,<br />
to 15-20 m tall, to 30 cm dbh,<br />
thers (and between the two thecae) is<br />
usually single-stemmed frondose trees; bark grayish.<br />
quite evident,<br />
black, and with irregular margins. The closely related<br />
Leafy branchlets glabrous, with soft pith in the cen-<br />
<strong>Rustia</strong> also has poricidal anthers, but<br />
ter,<br />
these<br />
easy to break, terete to subquadrangular, grassopen<br />
by<br />
two terminal pores (the thecae not converging). Both<br />
green; older branches terete, pale brown. Stipules<br />
<strong>Tresanthera</strong> and <strong>Rustia</strong> are exceptional in the Rubinarrowly<br />
triangular, acuminate, 3-7 cm long, ciliolate,<br />
aceae in having leaves with<br />
glabrous to sparsely puberulent outside, glabrous, with<br />
"pellucid glands." a basal area of colleters inside<br />
<strong>Tresanthera</strong> has protandrous (the stigma is<br />
(Fig. 3E), dark green,<br />
receptive<br />
only after the anthers are dried) flowers, with<br />
readily caducous, leaving a scar encircling the stem<br />
1.5-2.5 mm wide. Leaves 28-71 x 15-25 cm, L/W<br />
valvate-reduplicate aestivation, erect or nodding, and<br />
do not produce any particular fragrance. Because of<br />
1.5:1-3:1; elliptic, oblong, to oblanceolate; acute,<br />
obtuse to truncate at<br />
their particular morphology, the flowers are most<br />
base, acute to obtuse at apex,<br />
sometimes short deltoid-acuminate; blade dark<br />
probably buzz-pollinated by bees (as in some<br />
green<br />
species<br />
of <strong>Rustia</strong>), but their floral visitors/pollinators have<br />
above, pale green below, semi-coriaceous; drying olnever<br />
been observed. <strong>Tresanthera</strong> usually occurs as<br />
ive-green, stiff-chartaceous; glabrous above and be-
98 Flora Neotropica<br />
low, pellucid-punctate; primary and secondary veins<br />
glabrous, prominent below, secondary veins 16-25<br />
each side; tertiary veins starting subparallel and openly<br />
reticulate in the center, faintly evident above, very<br />
evident below; petioles 3-7(-10) cm long, 3-7 mm<br />
thick, terete to adaxially concave, glabrous. Inflorescence<br />
single pauciflorous axes to thyrsoid-panicles,<br />
to 100 cm long; lateral branches (when present) 1-2<br />
pairs, opposite to subopposite, (5-)15-28 cm long;<br />
basal portion of axis not branched 14-33 cm long;<br />
rachis terete to obtuse-compressed, rachis and<br />
branches glabrous; flowers on rachis and lateral<br />
branches solitary or in several cymules subtended by<br />
a bracteole; bracts subtending basal branches leaf-like,<br />
semi-caducous, to 17 x 7 cm, lanceolate; distal bracts<br />
subtending the cymules and the flowers narrowly triangular,<br />
4-15 x 3-7 mm. Flowers long-pedicellate,<br />
pedicels 10-25 mm long, glabrous; hypanthium<br />
obconical, 5-10 x ca. 3 mm, glabrous to puberulent.<br />
Calyx reduced to a minutely undulate (crenulate)<br />
margin, 0.5-1.5 x 4-6 mm, glabrous. Corolla campanulate<br />
with reflexed lobes, 20-30 mm long, orangered<br />
outside, yellowish white to greenish white inside,<br />
semi-fleshy to subcoriaceous when fresh; tube with<br />
medial constriction, 1.1-1.8 x 5-9 mm, glabrous outside<br />
and inside, without internal ring of hairs; lobes<br />
5, '/3-/2 of corolla length, 7-12 x 6-8 mm, ovate to<br />
deltoid, acute at apex, glabrous outside, microscopically<br />
(40x) papillose inside. Stamens 5, partially exserted,<br />
equal, attached 5-7 mm from the base of the<br />
tube; filaments 4-6 mm long, enlarged and flattened<br />
at base, glabrous; anthers convex toward the center,<br />
lanceolate, acute at apex, (8-)10-13 x their general shape, but differ from the latter two in<br />
having anthers strongly convex toward the center.<br />
Also, the anthers in <strong>Rustia</strong> are apically poricidal, while<br />
in Condaminea they dehisce by lateral slits.<br />
The detailed original description by Karsten (1858)<br />
and its hand-colored illustration provide sufficient<br />
information to grasp the identity of T. condamineoides.<br />
In the Vienna Herbarium (W) two Karsten<br />
specimens of <strong>Tresanthera</strong> are preserved: one with a<br />
flowering inflorescence and a cymule with old capsules,<br />
bearing the label "<strong>Tresanthera</strong> condamineoides<br />
Krst. - Cumbre de Valencia - Pt Cabello - Dr<br />
Karsten" (unknown handwriting); the other with a<br />
terminal branch with two fully expanded leaves, and<br />
two cymules with old capsules, bearing the label<br />
"<strong>Tresanthera</strong> condamineoides Krst - Cumbre de<br />
Valencia - Pt Cabello" (unknown handwriting), with<br />
"H. Karsten. Dr." embossed on the label, and the annotation<br />
"! Sch." (in K. Schumann handwriting). I<br />
have selected the former specimen (having flowers in<br />
anthesis and mature capsules) as the lectotype of T.<br />
condamineoides, and the latter as isolectotype. Additional<br />
isolectotypes might also be found at LE (Tryon,<br />
1963), but I did not examine material from that institution.<br />
A dubious fragment has been found at Field<br />
Museum (F), with a few anthers and a small piece of<br />
leaf; on the envelope is written "<strong>Tresanthera</strong> Condaminensis<br />
[orth. var. Karst. - Karsten." No original labels<br />
are attached to the herbarium sheet, and the identity<br />
of the fragments preserved at F remain doubtful.<br />
The label of Eggers 5812 (holotype of T. pauciflora)<br />
preserved at Munich (M) bears the following<br />
1.3-2.8 mm, annotation in Schumann's handwriting: "Certe non!<br />
base rounded and bilobed, smooth throughout, dehisc- potius <strong>Rustia</strong> spec. nov., <strong>Rustia</strong> pauciflora m."<br />
ing by common lateral triangular pore, 2.5-4 mm Solereder (1890) simply cited the name <strong>Rustia</strong>paucibelow<br />
the apex. Pollen exine foveolate (non-echinate). flora from Schumann's herbarium annotation, but<br />
Style exserted, terete, 20-28 mm long, glabrous; style without description, thus producing an invalid specific<br />
branches elliptic to narrowly ovate, 2.5-3 mm long, epithet. Schumann (1891), in Die Natirlichen Pflanzenstigmatic<br />
surface smooth; immature fruits elliptic, familien, simply listed "T. pauciflora (Solered.) K.<br />
reddish green, and semi-camose when fresh. Capsules Schum.," also without description, causing this epinarrowly<br />
elliptic to obovoid, 15-24 x 8-12 mm, dark thet to remain a nomen nudum. Finally, Solereder<br />
brown, sometimes with few lenticels, glabrous; disk (1893) published a detailed description of T. pauciobviously<br />
exceeding the calyx, black; disk septicidal flora, validating Schumann's specific epithet and statdehiscence<br />
present in some old capsules. Seeds 1-1.5 x ing that the single specimen to which he referred was<br />
0.33-0.53 mm.<br />
"Eggers 5812 (Herb. Monac. [M!])", where the holotype<br />
is still preserved.<br />
<strong>Tresanthera</strong> condamineoides is a rare species in- The main characters that Steyermark (1974) used<br />
creasingly threatened with extinction by human pres- to distinguish the three <strong>Tresanthera</strong> species recogsure.<br />
It is easily distinguishable within the family by nized by him were corolla color, inflorescence morits<br />
laterally poricidal anthers and pellucid-punctate phology, and leaf shape. Examining complete sets of<br />
leaves (with <strong>Rustia</strong>). The flowers of T. condamine- various collections, I noticed that the corolla color of<br />
oides vaguely resemble those of <strong>Rustia</strong> occidentalis <strong>Tresanthera</strong> ranges from yellowish orange to orangeand<br />
Condaminea corymbosa in being reddish outside red outside, and from yellowish white to greenish<br />
and greenish white inside, fleshy when fresh, and in white (to pearl-white) inside (as in Condaminea
Systematic Treatment of the Genera Studied 99<br />
I _ n .*<br />
s<br />
f<br />
4<br />
*?<br />
^p , a<br />
II ?11<br />
FIG. 41. Distribution of <strong>Tresanthera</strong> condamineoides var. condamineoides (circles) and T. condamineoides var.<br />
thyrsiflora (star).<br />
corymbosa), making this character taxonomically insignificant.<br />
The inflorescence morphology of <strong>Tresanthera</strong><br />
varies from pauciflorous to sparsely cymose within<br />
the same set of collections (probably from single individuals),<br />
and is densely thyrsoid only in Bernardi<br />
5829. In <strong>Rustia</strong>, the closest genus to <strong>Tresanthera</strong>, I<br />
have observed subsecund inflorescences and erect<br />
inflorescences in the same individual. I have not observed<br />
individuals of <strong>Tresanthera</strong> in the field, but it<br />
is expected that the same is true for this genus, and<br />
therefore the secundiflorous or erect inflorescences are<br />
not considered a reliable taxonomic character.<br />
The leaves of <strong>Tresanthera</strong> are usually oblanceolate<br />
(but elliptic-oblong in Bernardi 5829), and the leaf<br />
bases vary from acute to obtuse (but are truncate in<br />
Bernardi 5829). The fruits of <strong>Tresanthera</strong> change in<br />
shape as they go through maturation, early on elliptic-ovoid<br />
and becoming narrowly obovoid with age.<br />
After proposing the three <strong>Tresanthera</strong> species,<br />
Steyermark (1974) stated that they might be reduced<br />
to one or two species as further collections accumulated.<br />
Steyermark did not study the types housed in<br />
the European herbaria, which are crucial for understanding<br />
the identity of these taxa. Additional recent<br />
collections (and their accurate label data) throughout<br />
its range have helped me to unravel the confused taxonomy<br />
of <strong>Tresanthera</strong>. I conclude that the characters<br />
used by Steyermark (1964, 1974) to separate the three<br />
taxa are of little diagnostic merit, and this genus is<br />
here treated as monotypic: T. thyrsiflora is reduced<br />
to varietal rank, and T. pauciflora is placed in synonymy<br />
with var. condamineoides.<br />
Key to the varieties of<br />
<strong>Tresanthera</strong> condamineoides<br />
1. Leaf blades oblanceolate to obovate, acute at<br />
base, L/W ca. 3:1; inflorescence a sparsely<br />
flowered thyrsoid panicle, with a single axis<br />
or with a pair of basal branches; corolla campanulate<br />
with reflexed lobes, ventricose at base,<br />
9-10 mm wide at base ......... la. var. condamineoides<br />
1. Leaf blades elliptic to oblong, truncate at base,<br />
L/W 1.5:1 to 2.5:1; inflorescence a densely<br />
flowered thyrsoid panicle, with a pair of long<br />
basal branches; corolla subcampanulate with<br />
patent lobes, conical at base, 4-5 mm wide at<br />
base ................................................. b. var. thyrsiflora<br />
la. <strong>Tresanthera</strong> condamineoides H. Karsten var.<br />
condamineoides. <strong>Tresanthera</strong> condamineoides H.<br />
Karsten, Fl. Columb. 1: 37, pl. 19. 1869. Type.<br />
Venezuela. Aragua: Puerto Cabello, Cumbre de<br />
Valencia (near the village of Campanero), hot wet<br />
forest on the slopes of coastal Cordillera, 100-200<br />
m, 1844-1847[?] Karsten s.n. (B*; lectotype, W,<br />
here selected; isolectotypes, W, LE?; dubious<br />
isolectotype, fragment at F).<br />
Figs. 3E, 4D, 7C, 15B, 41, 42A-F<br />
<strong>Tresanthera</strong> pauciflora K. Schumann ex Solereder, Bull.<br />
Herb. Boissier 1: 280. 1893. <strong>Tresanthera</strong> pauciflora<br />
K. Schumann, nom. nud., in Engler & Prantl, Nat.<br />
Pflanzenfam. 4(4): 19. 1891. <strong>Rustia</strong> pauciflora K.<br />
Schumann ex Solereder, nom. nud., Ber. Deutsch.<br />
Bot. Ges. 8: 99. 1890. Type. Tobago. Morue d'Or,<br />
forest of Cremorue River, 360 m, Nov 1889 (fl),<br />
Eggers 5812 (holotype, M; isotypes, GB, K, US).
100 Flora Neotropica<br />
FIG. 42. <strong>Tresanthera</strong> condamineoides var. condamineoides (A from Eggers 5812, US, isotype; B from Steyermark et<br />
121534, NY; C-E from Sandwith 1757, K; F from Fendler 2366, K). A, B. Inflorescence variation. A. Pauciflorous inflo-<br />
rescence with immature fruits, and mature leaf. B. Multiflorous inflorescence with mature fruits. C. Open flower at anthe-<br />
sis with dehished anthers, before style elongation. D. Stamen, adaxial view. E. Stamen, lateral view. F. Mature capsule.<br />
A
Systematic Treatment of the Genera Studied 101<br />
Leaf blades oblanceolate to obovate, acute at base,<br />
L/W ca. 3:1. Inflorescences single pauciflorous axes<br />
to sparsely flowered thyrsoid panicles with a pair of<br />
basal branches; bracts subtending basal branches nar-<br />
rowly triangular. Corollas campanulate with reflexed<br />
lobes, 25-30 mm long; tube ventricose at base, 13-<br />
18 x 9-10 mm (at base); lobes 13-/2 of corolla length,<br />
7-10 mm long. Stamens with filaments 5-6 mm long;<br />
anthers lanceolate, acute at apex, 11.5-12.5 x 2.2-2.5<br />
mm. Pollen exine regularly foveolate. Style 20-22<br />
mm long; immature fruits ovoid-elliptic. Capsules<br />
narrowly obovoid, 18-24 x 8-12 mm. Seeds 1.33-<br />
1.5 x 0.37-0.53 mm.<br />
Distribution (Fig. 41) and ecology. Lowland rain<br />
forests and cloud forests of medium elevations, in<br />
association with Gyranthera caribensis and Iriartea<br />
fusca, 100-1400 m, along the Coastal Cordillera of<br />
Venezuela and the forested slopes of Tobago. Flow-<br />
ering specimens were collected in August, Novem-<br />
ber, and December. Specimens with mature capsules<br />
have been collected in July and December.<br />
pauciflorous to multiflorous panicles, with lateral<br />
cymules ranging from single-flowered to few flowered;<br />
all the possible intergradations were found<br />
throughout the specimens examined. Unfortunately,<br />
in Bernardi 5829 (var. thyrsiflora) only young capsules<br />
are available, so to it was impossible to reliably<br />
compare the fruits of these two varieties.<br />
lb. <strong>Tresanthera</strong> condamineoides H. Karsten var.<br />
thyrsiflora (Steyermark) Delprete, var. et stat.<br />
nov. <strong>Tresanthera</strong> thyrsiflora Steyermark, Bol. Soc.<br />
Venez. Cienc. Nat. 25: 248. 1964. Type. Venezuela.<br />
Miranda: Merida, rain forest of Guatopo,<br />
400-600 m, small tree of the lower level of canopy,<br />
flowers red-brown, 23 Nov 1956 (fl), Bernardi<br />
5829 (holotype, NY[2]; isotypes, K[3], NY,<br />
VEN[2]). Figs. 7A,B, 15A, 41, 43A-E<br />
Leaf blades elliptic to oblong, truncate at base; L/<br />
W 1.5:1 to 2.5:1. Inflorescences densely flowered<br />
thyrsoid panicles, with a pair of long basal branches;<br />
Specimens examined. VENEZUELA. ARAGUA: Near bracts subtending basal branches foliose, 10-15 x 4-<br />
Colonia Tovar, 1856-1857 (fr), Fendler 2366 (frag-F, GH, 8 cm. Corollas subcampanulate with patent lobes, 18-<br />
K). DISTRITO FEDERAL: N slope of Cordillera de la Costa, S 25 mm long; tube conical at base, 10-13 x 4-5 mm<br />
of Camuri grande, Topo Tacamahaco [Tacamanaco], wide; lobes ca. '/3 of corolla length, 9-10 x 6.5-7 mm.<br />
quebrada Rio Grande, 9 km E of Tanques de la Electricidad Stamens with filaments 4-5 mm long; anthers lande<br />
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-<br />
(MY, VEN[3]); between Fila Maestra and Rio Caruao, San ine irregularly foveolate. Style 22-28 mm long; im-<br />
Jorge de Caruao, 900-1150 m, 10?32'N, 66?22'W, 16-19 Apr mature fruits ovoid-elliptic. Capsules not seen. Seeds<br />
1993 (fr), Meyer & Llamosa 3627 (VEN); N slope, above<br />
1-1.2 x 0.33-0.5 mm.<br />
Naiguata, quebrada Rio Camuri Grande, 800-850 m,<br />
10034'N, 66?42'W, 11-12 Apr 1992 (fr), Meyer et al. 2039<br />
(VEN). SUCRE: Quebrada Nivardo, affluent of Rio Local name. Venezuela: cambur6n<br />
Oscuro,<br />
(Bernardi 5829).<br />
above Mundo Nuevo, W of Cerro Humo, Paria peninsula,<br />
700-750 m, 7 Aug 1966 (fl), Steyermark & Rabe 96163 (K, Distribution (Fig. 41) and ecology. This variety<br />
VEN[2]); Dto. Cagigal, Peninsula de Paria, Rio Cumana, is known only from a single collection (Bernardi<br />
below trail between El Paujil and El Brasil, 10?40'N, 5829) made in the rain forests of Guatopo (State of<br />
62?43'W, 850 m, 21 Feb 1980 (fr), Steyermark et al. 121534 Miranda, Venezuela). On its large inflorescence, col-<br />
(NY, VEN); Peninsula de Paria, Cumbre de La Estrella, W lected in November, are present floral buds, flowers<br />
of Manacal, N of El Paujil, 800-850 m, 10?40'N, 62041'W, in anthesis, and immature ovoid fruits.<br />
17 Oct 1984 (fl), Knapp & Mallet 6764 (BH).<br />
TOBAGO: Rd. Roxborough-Parlatuvier, 800-1300 m,<br />
The<br />
14 Oct 1937 (fl), Sandwith 1757 (K[2]); Tobago Forest Re- original description of <strong>Tresanthera</strong> thyrsiflora<br />
serve, along Roxborough-Parlatuvier, between 4 & 5 mi was based on the terminal portion of the inflorescence,<br />
posts, 1200-1400 m, 20 Aug 1959 (fl), Webster et al. 9805 not the complete inflorescence, and was described by<br />
(A, CAS, DAV); Easterfield, 20 Dec 1912 (fl-fr), Broadway Steyermark (1964a) as a closely flowered thyrse.<br />
3076 (F[2], G[6], GH, NY[2], P, US[2]).<br />
Having available the complete set of Bernardi 5829,<br />
I was able to match the lateral branches of T. thyrsi-<br />
Variety condamineoides differs from var. thyrsi- flora, realizing that its inflorescence is instead a longflora<br />
in having oblanceolate, basally acute (vs. oblong, pedunculate, densely flowered, thyrsoid panicle, with<br />
basally truncate) leaf blades (Fig. 42A), larger and less a pair of long basal branches (Fig. 43A). In addition,<br />
constricted corollas with reflexed (vs. sub-erect) lobes, the flowers of Bernardi 5829 are clustered in short<br />
and larger anthers. The inflorescences of var. cymules as illustrated for T. condamineoides (in<br />
condamineoides are quite variable, ranging from Karsten, Fl. Columb. 1, pl. 19) and orange-red (as in
102 Flora Neotropica<br />
FIG. 43. <strong>Tresanthera</strong> condamineoides var. thyrsiflora (Bernardi 5829, K and NY). A. Habit of inflorescence and mature<br />
leaf. B. Open flower at anthesis with dehisced anthers, before style elongation. C. Stamen, adaxial view. D. Stamen,<br />
lateral view. E. Immature capsule.
Systematic Treatment of the Genera Studied 103<br />
T. condamineoides); the calyx crenulations of<br />
Bernardi 5829, to which Steyermark (1964a) referred<br />
in the protologue, are similarly illustrated for T.<br />
condamineoides (in Karsten, Fl. Columb. 1, pl. 19).<br />
CONDAMINEA<br />
3. Condaminea A. P. de Candolle, Prodr. 4(4): 402.<br />
Sep 1830; Ruiz & Pavon, Fl. Peruv. 2: 48-49, figs.<br />
188, 190, 198, 199. 1799; Schumann in Martius,<br />
Fl. Bras. 6(6): 257. 1889; Poiret in Lamarck,<br />
serted because of spreading lobes, subequal; anthers<br />
dorsifixed near the base or at medial zone, opening<br />
inward by longitudinal splits. Pollen tricolporate,<br />
exine reticulate. Style exserted; style branches membranous,<br />
ovate, reversed at maturity. Ovary 2-celled,<br />
obconical to obovate, glabrous, placentation axile;<br />
ovules many in each locule, horizontally inserted;<br />
immature fruits green and semi-fleshy when fresh.<br />
Capsules oblong, obovate to turbinate, woody; disk<br />
septicidal dehiscence absent even in old capsules.<br />
Seeds irregularly 3-4-angular, trapezoid, compressed,<br />
testa reticulate.<br />
Encycl. Suppl. 3: 570-571. 1814; Kunth in<br />
Humboldt, Bonpland & Kunth, Nov. Gen. & Spec.<br />
3: 311. 1818; A. Richard, Mem. Fam. Rubiacees<br />
198. Dec 1830; Endlicher, Gen. P1. 553 (N. 3262).<br />
1837; Meisner, PI. Vasc. Gen. PI. 158 (114). 1838;<br />
Hooker in Bentham & Hooker, Gen. PI. 2: 12, 44-<br />
45. 1873; Baillon, Hist. PI. 7:471. 1880; Standley,<br />
Publ. Field Columbian Mus., Bot. Ser. 7: 26, 203,<br />
278. 1930, 1931; Standley, Field Mus. Nat. Hist.,<br />
Bot. Ser. 13: 55-57. 1936; Steyermark in Lasser<br />
& Steyermark, Fl. Venez. 9(1): 241-244. 1974;<br />
Dwyer, Fl. Panama, Ann. Missouri Bot. Gard. 67:<br />
115-117. 1980; Burger & C. M. Taylor in Burger,<br />
Fl. Costar., Fieldiana, Bot. n.s. 33: 107-108. 1993;<br />
Delprete in Harling & Andersson, Fl. Ecuador (in<br />
press). 1999. Type species. Condaminea corymbosa<br />
(Ruiz & Pav6n) A. P. de Candolle.<br />
Shrubs to small trees, much-branched shrubs,<br />
rarely single-stemmed treelets; bark pale brown to<br />
grayish. Stipules interpetiolar and intrapetiolar, 4<br />
(rarely 2) at each node, connate at base, overlapping<br />
in bud, foliose, lanceolate, acuminate, with one central<br />
vein with strongly ascending secondary veins and<br />
several parallel veinlets departing from base, caducous.<br />
Leaves sessile to short-petiolate, elliptic to lanceolate<br />
to oblanceolate, rounded, cordate, to auriculate<br />
at base, acute to obtuse, usually acuminate at apex,<br />
dark green and waxy above, flat to undulate, thickfoliaceous<br />
to very coriaceous; petioles (when present)<br />
short and stout; domatia absent. Inflorescences terminal,<br />
solitary, long pedunculate, open corymbose<br />
panicle with a pair of opposite branches terminating<br />
in cymules. Flowers protandrous, hypanthium<br />
obconical to obovate. Calyx cupular, truncate to<br />
lobed; lobes (when present) 4-5, rounded to ovate;<br />
Condaminea is the only <strong>Rubiaceae</strong> to have largefoliose,<br />
acuminate stipules made of four units, two<br />
interpetiolar and two intrapetiolar, usually connate at<br />
base, forming a short sheath above the petiole attachments<br />
(Figs. 3F,G, 44A, 45C,D). These four stipular<br />
units have been interpreted either as two interpetiolar<br />
bifid stipules per node or four (two inter- and two<br />
intrapetiolar) entire stipules per node, depending on<br />
the basal connections.<br />
Condaminea is also easily detectable by its sessile<br />
to short-petiolate leaves, which are commonly thickcoriaceous<br />
and have undulate margins. Because of its<br />
variable leaf shape (see Fig. 47A-H) and vestiture,<br />
various specific and varietal names have been assigned<br />
to this genus, which are all reduced to synonymy under<br />
C. corymbosa.<br />
The flowers of Condaminea are thick and fleshy,<br />
and with valvate aestivation. During anthesis the filaments<br />
bend toward the lower portion of the flowers,<br />
and the anthers open upward by longitudinal slits,<br />
forming a landing platform for visitor/pollinator insects<br />
(commonly bees). Flowers in Condaminea are<br />
usually flesh-red outside and pearl-white inside.<br />
Seeds in Condaminea are minute, horizontally inserted,<br />
non-winged, truncate at apex, and usually (3-)<br />
4-angled. The capsules of Condaminea could be confused<br />
with those of <strong>Rustia</strong>, the latter characterized<br />
by their persistent calyx (readily caducous in Condaminea).<br />
Condaminea is commonly found in almost pure<br />
stands of single-stemmed shrubs in disturbed soil, and<br />
secondary vegetation, but exceptionally could grow<br />
into small trees. Condaminea is here recognized as a<br />
genus of two species (C. elegans of dubious identity,<br />
see below), ranging from Costa Rica to Bolivia.<br />
calyx caducous, leaving a white scar on capsules.<br />
Corolla salverform to infundibuliform (tubular with<br />
spreading lobes), aestivation valvate narrowly reduplicate<br />
with contact zone, camose, red to purple outside,<br />
greenish white inside. Stamens 5, partially ex-<br />
Taxonomic History<br />
Condaminea was founded by A. P. de Candolle<br />
(1830) and was dedicated to Charles Marie de La<br />
Condamine. La Condamine directed an expedition to
104 Flora Neotropica<br />
?iJ<br />
FI.44 onanie crrnoa AD rm rshmteil n Dlree& edua642 TX EfomDlpee<br />
cec. .Opnflwr,sown zgmopicpsiin fathr, n eogae rcptv sima . aur apue
Systematic Treatment of the Genera Studied 105<br />
Ecuador to-prove that the earth is not an exact sphere,<br />
but rather flattened at the poles. He later descended<br />
the Amazon River (La Condamine, 1745) on his way<br />
back to France. In addition, La Condamine was the<br />
first to describe Cinchona, as reported in the Spanish<br />
translation by Ruiz (1778). La Condamine's description<br />
and illustration were later used by Linnaeus<br />
(1753) for his description of Cinchona.<br />
Condaminea corymbosa was first described and<br />
illustrated by Ruiz and Pav6n (1799) as Macrocnemum<br />
corymbosum. Kunth (1818) included M. corymbosum<br />
into "Macrocnemum sect. 2) bracteis minutis,<br />
sessilibus (An genus distinctum?)" [with M. tinctorium<br />
(= Simira rubescens (Bentham) Bremek. ex<br />
Steyerm.)], which was the first suggestion to treat this<br />
taxon as a separate genus. Macrocnemum corymbosum<br />
was also accepted by A. Richard (1830).<br />
A number of species were transferred by de<br />
Candolle (1830) from Macrocnemum (probably influenced<br />
by Kunth, 1818) to the newly founded Condaminea<br />
as C. corymbosa, C. tinctoria (= Simira rubescens<br />
(Bentham) Bremek. ex Steyerm.), C. microcarpa<br />
(= Bathysa sp.?), C. venosa (not Condaminea!), and C.<br />
glabrata (not Condaminea!), reaching the highest<br />
number of species ever attributed to this genus.<br />
Schumann (1889) described C. corymbosa var.fi<br />
pubescens Spruce ex Schum., which is here treated<br />
as synonymous with C. corymbosa. Schumann (1891)<br />
maintained the species of Condaminea previously<br />
recognized by de Candolle (1830), but placing C. tinctoria<br />
as a Sickingia [= Simira].<br />
Rusby (1896) described C. angustifolia from narrow-leaved<br />
specimens collected in Bolivia, and<br />
Standley (1929) described C. breviflora, which he<br />
separated from C. corymbosa because of smaller flowers<br />
and petiolate leaves. I consider both of these species<br />
to be synonymous with C. corymbosa.<br />
Standley (1936) tentatively recognized three species<br />
of Condaminea: C. microcarpa, C. corymbosa<br />
(with two varieties), and C. macrophylla; in addition<br />
he discussed the identity of C. glabrata and C. venosa<br />
(see Excluded Species, below).<br />
Steyermark (1974) stated (translated from Spanish):<br />
"Four or five species [of Condaminea] are known<br />
from Panama to Peru and Bolivia; only one is known<br />
in Venezuela [C. corymbosa]."<br />
Dwyer (1980) stated that "there are about three<br />
species in this genus." In Panama he recognized two<br />
species: C. corymbosa and his newly described C.<br />
petiolata (= Rondeletia panamensis DC.; see Excluded<br />
Species).<br />
Burger and Taylor (1993) also recognized Condaminea<br />
as a genus of"four or five species ranging from<br />
Costa Rica to Venezuela, Peru, and Bolivia."<br />
Key to the species of Condaminea<br />
1. Stipules (2-)3-7(-9) cm long, at least 4 mm<br />
wide at base; corollas 2-3 cm long; inflorescences<br />
lax corymbs, with distal cymules of<br />
>4 flowers; leaves elliptic, ovate to obovate<br />
(rarely lanceolate), L/W 1.5:1 to 3.5:1 (widespread<br />
from Costa Rica to Bolivia) .. 1. C. corymbosa<br />
1. Stipules 0.5-1.7 cm long, 2-4 mm wide at base;<br />
corolla 0.7-1.5 cm long; inflorescences very<br />
reduced corymbs, with distal cymules of 2-4<br />
flowers; leaves lanceolate to narrowly elliptic,<br />
L/W 3:1 to 4:1 (Peru: Madre de Dios, Manu<br />
National Park) ........................................ 2. C. elegans<br />
1. Condaminea corymbosa (Ruiz & Pav6n) A. P. de<br />
Candolle, Prodr. 4: 402. 1830. Macrocnemum<br />
corymbosum Ruiz & Pav6n, Fl. Peruv. 2: 48, fig.<br />
189.1799. Type. Peru. Chinchao, 1795 (fl), Tafalla<br />
& Manzanilla s.n. (lectotype, MA, selected by<br />
Delprete, 1999b; photo-MA at F, MO, TEX).<br />
Figs. 3F-H, 4F, 7D-F, 15C-F, 44A-D, 45A-C,<br />
46A-F, 47A-H, 48, 49<br />
Condaminea breviflora Standley, Publ. Field Columbian<br />
Mus., Bot. Ser. 6: 264. 1929. Type. Ecuador. Zamora-<br />
Chinchipe: E slopes of E Andes of Loja, 800-1200<br />
m, Oct-Jan 1906 (fl), Lehmann 5651 (holotype, F,<br />
N.578,485; photo-F at G, VEN; isotype, K).<br />
Condaminea angustifolia Rusby, Mem. Torrey Bot. Club<br />
6: 45. 1896. Type. Bolivia. Between Guanay and<br />
Tipuani, Apr-Jun 1892 (fl), Bang 1429 (holotype,<br />
NY; isotypes, F, G, K, MO, M, NY[2], US).<br />
Condaminea corymbosa (Ruiz & Pav6n) A. P. de<br />
Candolle var. pubescens Spruce ex K. Schumann in<br />
Martius, Fl. Bras. 6(6): 258. 1889. Type. Peru. San<br />
Martin: Vie. of Tarapoto, slopes close to Rio Mayo,<br />
Jul-Aug 1856 (fl), Spruce 4579 (lectotype, K, selected<br />
by Delprete, 1999b; isolectotypes, BR, F, G[2],<br />
GH, K[2], NY).<br />
Shrubs 2-5 m tall (exceptionally small trees to<br />
15 m tall, to 30 cm dbh), multicaulous shrubs, excep-<br />
tionally single-stemmed trees; bark fissured, grayish<br />
brown to yellowish brown. Leafy branchlets terete<br />
to subterete, glabrous, dark green; older branches rug-<br />
ose, pale brown; lenticels sparse, whitish, ca. 1 mm<br />
wide. Stipules 4 at each node, connate at base (basal<br />
2-8 mm), foliose, long lanceolate, acuminate, gla-<br />
brous to puberulent to hirsutulous outside, glabrous<br />
with basal area of colleters inside; (2-)3-7(-9) x 4-<br />
9(-13) mm, persistent, pale green (to reddish green),<br />
caducous, leaving a white-grayish linear scar. Leaves<br />
sessile, subpetiolate to short-petiolate, (15-)25-55<br />
(-65)x (10-)15-29cm,L/W 1.5:1 to 3.5:1; lanceolate<br />
to elliptic to oblanceolate to obovate, rounded to cor-
106<br />
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Flora Neotropica<br />
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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<br />
in apical bud, before opening into four units. D. Stipules in young vegetative bud (ca. 6 cm long).<br />
.
Systematic Treatment of the Genera Studied 107<br />
'::<br />
f:x<br />
..<br />
108 Flora Neotropica<br />
m!. . 4<br />
G. C.<br />
angustfolia, Bang 1429 type. H. Lewis 88914.<br />
;<br />
C. P er 5251. D. Cro 69930. . Subsessile leaf, Brandbyge & Asanza 30431. F-H. Petiolate leaves. Schunke 9494.<br />
"<br />
... _? .<br />
. v g . ..<br />
S g<br />
T. -I<br />
FIG. 47. Leaf variation in Condaminea corymbosa. A-D. Sessile leaves. A. Steyermark 56659. B. Skutch 3981.<br />
C. Pittier 5251. D. Croat 69930. E. Subsessile leaf, Brandbyge & Asanza 30431. F-H. Petiolate leaves. F. Schunke 9494.<br />
G. C. angustifolia, Bang 1429 type. H. Lewis 88914.
Systematic Treatment of the Genera Studied 109<br />
Adir. IIcrb.rzaNr,vg,m<br />
M04ab,earA4 t AM<br />
'"s.^' ...' 'I<br />
rr rra.e e.Iirpab 4 _M<br />
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48. M<br />
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ascending at medio-distal portion of blade; tertiary<br />
veins subparallel; quaternary veinlets openly reticu-<br />
late, evident below; petioles (when present) to 25 mm<br />
long, 3-7 mm thick, terete to flattened adaxially. In-<br />
4a.SAr*<br />
?..<br />
~1. l<br />
o<br />
7<br />
florescences open corymbose panicles with 1-2(-3)<br />
pairs of opposite branches terminating in variably<br />
reduced cymules; 18-50 cm long, basal branches 4-<br />
25 cm long; basal portion of axis not branched 2-30
110 Flora Neotropica<br />
cm long; rachis terete to decussately compressed, rachis<br />
and branches glabrous to puberulent to hirtellous;<br />
flowers on terminal cymules; distal bracts 2-7 x 2-4<br />
mm, deltoid to narrowly triangular, sometimes similar<br />
to stipules (foliose, many-nerved) to 2.5 x to 5 mm,<br />
lanceolate, acuminate; bracteoles subtending flowers<br />
1-3 x 1-2 mm, deltoid. Flowers pedicellate, pedicels<br />
1-10 mm long, glabrous to puberulent to hirtellous;<br />
hypanthium obconical to narrow-obovate, 6-12 x 3-<br />
6 mm, glabrous to hirtellous, sometimes verrucate;<br />
flower buds short-clavate, apiculate at tip. Calyx<br />
cupular, truncate to evidently lobed, 3-10 x 5-12 mm,<br />
glabrous to sparsely puberulent; lobes (when present)<br />
(4-)5, deltoid to rounded, 1-6 mm long; leaving a<br />
white scar on capsule. Corolla salverform (tubular<br />
with spreading lobes), 2-3 cm long, carose, creamwhite<br />
to purple outside, greenish white inside; tube<br />
cylindrical, 7-16 mm long, 3-6 mm wide at base and<br />
6-8 mm wide at orifice, glabrous outside, glabrous to<br />
sparsely pilose inside, with a ring of short-sericeous<br />
hairs 4-7 mm from the base, at the same point of filament<br />
attachment; lobes (4-)5, /3--2 of corolla length,<br />
(3-)5-9 x 3-7 mm, ovate to oblong, pointed at apex,<br />
glabrous outside and inside. Stamens 5, subequal,<br />
attached 4-9 mm from the base of the tube and bending<br />
toward lower portion at anthesis; filaments 4-8<br />
mm long, terete distally, flattened and widened at base,<br />
white, sparsely pubescent at base; anthers narrowly<br />
elliptic, 5-8 x 1-1.5 mm, dorsifixed at medial zone,<br />
base elliptic to sagittate, smooth throughout, dehiscing<br />
by longitudinal slit toward the inside, yellow.<br />
Pollen exine reticulate, columellate. Style exserted,<br />
15-22(-26) mm long, terete to moderately compressed,<br />
stout, glabrous, grass-green; style branches<br />
ovate, membranous, often unequal, reversed at maturity,<br />
3-5 mm long, stigmatic surface microscopically<br />
(40x) papillose; immature fruits green to reddish,<br />
semi-fleshy, glabrous to minutely puberulent. Capsules<br />
obovoid to oblong-turbinate, acute to obtuse at<br />
base, shallowly obtuse at apex, (10-)14-20 x the year except March and September. Fruiting specimens<br />
were collected throughout the year except July<br />
and November.<br />
Ecology. This species is rather frequent on disturbed<br />
slopes of the South American Andes, and occurs<br />
sometimes as the first successional woody shrub<br />
(often in pure stands of individuals 2-3 m tall) on landslides<br />
and in cleared or freshly disturbed soil of recently<br />
built mountain roads. Because of this, Condaminea<br />
corymbosa plays an important role in<br />
containing the frequent landslides in the rainy Andean<br />
slopes and protecting the young seedlings of endemic<br />
tree species. I would very much encourage artificial<br />
dissemination of this short-lived shrub for rapid establishment<br />
of woody species on disturbed slopes so<br />
as to protect newly built Andean roads from frequent<br />
landslides.<br />
Reproductive biology. The flowers of Condaminea<br />
corymbosa are carnose, purple to flesh-red outside<br />
and perlaceous inside, and are historically described<br />
as strictly actinomorphic. Personal observations<br />
(Fig. 46A-F) have revealed that the flowers of this<br />
species are obliquely presented (sometimes erect),<br />
never pointing downward, and that during anthesis the<br />
upper filaments bend toward the lower portion of the<br />
corolla throat (not caused by their weight alone), arranging<br />
the anthers parallel to each other and dehiscing<br />
upward by longitudinal furrows, forming an easy<br />
landing platform for possible pollinators (Fig. 46A-<br />
F). In Ecuador the most frequent visitors are bees (not<br />
identified), which are probably the pollinators of this<br />
species ("lap-pollination"). Occasional individuals of<br />
C. corymbosa were observed to be associated with ants<br />
(pers. obs.). I suspect these ants to be only casual visitors,<br />
and no obvious mutualism was observed.<br />
6-11<br />
mm, light to dark brown, without lenticels, glabrous<br />
to puberulent throughout, with a cream-white circular<br />
calyx scar, 0.5-1.1 mm wide, 5-8 mm diam. Seeds<br />
0.57-0.77 x 0.13-0.3 mm; irregularly 3-4-angular,<br />
trapezoid, compressed, testa reticulate.<br />
Representative specimens examined. COSTA RICA.<br />
CARTAGO: Field of Inst. Interamericano de Ciencias<br />
Agricolas, Turrialba, 600 m, Feb 1949 (fr), Le6n 1484 (US).<br />
HEREDIA: Near Rio Col6n, between San Isidro and Col6n, 6<br />
Aug 1971 (fr), Dwyer et al. 523 (NY). LIMON: Sukut, Alto<br />
Uren, margin of Rio Ur6n, 26 Oct 1985 (fl), G6mez et al.<br />
23844 (MO, NY); Reserva Indigena Talamanca, confluence<br />
of Rio Sukut with Rio Ur6n, 09?24'N, 82?58'W, 325 m, 3<br />
Jul 1989 (fl), Hammel et al. 17540 (MO). PUNTARENAS: Be-<br />
Distribution (Fig. 49) and ecology. Common<br />
shrubs (see Fig. 45A), exceptionally single-stemmed<br />
trees, in disturbed soil and roadside vegetation on<br />
mountain slopes, in dry, deciduous or moist primary<br />
and secondary forests, coastal and mainland rain forests<br />
to cloud forests on eastern and western Andean<br />
slopes, 120-2000 m, from Costa Rica to Bolivia.<br />
Flowering specimens were collected in all months of<br />
tween Villa Neilly and Agua Buena, Golfito, 500 m, 21 Oct<br />
1964 (fl), Jimenez 2451 (F[2], NY); Rio Sonador, El General<br />
Valley, 600 m, 31 Jan 1965 (fr), Williams et al. 28805<br />
(F); 24.7 km SE of Brujo, on rd. Coto Brus-San Vito, 300-<br />
350 m, 17 Mar 1978 (fl-fr), Almeda et al. 4264 (CAS); Callas<br />
Gordas, Coto Brus, 1160 m, 14 Nov 1985 (fl), Gdmez-<br />
Laurito 10694 (F); forest of Boruca, Feb 1891 (fr), Tonduz<br />
3947 (BR[2]). SAN Jos:: Vic. of El General, 640 m, Jan 1939<br />
(fl), Skutch 3981 (GH, K, NY, US[2]).
Systematic Treatment of the Genera Studied 111<br />
PANAMA. CHIRIQUi: 20Ykm N of David, 14 Dec 1966<br />
(fl), Lewis et al. 692 (GH, K, MO, NY, UC, US); Boqueli,<br />
1500 m, 26 Jun 1938 (fr), Davidson 792 (A); vic. of San<br />
Felix, E of Chiriqui, 120 m, Dec 1911 (fl), Pittier 5251 (F,<br />
GH, NY, US). DARItN: 4 km S of Jaque, 2 Feb 1981 (fr),<br />
D 'Arcy & Sytsma 14584 (MO, NY); Piifas, 2 Mar 1967 (fr),<br />
Duke 10621 (ECON, MO).<br />
COLOMBIA. ANTIOQUIA: Vic. of Medialuna, 1700 m,<br />
Chama, 19 May 1944 (fl-fr), Steyermark 56659 (F[2], US[2],<br />
VEN); Dto. Libertador, between San Rafael and Via El<br />
Morro, 18 Aug 1972 (fl), Benitez de Rojas 1513 (MY[2]);<br />
between Merida and San Rafael del Chama, 1600 m, 16 May<br />
Oct 1945 (fr), Molina s.n. (US); Quebrada Verruga, near<br />
Uramita, 1600 m, 8 Apr 1944 (fr), Core 566 (US). CALDAS:<br />
8 km up Rio Apii, 1015 m, 29 Nov 1944 (fl), St. John 20871<br />
(UC, P). CAQUETA: Florencia, 420 m, Dec 1930 (fl) Perez-<br />
Arbeldez 697 (COL, US). CAUCA: El Tambo, vic. of La Paz,<br />
1700 m, 29 Aug 1949 (fl), Idrobo 287 (US); Tierra Adentro,<br />
Rio Paez drainage, Segovia, Jun 1943 (fl), Hurtado 5 (US).<br />
CHOC6: Mecana, N of Bahia Solano, 06016'N, 77?21W, 3<br />
Jan 1984 (fl), Juncosa 1660 (MO); Bahia Solano (Puerto<br />
Mutis), 0-20 m, 4 Jan 1973 (fr), Gentry & Forero 7195<br />
(VEN); rd. Tutunend6-El Carmen, Alto Rio Atrato, 29 Apr<br />
1979, Forero et al. 6093 (COL). CUNDINAMARCA: Intersection<br />
of rds. from Pandi and from Arbelaez, 1300 m, 21 Feb<br />
1970, Uribe-Uribe 6385 (COL); S of Silvania, on rd. to<br />
Fusagasuga, 1400 m, 26 May 1972, Barclay 3463 (COL).<br />
HUILA: S of Acevedo, 16 Aug 1944 (fl), Little 8490 (MO,<br />
US). META: Valley of Rio Tigre, Colombia-Uribe trail,<br />
03?14'N, 74?36'W, 18 Dec 1942 (fl), Fosberg & Holdridge<br />
19409 (UC). NARISO: Mallama, trail Planada-Pialapi, 1700<br />
m, 27 May 1985 (fr), Benavides 5651 (MO). NORTE DE<br />
SANTANDER: Region Sarare, riverside of Rio Cubug6n, El<br />
Indio, 13 Nov 1941, Cuatrecasas 13074 (COL). PUTUMAYO:<br />
Near Ecuadorian border, Rio San Miguel, between the affluents<br />
Bermeja and Conejo, 300 m, 13 Dec 1940 (fl),<br />
Cuatrecasas 11059 (US). SANTANDER: La Corcova, Tona,<br />
1866 m, 12 Oct 1977 (fr), Renteria et al; 658 (COL, MO);<br />
Bucaramanga, May 1948 (fl), Sandemer 6055 (K). TOLIMA:<br />
Region of Herrera, Rio Saldafia, 03?22'N, 75?56'W, 1800<br />
m, Jan 1945 (fr), Hurtado 111 (US[2]); canyon of Rio Ata,<br />
SW of La Colonia, 5 Oct 1944 (fl), Little 8757 (COL, MO);<br />
Prov. Mariquita, Ibagu6, 2000 m, May 1854, Triana 1782<br />
(3262.1) (COL); Central Cordillera, Valley of Rio<br />
Bermellon, near Ibagu6, 1220 m, 29 Oct 1952 (st), Humbert<br />
26973 (NY). VALLE: Tulua, at limits of botanical gardens,<br />
1300 m, 30 Jun 1984 (fl), Devia 558 (COL, MO[2]); Mun.<br />
Argelia, rd. La Argelia-Toro, 1700 m, 2 Sep 1983 (fl), Devia<br />
316 (COL, MO).<br />
VENEZUELA. BARINAS: Dto. Pedraza, W of La<br />
Escampa, Parque Nacional Sierra Nevada, 08031'N,<br />
70046'W, 27 Jun 1988 (fl), Dorr et al. 5719 (NY). MERIDA:<br />
Near M6rida, 1700 m, 24 Feb 1964 (fl-fr), Breteler 3612<br />
(NY, US, VEN[2]); Rio Chama, 1500 m, 15 Nov 1953 (fr),<br />
Little 15872 (VEN); Dto. Libertador, 2 km from Belsante,<br />
quebrada Carvajal, Los Curos, 1800 m, 16 Nov 1962 (fr),<br />
Ruiz Terdn 1204 (VEN); Aricagua, 950 m, Jun 1955 (fl),<br />
Bernardi 2209 (VEN); Dto. Arzobispo Chac6n, rd. Canagua<br />
(1500 m) - Mucuchachi 1964 (fr), Ijjfsz-Madriz 422 (VEN).<br />
ECUADOR. CARCHI: Trail Chical-Quinyul, 78?13'W,<br />
01?02'S, 1200 m, 13 Jan 1983 (fr), Barford 41389 (AAU);<br />
E of Maldonado, 23 Aug 1943 (fr), Steere 8060 (F);<br />
Maldonado, 1500 m, 4 Oct 1981 (fl), Werling & Leth-Nissen<br />
226 (QCA[2]); Maldonado, rd. Maldonado-Chical,<br />
1500-1700 m, 14 Aug 1994 (fl), Delprete & Verduga 6412<br />
(NY[3], QCA, QCNE, TEX, UPS[2]). COTOPAXI: Rio Pilalo,<br />
km 52 on Quevedo-Latacunga, 750-900 m, 29 Oct 1981 (st),<br />
Dodson & Dodson 12001 (F). EL ORO: Ca. 10 km SE of<br />
Portovelo, 850 m, 8 Oct 1944 (fl), Wiggins 10947 (DS, F);<br />
El Placer, W of Morona, 340 m, 28 May 1979 (fl), Albert de<br />
Escobar 1373 (QCA). ESMERALDAS: Lita, on Ibarra-San<br />
Lorenzo, 550 m, 8 Jun 1978 (fr), Madison et al. 5051 (F,<br />
QCA, SEL). IMBABURA: Between Guayupe and Peias<br />
Negras, 78014'W, 00?44'S, 1050 m, 6 Dec 1980 (fl), Holm-<br />
Nielsen & Jaramillo 28968 (AAU); rd. Salinas-Lita, 4 km<br />
before Lita, 78?26'W, 00?50'N, 580 m, 25 Jul 1992 (fl),<br />
Freire-Fierro et al. 2333 (QCA). MORONA-SANTIAGO: Rd.<br />
Suc6a-Macas, 880-1080 m, 78?07'W, 02?20'S, 26 Sep 1979<br />
(fr), Holm-Nielsen et al. 20567 (AAU, QCA); Bomboiza,<br />
15-18 km SE of Gualaquiza, 700 m, 03?27'S, 78?34'W, 1-<br />
3 Nov 1986 (fl), Zaruma 802 (MO, NY, QCNE, QCA).<br />
NAPO: Canton Archidona, rd. Hollin-Loreto, 01?04'N,<br />
77?36'W, 1300 m, 3 Oct 1992 (fr), Delprete & Luteyn 6205<br />
(AAU, MO, NY, QCA, QCNE, TEX), 6207 (NY); Rio<br />
Quijos with Rio Salado, below Baeza, 2 Mar 1944 (fr),<br />
Ownbey 2686 (F[2], MO[2], US[2]). PASTAZA: Rio Bobonaza,<br />
between Cachitama and outlet of Rio Bufeo, 74?40'W,<br />
02?20'S, 19 Jul 1980 (fl), 0llgaard et al. 34667 (AAU,<br />
QCA); Mera, 2 km toward valley, 27 Jun 1992 (fl), Delprete<br />
et al. 6049 (AAU, MO, NY, QCA, QCNE, TEX); Mera, bank<br />
of Rio Pastaza, ca. 1100 m, 11 Nov 1955 (fl),Asplund 18347<br />
(B). PICHINCHA: Rd. La Uni6n del Toachi-San Francisco de<br />
las Pampas, km 3, 1100-1200 m, 19 Mar 1985 (fl), Harling<br />
& Andersson 23151 (QCA); Bosque Protector Mindo, ridge<br />
between Rio Mindo and Rio Bagasal, 00?02'S, 78?48'W,<br />
1400 m, 15 Jul 1992 (fl), Delprete & Webster 6104 (AAU,<br />
DAV, F, MO, NY[2], QCA, QCNE, TEX, UPS); Canton<br />
Santo Domingo de los Colorados, Uni6n del Toachi, 9 Oct<br />
1992 (fl), Delprete et al. 6218 (NY, QCA). SucUMBmOS: Section<br />
NW of San Rafael waterfall, 1300 m, 11 Oct 1990 (fl),<br />
Jaramillo et al. 13199 (QCA). TUNGURAHUA: Rio Negro,<br />
00?15'S, 78020'W, 1200 m, 18 Sep 1987 (fl), Zak &<br />
Jaramillo 3577 (AAU, GB); rd. Bafios-Puyos, km 11.3-18,<br />
78?15'W, 01?24'S, 1650 m, 30 Apr 1992 (fl), Freire-Fierro<br />
& Luteyn 2159 (QCA). ZAMORA-CHINCHIPE: Mtn. ridge N<br />
of Zamora, between Rio Janeiro and Rio Yacuambi, 29 Jun<br />
1944 (fr), Camp A-C25 (F); rd. Zumba-Valladolid, between<br />
Palanuma and Palanda, 04?40'S, 7908'W, 1200 m, 13 May<br />
(974 m), Andean Cordillera, Cerro 1985 (fl), Stein & D 'Alessandro 2776 (QCNE); Lim6n (Gende<br />
Las Flores, s.d. (fl), Ruiz Terdn & L6pez Figueras 658 eral Plaza Gutierrez), rd. San Bartolo-Uni6n, 24 Aug 1994<br />
(VEN[3]), 4 Jun 1953 (fr), Bernardi 563 (NY); between La<br />
Trampa and Lagunillas, 1188-2430 m, 29 Apr 1944 (fl),<br />
Steyermark 56208 (VEN); Mt. San Jacinto, N side of Rio<br />
(fl), Delprete & Verduga 6436 (NY[2], QCA, TEX).<br />
PERU. AMAZONAS: Quebrada Sasa, 600 m, 25 Aug 1975<br />
(fl), Kayap 2012 (F, MO, NY); Rio Cenepa, vic. of Huam-
112 Flora Neotropica<br />
* as- 1'''--"-'-'-40 '<br />
\r~ - p<br />
V -,<br />
0~~~~ -~~. pi -<br />
I~~~~~~~~I<br />
6. ~ ~ ~a 1' :<br />
- . 2~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I<br />
t i~~~~~~~~~~~~~~~~~U<br />
* m ~~~~~~~~~~~~~~~~~~??<br />
ma-wa ~ ~ ~ ~ . ?<br />
:Me~~~~~~~~~~~~<br />
*~~~~~~~~~~~~~~~~<br />
FIG. 49. Distribution of Condaminea<br />
ED~~~~~~~~~~~~~~~~<br />
corymbosa (circles) and C. elegans (star).<br />
* 's M -usessa<br />
FI. 9 Dstiutono Cndmne crmbsa(irle)an . elgas str<br />
pami, 5 km E of Chavez Valdivia, 78?30'W, 04?30'S, 200<br />
m, 15 Aug 1978 (fr), Ancuash 1509 (MO); Prov. Bagna,<br />
Valley of Rio Maration, above Cascadas de Mayasi, near<br />
Campamento Montenegro (kms 276-280 of Marafion rd.),<br />
450 m, 3-4 Sep 1962 (fl), Wurdack 1831 (VEN).<br />
CAJAMARCA: Prov. Ja6n, 800 m N of Chirinos, 1700 m, 30<br />
Jul 1943 (fl), Hodge 6048 (ECON). Cuzco: San Miguel,<br />
Urubamba Valley, 1800 m, 26 May 1915 (fl), Cook & Gil-<br />
bert 913 (F, US); Prov. La Convenci6n, E of Quillabamba,<br />
1200 m, 20 Apr 1981 (fl), Young & Eisenberg 255 (MO,<br />
NY). HuANuco: Prov. Pachitea, Parque Nacional de Iparia,<br />
Rio Pachitea, 1 km above Tournavista, 22 Jun 1967 (fl),
Systematic Treatment of the Genera Studied 113<br />
Schunke 2072 (F, NY); Chinchao, Pampayacu, 1 km E of Condaminea corymbosa is easily recognizable for<br />
Cochero, 1050-1200 m, 16 Feb 1973 (fl),Schunke Vigo 5807 its four long-lanceolate foliose stipules (two<br />
(CAS, VEN); Prov. Leoncio Prado, Dto. Rupa Rupa, Calpar interpetiolar and two intrapetiolar) at each node (Figs.<br />
Bella, Rio Monz6n, 700-900 m, 2 Jul 1976 (fl), Schunke<br />
3F-H, 44A; Fig. 45C,D for stipules in apical buds),<br />
9494 (F, MO, NY); Rupa Rupa, environs of Tingo Maria, its<br />
09?18'S, 75059'W, 680 m, 24 Mar 1982 (fl), King & Ramirez- fleshy flowers, purple outside and perlaceous in-<br />
Rengifo 372 (B); Chinchao, 1 Aug 1964 (fl-fr), Dwyer 6178 side, and by its sessile to short-petiolate coriaceous<br />
(MO[4], UC). JUNIN: Huacapistana, 1800-2400 m, 5-8 Jun<br />
leaves. Condaminea corymbosa is most commonly a<br />
1929 (fl), Killip & Smith 24130 (F, NY, US); La Merced, short-lived, single-stemmed shrub 2-4 m tall, but I<br />
700 m, May-Jun 1929 (fl), Killip & Smith 23506 (F, NY, have encountered it as a tree 15 m tall with 30 cm dbh.<br />
US); Prov. Tarma, gorge of Rio Huasahuasi, 7 km E of The type of Macrocnemum corymbosum was de-<br />
Huasahuasi, 12 Aug 1957 (fr), Hutchison 1153 (F, NY, UC); scribed and beautifully illustrated in Ruiz and Pav6n's<br />
Prov. Chanchamayo, Rio Tulumayo valley, rd. from San famous Flora Peruviana et Chilensis (1799: pl. 189);<br />
Ram6n to Vitoc and Chilpes, 1000-1250 m, 11o15'S, no precise type was mentioned and the protologue<br />
75?15'W, 9 Mar 1985 (fl-fr), Stein & Todzia 2358 (MO, NY). vaguely states: "Habitat in collibus et runcationibus<br />
LORETO: Above Pongo de Manseriche, hills left of Rio<br />
callidis, versus Chinchao, Acomayo, Pillao et Mufia<br />
Marafion, 250 m, 26 Dec 1931 (fr), Mexia 6351 (CAS, F, K, vicos." Most of the material collected<br />
MO, NY, UC, US). PUNO: Prov. Sandia, near Ketlin, 2000<br />
by Ruiz and<br />
Pav6n in this area was lost on the<br />
m, 4 Jun 1942 (fl), Metcalf30676 (A, UC[2]). SAN MARTIN:<br />
ship San Pedro de<br />
Boqu6ron Pass, 92 km from Tingo Maria, 400 m, 16 Dec Alcantara returning to Spain, in a shipwreck along the<br />
1949 (fl), Allard 22129 (UC); Zepelacio, near coast of<br />
Moyobamba, Portugal. Ruiz and Pavon returned to Spain<br />
1100 m, Jun 1934 (fl), Klug 3672 (A, CAS, F, MO); Prov. in 1788, leaving the two botanists Juan Jose Tafalla<br />
Lamas, trail from Tabalosos to Lamas, E of Rio Mayo, 13- and Juan A. Manzanilla (between 1793 and 1797) to<br />
15 Sep 1937 (fr), Belshaw 3422 (F, GH, MO, NY, UC, US); re-collect in sites of the collections lost by their pre-<br />
Prov. San Martin, hills NE of Tarapoto, on rd. to decessors (Estrella, 1991). Tafalla and Manzanilla sent<br />
Yurimaguas, 1000 m, 06030'S, 76?20'W, 11 Feb 1985 (fl- their collections to Spain, where they were then filed<br />
fr), Stein & Todzia 2142 (MO, NY); Prov. Rioja, Disto. Pardo in the Herbarium Peruvianum by Ruiz and Pavon and<br />
Miguel, 1.5 km W of Aguas Verdes, 1320-1440 m, 24 Jun included in Flora Peruviana et Chilensis. In the Her-<br />
1997 (fr), Sdnchez Vega & Dillon 8906 (NY), 26 Jun 1997<br />
barium Peruvianum of Madrid (MA) are preserved<br />
(fl), 9052 (NY). UCAYALI: Prov. Portillo, Bosque Von<br />
four<br />
Humboldt, quebrada Tahuahillo, 200 m, 20 Jun 1981 (fl),<br />
specimens annotated as Macrocnemum<br />
Young 942 (MO, NY); rd. Fed. Basadre, between Pucallpa corymbosum by Pav6n (in his handwriting) with the<br />
and San Alejandro, 08?22'S, 74?31'W, 21 Jan-3 Feb 1987 following labels: "Pentand. Monog. - Macrocnemum<br />
(fl), Trigoso 1025 (K).<br />
corymbosum<br />
BRAZIL. MINAs GERAIS: Mun. Ituiutaba, San Vicente,<br />
29 May 1950 (fl), Macedo 2402 (MO[2], US[2], TEX).<br />
BOLIVIA. COCHABAMBA: Prov. Chapare, vic. of San<br />
Onofre, 1600 m, 12 Feb 1929 (fl), Steinbach 9292 (GH, K).<br />
LA PAZ: Yungas, 1890 (fl), Bang 610 (DS, F, K, NY[2], UC,<br />
US); Nord Yungas, Covipata, Hazienza El Choro, 1700 m,<br />
28 Jun 1930 (fl), Buchtien 8186 (NY, US); Sur Yungas, basin<br />
of Rio Bopi, San Bartolom6, near Calisaya, 750-900 m, 1-<br />
22 Jul 1939 (fr), Krukoff 10032 (F, NY); Inquisivi, bridge<br />
over Rio Khara, 2 km NW of Licoma Pampa, 16?48'S,<br />
67?13'W, 15 Jun 1988 (fr), Lewis 88825 (MO[3]); Apolo,<br />
27 Feb 1902 (fl), Williams 322 (NY[2], US); Coroico,<br />
Pajonales, 1500 m, 19 Mar 1928 (fl), Troll 1745 (B); Prov.<br />
Sur Yungas, W of Chulumani, 16?23'S, 67?34'W, 1900 m,<br />
17 Sep 1947, Fosberg 28672 (CAS, US[2]).<br />
Local names and uses. Venezuela: anime chiquita<br />
(Steyermark 56208), cacahuito (Steyermark 56306),<br />
guamo (Steyermark 56208). Colombia: lengua de vaca<br />
(Goudot s.n.). Peru: caspi blanco (Trigoso 1025),<br />
chullachasqui (Trigoso 1025), larangui (Hodge 6048),<br />
mun baikuanin (Berlin 1812).<br />
Leaves used for wrapping food, and wood used in<br />
sugar mills (Venezuela, Steyermark 56306).<br />
- Vulgo. Carato. - F. P. c. 1. NO 212. -<br />
Ex Chinchao. - Afio 95" (Fig. 48), "Macrocnemum<br />
corymbosum - Vulgo. Carato insilbatis - Fl. Per. Tab.<br />
189 - Cochero," "Macrocnemum corymbosum - Fl.<br />
Peruv. 2. Ta. 189," and "Macrocnemum corymbosum<br />
F. P." The first specimen, which was collected by<br />
Tafalla and Manzanilla in 1795 and has attached to it<br />
the complete description of this species (Pav6n's<br />
handwriting), was selected by Delprete (1999b) as the<br />
lectotype (Fig. 48); the other specimens are treated as<br />
paratypes.<br />
Condaminea corymbosa has leaves that vary from<br />
sessile through subsessile, to short-pedicellate (Fig.<br />
47A-H) and glabrous (mostly in Central America and<br />
northern South America) to short-petiolate and glabrous<br />
to pubescent (mostly in Peru and Bolivia); in<br />
shape they vary from broadly elliptic to narrowly lanceolate<br />
(Fig. 47A-H). The floral morphology is relatively<br />
constant throughout its distribution, except for<br />
the calyx, which tends to be slightly more expanded<br />
in southern South America. Because of its variation<br />
in shape and vestiture of the vegetative parts throughout<br />
its range, several specific and varietal epithets have<br />
been assigned to this species.
114 Flora Neotropica<br />
This species is known from only one Brazilian ing olive-green and chartaceous, glabrous above and<br />
collection made in the state of Minas Gerais (Mun. below; primary and secondary veins glabrous, fine,<br />
Ituiutaba, San Vicente, Macedo 2402), a locality quite prominent below; secondary veins 18-22 each side;<br />
distant from the southernmost Bolivian Andes. Nev- tertiary veins subparallel, quaternary veinlets openly<br />
ertheless, this disjunct collection does not differ in any reticulate, faintly evident below; petioles 9-16 mm<br />
morphological character from its Andean relatives. long, 1-2 mm thick, subterete. Inflorescences very<br />
I have attempted to maintain Condaminea reduced corymb, long pedunculate, with or without a<br />
corymbosa var. pubescens from the Andes of south- pair of opposite branches terminating in extremely<br />
ern Peru and Bolivia, where the leaves of this species<br />
reduced cymules of 2-4 flowers; 16-22 cm long,<br />
are almost consistently petiolate, with blades thick- branches (when present) 3.5-5 cm long; basal portion<br />
coriaceous, often lanceolate, undulate and of axis not branched 10.5-14.5 cm<br />
commonly<br />
long; rachis terpuberulent<br />
to pubescent beneath. However, as a re- ete, rachis and branches glabrous; distal bracts reduced<br />
sult of field observations of various to scales 1-2 mm<br />
populations<br />
long, deltoid; bracteoles subtendthroughout<br />
its range and critical study of herbarium ing flowers ca. 0.5 mm long, deltoid. Flowers pedimaterial<br />
of C. corymbosa, it is obvious that the above cellate, pedicels 6-10 mm long, glabrous; hypanthium<br />
set of vegetative characters intergrade clinally into the obconical, ca. 6 x ca. 3 mm, glabrous; flower buds<br />
not seen.<br />
more typical elements and that Calyx cupular, evidently lobed, 2-5 x 5-7<br />
infraspecific recognition<br />
for the populations concerned is unwarranted. mm, glabrous; lobes 5, ovate, 2-4 x 3-4 mm; calyx<br />
The specimens Boeke 1501 (NY-2 sheets) have<br />
caducous, leaving a scar on mature capsule. Corolla<br />
yellow-velutinous pubescence on vegetative and re- campanulate, ca. 12 mm long; tube subcylindrical, ca.<br />
6 x ca. 4 mm,<br />
productive parts, capsules abruptly truncated at<br />
glabrous outside and inside; lobes 5,<br />
apex, ca. /2 of corolla<br />
and label data that reports "Corolla<br />
length, ca. 6 x ca. 3 mm,<br />
yellow." As stated<br />
narrowly<br />
triangular, glabrous outside and inside. Stamens and<br />
above, the clinal gradation in external vestiture does<br />
style not seen (the single flower available is in depnot<br />
represent a reliable taxonomic character, and the<br />
auperate condition). Pollen exine reticulate, columelyellow<br />
flowers reported by Boeke are probably an late. Capsules oblong to obovate, acute at base, shalerroneous<br />
observation.<br />
2. Condaminea elegans Delprete, sp. nov. Type.<br />
Peru. Madre de Di6s: Prov. Manu, Parque Nacio-<br />
nal Manui, Cocha Cashu uplands, 400 m, 11?45'S,<br />
71000'W, 13 Sep 1986 (fl-fr), Nunfez 6104 (holo-<br />
type, F; isotypes, MO, NY). Figs. 7H, 49, 50A,B<br />
lowly obtuse at apex, 9-12 x 8-9 mm, black, glabrous<br />
throughout, without lenticels; disk flat, with a pale<br />
brown circular calyx scar at edge of apex, ca. 1 mm<br />
wide, 6-7 mm diam. Seeds 1.3-1.87 x 0.9-1.33 mm;<br />
irregularly 3-4-angular, trapezoid, compressed, yel-<br />
lowish beige, testa reticulate.<br />
Distribution (Fig. 49) and ecology. Known only<br />
from the hills (350-400 m) of the Amazonian forests<br />
in Parque Nacional Manu, Madre de Di6s Province,<br />
Peru. Specimens in end-flowering/early-fruiting stage<br />
were collected in September. Fruiting specimens were<br />
collected in October.<br />
Arbores parve, a C. corymbosa stipularum laminis 5-17<br />
mm longis basi 2-4 mm latis (nec ultra 20 mm longis basi<br />
ultra 4 mm latis), inflorescentiis 2-4-floris (nec multi-floris),<br />
necnon floribus quam 15 brevioribus distantes.<br />
Specimens examined. PERU. MADRE DE Di6s: Parque<br />
Nacional Manu, Rio Manu, Cocha Jarez, hills N of Cocha<br />
Small trees 4-7 m tall. Leafy branchlets terete, on side trail 3 km N of Mirador trail, 350 m, 12?05'S,<br />
thin and delicate, 3.5-4.5 mm thick, densely rugulose, 71?04'W, 5 Oct 1989 (fr), R. Foster & E. Vivar 13333<br />
pale brown, glabrous. Stipules 2 each node, com- (MO, US).<br />
pletely connate, induplicate in bud, splitting at maturity,<br />
lanceolate, glabrous outside and inside, with a This species is tentatively assigned to Condasmall<br />
group of colleters at base inside, 5-17 x 2-4 minea, because the flowering material is depauperate<br />
mm, reddish brown, persistent, leaving a grayish lin- and only partially known. Its stipules, although much<br />
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<br />
lanceolate to oblanceolate, decurrent to acute (rarely (Fig. 7H) are thicker and larger than C. corymbosa<br />
obtuse) at base, acute and short-acuminate at apex, the and resemble those of<strong>Rustia</strong>. No pollen material was<br />
acumen to 5 mm long; olive-green and dull (not wax- available in any of the specimens studied. Unfortushiny)<br />
above, grass-green below, thin-foliaceous; dry- nately, there was only a single depauperate corolla
Systematic Treatment of the Genera Studied 115<br />
.I Y<br />
G<br />
5.<br />
? ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~?<br />
? ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ?<br />
....?.<br />
..:.<br />
..., ?<br />
~~~~~~~~~~.?<br />
.' ? . ? ~'\<br />
r.;<br />
,_,..._"<br />
__ A-<br />
T~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
FIG. 50. Condaminea elegans (NriSez 6104, F). A. Habit of innlorescence, typical stipules, and iong-petiolate leaves.<br />
B. Young capsule with deciduous calyx still attached,<br />
i ' iI
116 Flora Neotropica<br />
without anthers or style. Some old calices were present<br />
on some immature capsules of N~uez 6104; these are<br />
larger (5-6 mm long) and more lobed than any known<br />
member of <strong>Rustia</strong>, and closely resembled the calices<br />
of Condaminea. Fruit size and shape could place this<br />
species in either <strong>Rustia</strong> or Condaminea. The seeds of<br />
Nunez 6104 resembled those of <strong>Rustia</strong> in their size,<br />
shape, and external sculpturing, and were much larger<br />
than any known Condaminea species. Because of the<br />
thickness of the much-enlarged deciduous calyx (re-<br />
duced and persistent in <strong>Rustia</strong>), the bifid short-lan-<br />
ceolate stipules, the absence of pellucid glands in its<br />
leaves, and the much-reduced inflorescences, I pro-<br />
visionally place this species in Condaminea (but it<br />
might as well be a new genus!). The definitive place-<br />
ment of this species will be possible only when flow-<br />
ering collections (and pollen material) become avail-<br />
able.<br />
Condaminea macrophylla Poeppig & Endlicher, Nov.<br />
Gen. Sp. 3: 30. 1845. Type. Peru. Loreto: Prov.<br />
Maynas, vic. of Yurimaguas, May 1929-1932 (fl),<br />
Poeppig 2463 (lectotype, W, here selected; photo-<br />
W at A, F, MO, NY, VEN; isolectotype, frag-F)<br />
= Capirona decorticans Spruce, J. Linn. Soc. 4:<br />
200. 1860.<br />
Its large intrapetiolar stipules, large broadly ovate<br />
and long-petiolate leaves, and flower morphology<br />
are typical of Capirona decorticans, with which it is<br />
synonymous.<br />
Condaminea microcarpa (Ruiz & Pav6n) A. P. de<br />
Candolle, Prodr. 4: 402. 1830. Macrocnemum<br />
microcarpon Ruiz & Pav6n, Fl. Peruv. 2: 49, pl.<br />
188a. 1799. Type. Peru. Chinchao, 1795 (fl-fr)<br />
Tafalla & Manzanilla s.n. (lectotype, MA, here<br />
selected; photo-MA at F, GH, MO, NY, TEX,<br />
VEN) = Not Condaminea! (Bathysa?, Elaeagia?).<br />
The specimen preserved in the Herbarium<br />
Peruvianum of Madrid (MA), was collected by Tafalla<br />
and Manzanilla (not by Ruiz and Pav6n; see discussion<br />
under Condaminea corymbosa, and Estrella,<br />
1991) in 1795 at Chinchao (Peru). The specimen label<br />
reads "Pentand. Monog. - Macrocnemum microparpon<br />
Rejected and Doubtful Species<br />
Condaminea glabrata Bartling ex A. P. de Candolle,<br />
Prodr. 4: 402. 1830. Type. Peru. Mountains of Huanuco,<br />
Haenkes. n. (type not found) = not Condaminea!<br />
De Candolle (1830) described this species as having<br />
bilobed stipules (which is characteristic of Condaminea)<br />
with rounded-obtuse lobes (which exclude<br />
it from Condaminea), and inflorescences with glomerulate<br />
flowers (which exclude it from Condaminea).<br />
Standley (1936) might have overlooked this<br />
species, since it was not cited in his treatment of Peruvian<br />
<strong>Rubiaceae</strong>. The type specimens were reportedly<br />
annotated by Bartling as "Macrocnemum (?)<br />
glabratum in Herb. Haenke" and published in Condaminea<br />
by de Candolle (1830). The identity of this<br />
species cannot be ascertained from the brief description<br />
presented by de Candolle (1830) (no types were<br />
found at G, and PR material was not available), and it<br />
can only be concluded that it is not a Condaminea.<br />
- F. P. c. 1. N. 245 - Ex Chinchao. Aflo 95,"<br />
and it is this specimen that I select as lectotype. I have<br />
seen a set of detailed photographs of this specimen<br />
(sent to me by the staff of the Madrid Herbarium) and<br />
the original illustration, and I concluded that it is not<br />
Condaminea. Its terminal inflorescence, absence of<br />
calycophylls, small sessile flowers with exserted anthers,<br />
stamens attached at the medial portion of the<br />
corolla tube, small capsules dehiscing loculicidally,<br />
and many minute seeds readily suggest a member of<br />
the <strong>Rondeletieae</strong>, possibly Bathysa, Elaeagia, or<br />
Warszewiczia.<br />
Condaminea petiolata Dwyer, Ann. Missouri Bot.<br />
Gard. 67: 117. 1980. Type. Panama. Panama:<br />
Along Rio Maje, 2-3 km above waterfalls, near<br />
Baiano lake, 100 m, 5 May 1976 (fr), Croat 34706<br />
(holotype, MO; isotype, MO) = Rondeletia<br />
panamensis A. P. de Candolle.<br />
The general morphology, placentation, and capsule<br />
dehiscence of the depauperate type specimens suggested<br />
that this taxon belongs to Rondeletia. The absence<br />
of stipules and flowers makes it difficult to<br />
readily identify this species. Nevertheless, the type<br />
specimens have been identified by David Lorence<br />
(annotation label, and pers. comm.) as R. panamensis.<br />
Condaminea tinctoria (Kunth in Humboldt, Bonpland<br />
& Kunth) A. P. de Candolle, Prodr. 4: 402. 1830.<br />
Macrocnemum tinctorium Kunth in Humboldt,<br />
Bonpland & Kunth, Nov. Gen. Sp. 3: 399. 1820.<br />
Sickingia tinctoria (Kunth in Humboldt, Bonpland<br />
& Kunth) K. Schumann in Martius, Fl. Brasil. 6(6):<br />
228. 1889, non Simira tinctoria Aublet, Hist. PI.<br />
Gui. Fran9. 170. 1775. Type. Venezuela. Rio<br />
Orinoco, between Encamarada and Carichana,<br />
Humboldt & Bonpland (type not found). = Simira<br />
rubescens (Bentham) Bremekamp ex Steyermark<br />
(1972: 301).
Systematic Treatment of the Genera Studied 117<br />
This species was described by Kunth (1820) on the<br />
basis of flowering material. Recent fruiting collections<br />
showed that the seeds of this species are unique to<br />
Sickingia, a genus later synonymized with Simira by<br />
Bremekamp (1954a) and Steyermark (1972). This<br />
species is utilized for red dye used on fabrics and<br />
wooden artifacts.<br />
Condaminea utilis Goudot, Compt. Rend. 18: 260.<br />
1844. Type. Colombia. Cundinamarca: Vic. of<br />
Fusagasuga, s.d., Goudot s.n. (P?). = Elaeagia<br />
utilis (Goudot) Weddell (1849).<br />
This species is characteristic of Elaeagia in hav-<br />
ing a resin that exudes from its young vegetative buds,<br />
which is utilized in Colombia and Ecuador as a pro-<br />
tective coating ("varniz" or "varmiz del Pasto") for<br />
bowls and ornamental vases.<br />
Condaminea venosa (Ruiz & Pav6n) A. P. de<br />
Candolle, Prodr. 4: 402. 1830. Macrocnemum<br />
venosum Ruiz & Pavon, Fl. Peruv. 2: 49, pl. 190b.<br />
1799. Type. Peru. (Acomayo?, Pati?), 1793-1796,<br />
Tafalla & Manzanilla s.n. (lectotype, MA, here<br />
selected; photo-MA at F, GH, MO, NY, TEX,<br />
VEN) = not Condaminea!.<br />
There are two specimens of this species preserved<br />
at MA (both specimens without date or locality) and<br />
annotated by Pav6n: one as "Macrocnemum venosum<br />
Sp. P1. Fl. Per." and the other as "Macrocnemum<br />
venosum Fl. P." I select the former specimen as lectotype.<br />
Both specimens were probably collected by<br />
Tafalla and Manzanilla in 1793-1796, after Ruiz and<br />
Pav6n returned to Spain (see discussion under Condaminea<br />
corymbosa, and Estrella, 1991). Standley<br />
(1936) apparently overlooked this species, since it was<br />
not cited in his treatment of Peruvian <strong>Rubiaceae</strong>. I<br />
cannot position this species with certainty, but from<br />
the detailed photos obtained from MA and de<br />
Candolle's (1830) brief description, it is not a Condaminea.<br />
PICARDAEA<br />
4. Picardaea Urban, Symb. Ant. 3: 376. 1903.<br />
Grisebach, Cat. P1. Cub. 124. 1866; Urban, Symb.<br />
Ant. 7: 391. 1912; Standley, N. Amer. Fl. 32: 6-<br />
7. 1918; Liogier, Fl. Cuba 5: 26. 1963; Liogier,<br />
Fl. Espafiola 7: 355, 357. 1995. Type species.<br />
Picardaea haitiensis Urban [= Picardaea cubensis<br />
(Grisebach) Britton ex Urban].<br />
Shrubs to trees, much-branched shrubs, rarely<br />
single-stemmed trees; bark grayish brown, smooth.<br />
Stipules interpetiolar, adnate to the petioles, triangular,<br />
persistent. Leaves petiolate, lanceolate to oblanceolate,<br />
acute at base, obtuse and usually short-acuminate<br />
at apex, chartaceous; domatia often present.<br />
Inflorescences terminal, solitary, short pedunculate,<br />
corymbose trichotomous panicle terminating in lax 3-<br />
flowered cymules. Flowers protandrous, hypanthium<br />
obconical to obovate. Calyx cupular, truncate to<br />
lobed; lobes (when present) 4-5, rounded to ovate;<br />
calyx caducous, leaving a white scar on mature capsules.<br />
Corolla campanulate, aestivation valvate with<br />
contact zone, fleshy when fresh, red to purple outside,<br />
yellowish white inside; tube cylindrical, expanding<br />
distally; lobes 5, erect, triangular. Stamens 5, partially<br />
exserted, subequal; anthers dorsifixed near the base,<br />
opening by longitudinal splits. Pollen tricolporate,<br />
exine reticulate. Style exserted; style branches narrowly<br />
oblong, reversed at maturity. Ovary 2-celled,<br />
obconical, glabrous, placentation axile; ovules many<br />
in each locule, horizontally inserted; immature fruits<br />
green, semi-fleshy. Capsules oblong, woody; dehiscing<br />
loculicidally, the halves remaining attached at base<br />
and at apex, disk septicidal dehiscence absent even<br />
in old capsules. Seeds irregularly 3-4-angular, horizontally<br />
compressed, testa reticulate.<br />
The combination of fleshy valvate corollas, cap-<br />
sules that dehisce loculicidally with the halves remain-<br />
ing attached at apex and at base (like in<br />
Macrocnemum, Cinchoneae), and horizontal minute<br />
unwinged seeds make this genus unique in the Rubi-<br />
aceae.<br />
The floral morphology of Picardaea is similar to<br />
that of Condaminea, but its reduced inflorescences<br />
resemble those of Macbrideina (<strong>Rondeletieae</strong>). Its<br />
capsule dehiscence is similar to that of Macrocnemum<br />
(Cinchoneae), and its seed morphology is similar to<br />
that of <strong>Rustia</strong>, Pogonopus, and several other genera<br />
of the <strong>Rondeletieae</strong>. Because the seeds of Picardaea<br />
are minute, unwinged, and horizontally attached, this<br />
genus cannot be placed in the Cinchoneae as defined<br />
by Andersson and Persson (1991), which have seeds<br />
vertically attached and wings strongly bipolar. This<br />
genus is therefore included in <strong>Rondeletieae</strong> s.l.<br />
Taxonomic History<br />
Urban (1903), dedicated the name of this genus to<br />
Luis Picarda (1848-1901), botanical collector in<br />
Cuba, who collected the specimen selected as lecto-<br />
type ofPicardaea haitiensis. He separated this genus<br />
from <strong>Rustia</strong> and Condaminea as follows: "Ex affini-
118<br />
Flora Neotropica<br />
tate Condamineae DC. et <strong>Rustia</strong>e Klotzsch, quarum<br />
prior calycis limbo lobato v. dentato circumcisse<br />
deciduo, corollae fauce villosa, ramulis compressis,<br />
stipulis magnis elongatis 2-partitis chartaceis, cymis<br />
trichotome corymbosis, ulterior antheris basifixis,<br />
poro v. rimula terminali dehiscentibus, stipulis amplis<br />
elongatis deciduis discrepat."<br />
Urban (1912) added a second species to Picardaea<br />
by transferring Macrocnemum cubense Griseb., following<br />
a suggestion he encountered in an unpublished<br />
manuscript by Britton. Until recently, Picardaea has<br />
been treated as genus of two species (Standley, 1918;<br />
Liogier, 1963, 1995), but it is here recognized as<br />
monotypic (see discussion below).<br />
1. Picardaea cubensis (Grisebach) Britton ex Urban,<br />
Symb. Ant. 7: 391. 1912. Macrocnemum cubense<br />
Grisebach, Cat. PI. Cub. 124. 1866. Type. Cuba.<br />
Guantanamo: Rio Yumuri, 17 Sep 1860-64 (fl),<br />
Wright 2661 (holotype, GOET; isotypes, G, GH,<br />
HAC, K, P). Figs. 3M, 4E, 8F, 15G,H, 51A-D, 52<br />
Picardaea haitiensis Urban, Symb. Ant. 3: 377. 1903.<br />
Type. Haiti. Petionville, 500-600 m, Aug 1893 (fl),<br />
Picarda 1129 (B*; lectotype, GH (fragment), here<br />
selected).<br />
Shrub 4-6 m tall, exceptionally a tree to 15 m tall,<br />
commonly much-branched shrubs, rarely singlestemmed<br />
trees. Leafy branchlets glabrous, terete,<br />
dark-green; older branches terete, grayish brown; lenticels<br />
sparse, small. Stipules shallowly triangular,<br />
often short-acuminate, glabrous outside, glabrous with<br />
basal colleters inside, 1.5-3 x 2.5-3.5 mm, dark green.<br />
Leaves 8.2-16.5 x (3-)4-12 cm, L/W 1.5:1 to 2.5:1;<br />
oblanceolate to obovate, cuneate to acute at base,<br />
obtuse to round at apex, sometimes tapering to a short<br />
acumen to 15 mm long; grass-green above and below,<br />
foliaceous; drying olive-green, stiff-chartaceous; glabrous<br />
above and below; primary and secondary veins<br />
glabrous, prominent below, secondary veins 8-12<br />
each side; tertiary veins openly reticulate; petioles 7-<br />
15 mm long, 0.7-1 mm thick, adaxially concave to<br />
flattened; domatia (when present) a tuft of yellowish<br />
hairs in the axils of secondary veins. Inflorescences<br />
corymbose trichotomous paniculate, with opposite<br />
decussate lateral branches; (5-)7-15 x angular; bracteoles subtending flowers ca. 0.5 mm<br />
long, deltoid. Flowers pedicellate, pedicels (1-)2.5-<br />
4.5 mm long, glabrous to minutely puberulent; hypanthium<br />
obconical, 10-12 x 5-6 mm, glabrous to<br />
minutely puberulent; flower buds clavate. Calyx<br />
cupular, truncate to shallowly lobed (or 5 minute<br />
teeth), 2-3 x 7-10 mm, glabrous. Corolla campanulate,<br />
3.2-3.5 cm long, reddish purple (to pinkish) outside,<br />
yellowish white inside, camose when fresh; tube<br />
basally cylindrical, expanding distally, 10-12 mm<br />
long, 5-7 mm wide at base and 15-25 mm wide at<br />
the orifice, glabrous outside; pilose below filament<br />
attachments, glabrous above, inside; lobes 5, ca. 2/5<br />
of corolla length, 10-16 x 7-9 mm, triangular, glabrous<br />
throughout, minutely papillose at margins inside.<br />
Stamens 5, subequal, attached 10-12 mm from<br />
the base of the tube; filaments 9-12 mm long, glabrous,<br />
distally terete, basally flattened-adnate to tube;<br />
anthers narrowly elliptic, 5-7.5 x 1-1.5 mm, dorsifixed<br />
near the base, base rounded, smooth throughout.<br />
Pollen exine reticulate, columellate. Style exserted,<br />
25-30 mm long, glabrous; style branches narrowly<br />
oblong, 1.3-2 x 0.3-0.4 mm, stigmatic surface microscopically<br />
(40x) papillose. Capsules oblong, acute<br />
to obtuse at base, apex truncate, 15-19 x 6.5-8 mm,<br />
dark brown to black at maturity, with 3-5 parallel ribs,<br />
glabrous to minutely puberulent; disk glabrous. Seeds<br />
0.45-0.64 x 0.22-0.36 mm; irregularly 3-4-angular,<br />
horizontally compressed, beige, not winged, truncate<br />
at apex, testa reticulate.<br />
Distribution (Fig. 52) and ecology. In primary or<br />
secondary forests, and in remnants of thickets on<br />
cliffs, usually in proximity of rivers, in limestone soil,<br />
50-900 m, of E Cuba and Hispaniola. Flowering specimens<br />
were collected in January, February, July, August,<br />
October, and November. Fruiting specimens were<br />
collected in January, February, March, and August.<br />
Conservation status. This is a rare species much<br />
threatened by ongoing destruction of the natural vegetation<br />
of Hispaniola, and is encountered as a shrub<br />
in remnant forests on limestone cliffs (rarely as a tree).<br />
The reproductive biology of this species has not been<br />
studied.<br />
Specimens examined: CUBA. GUANTANAMO: Valley of<br />
7-10 cm, lat- Rio Yumuri, Baracoa, Feb 1941 (fl-fr), Le6n & Matos 19644<br />
eral branches 1-2 pairs, basal portion of axis not (GH, HAC, US); Altos del Yumuri, Aug 1939 (fl), Le6n &<br />
Victorin 17235 (GH, HAC, NY, TEX);<br />
branched (when present) up to 6 cm long; rachis<br />
Gorge of Rio Yamuri,<br />
7-9 Dec 1910 (fl), Shafer 7850 (NY, US); Valley of Rio<br />
decussately compressed, rachis and branches glabrous Yumuri, Baracoa, 16 Jan 1960 (fl), Liogier & Acuna 7673<br />
to minutely puberulent; distal cymules usually 3-flow- (HAC); Valley of Rio Yumuri, Baracoa, 25 Nov 1914 (fl),<br />
ered; distal bracts 0.5-1.5 x ca. 1 mm, narrowly tri- Ekman 3642 (S).
Systematic Treatment of the Genera Studied 119<br />
- .i. , .<br />
. .<br />
FIG. 51. Picardaea cubensis (A-C from Wright 2661, GH, isolectotype; D from Le6n 19644, GH). A. Habit of inflo-<br />
rescence with terminal leaves and foliage leaf. B. Open flower, with old anthers and exserted style. C. Open anther, dorsal<br />
view. D. Mature open capsule, showing typical dehiscence.
120 Flora Neotropica<br />
_ 70- 60<br />
- -A----- ----- --.-_ -<br />
--- --<br />
I!0 ^"W 1 IO'<br />
*?r3'~~~. ~F. Di I ..i<br />
.<br />
HAITI. Massif du Nord, Port-de-Paix, More Fourrise,<br />
400 m, 14 Aug 1925 (fl-fr), Ekman 4641 (A, G, GH[2], NY,<br />
US); Massif des Matheux, Grand-Bois, rd. Cormillon-<br />
Thamazeau, 800 m, 19 Mar 1926 (st), Ekman 5772 (GH, S);<br />
Massifde La Hotte, Miragoane, Jul 1927 (fl), Eyerdam 180<br />
(US); Massif de la Hotte, Group More Rochelois, Mira-<br />
goane, path Le Brum-Quatre Chemins, 700 m, 30 Mar 1927<br />
(fr), Ekman 7919 (G, S); Massifde la Selle, Croix-des-Bou-<br />
quets, Badeau, Trou-a-1'Eau, 1000 m, 15 Mar 1927 (fr),<br />
Ekman 7859 (S); Massifde la Selle, Gd. Goave, Papelle, 29<br />
Nov 1927 (fl), Ekman 9384 (S); Massif de la Hotte, dept.<br />
Sud, Citadelle des Platons, 30 km NNW from Torbec,<br />
18?16'N, 73058'W, 750 m, 10 Mar 1983 (fr), Zanoni et al.<br />
25634 (NY).<br />
DOMINICAN REPUBLIC. Prov. Barahona, La Fili-<br />
pana, Loma Travesia, Barahona, 700 m, 5 Jan 1977 (fl),<br />
Liogier & Liogier 26174 (GH, NY); Prov. Barahona, Las<br />
Filipinas, 450 m, Nov (fl), Fuertes 649 (G[2], NY); Prov.<br />
Barahona, Sierra de Bahoruco, Los Naranjos, confluence of<br />
Rio Bahoruco with Rio Brazo Seco, 18?05'N, 71?08'W, 50<br />
m, 17 Jan 1986 (fl-fr), Zanoni & Pimentel 35991 (NY, US);<br />
Prov. Barahona, Sierra de Bahoruco, Arroyo La Travesia,<br />
18?07'N, 71?07'W, 850-900 m, 12 Jan 1988 (fr), Zanoni et<br />
al. 40662 (NY); Prov. Barahona, Sierra de Bahoruco, Ar-<br />
royo El Maniel, 2 km from Los Patos de Paraiso, 17?58'N,<br />
71013'W, 350-470 m, 17 Jan 1985 (fl), Zanoni etal. 33125<br />
80<br />
FIG. 52. Distribution of Picardaea cubensis.<br />
(NY); Polo, Los Arroyos, 800 m, 28 Feb 1969 (fr), Liogier<br />
14353 (GH, NY); Prov. Pedernales, 13 km N of Pedernales,<br />
along Rio Mulito, 18009'N, 71?46'W, 230 m, 21 Oct 1991<br />
(fl), Thompson et al. 9859 (CM).<br />
Urban (1912) transferred Macrocnemum cubense<br />
to Picardaea, separating it from P. haitiensis by its<br />
yellow-white corolla [as reported on Wright 2661],<br />
stipule shape, and leaf size and shape. Standley (1918)<br />
also maintained the two names. Recent collections [Leon<br />
& Victorin 17235, Ekan 4641,Liogier &Liogier26174]<br />
showed that the corollas of Picardaea are purple-red<br />
outside and yellowish white (rarely reddish white)<br />
inside (see type label data, above), the stipules are<br />
constantly interpetiolar and adnate to the petioles, deltoid<br />
and variably acuminate, and the leaf blades vary<br />
in size and shape throughout the range of the species.<br />
For the above reasons, the characters that were used<br />
to separate P. cubensis from P. haitiensis are no longer<br />
valid, and the two taxa are here treated as synonymous,<br />
the previous having taxonomical priority.<br />
Among the duplicates of Wright 2661 examined,<br />
only the one present at the Herbarium Grisebachianum<br />
(GOET) bears Grisebach's handwriting and is here<br />
20<br />
30
Systematic Treatment of the Genera Studied 121<br />
treated as the holotype of Macrocnemum cubense (= Shrubs, rarely treelets, much-branched; outer<br />
P. cubensis). The label [manu Wright] has the follow- branches often scandent, main trunk (when present)<br />
ing description: "A small tree or large bush 10-20 feet short. Stipules interpetiolar, free at base, sometimes<br />
on farallones. Corolla valvate, thick greenish tinge adnate to the petioles, small, deltoid to shallowly triwith<br />
red externally and dull red within."<br />
angular, persistent; stipules in apical buds with axil-<br />
The typification of Picardaea haitiensis is diffi- lary colleters that secrete a sticky yellow resin. Leaves<br />
cult. The specimens of Picarda 1129, examined by petiolate, elliptic to obovate, foliaceous; petioles short,<br />
Urban at Berlin, were destroyed; no photos of that thickened at base; domatia absent, or tuft domatia with<br />
specimen were found. Among the herbaria from which sparse hairs (rarely pocket domatia). Inflorescences<br />
loans were received, only one leaf-fragment was lo- terminal, frondose, pyramidal, laxly paniculate, each<br />
cated at the Harvard herbarium (GH). It is unlikely lateral branch subtended by leaf-like bracts, terminatthat<br />
other duplicates of Picarda 1129 are extant, and ing in a small cyme. Flowers protandrous; hybecause<br />
the leaves of the GH specimen are typical of panthium obconical to obovoid (rarely turbinate).<br />
this genus, I have selected the leaf-fragment at GH as Calyx extremely reduced, caducous; lobes 5; 1-5<br />
lectotype of P. haitiensis.<br />
calyx lobes expanded into colorful calycophylls in<br />
some flowers. Calycophylls shortly stalked and<br />
blades ovate to obcordate to reniform. Corolla tubular<br />
with short<br />
POGONOPUS<br />
lobes, membranaceous or fleshy when<br />
fresh; tube cylindrical, basally constricted, with a ring<br />
5. Pogonopus Klotzsch, Monatsber. Koenigl. Preuss. of hairs at base inside; lobes 5, erect to spreading, tri-<br />
Acad. Wiss. Berlin 1853: 500. 1853; de Candolle, angular to ovate; aestivation valvate-reduplicate with<br />
Prodr. 4: 403. Sep 1830; A. Richard, Mem. Rubiac. contact zones. Stamens 5, exserted well above the<br />
199. Dec 1830; Weddell, Ann. Sci. Nat. 4: 66-71.<br />
corolla, attached near the base of the corolla tube (just<br />
1854; Linden & Planchon, Bot. Zeit. 12:365. 1854; above the constriction); filaments thin, with a tuft of<br />
Orsted, Centr.-Amer., fig. 13. 1863; Hooker in hairs at base; anthers elliptic-oblong, dorsifixed at<br />
Bentham & Hooker, Gen. PI. 2: 47. 1873; medial zone, dehiscing by longitudinal slit. Pollen tri-<br />
Schumann in Martius, Fl. Bras. 6(6): 264-266, fig. colporate, exine reticulate. Style exserted well above<br />
126. 1889; Schumann in Engler & Prantl, Nat. the corolla; style branches linear to oblong. Ovary 2-<br />
Pflanzenfam. 4(4): 21, fig. 6P. 1891; Werham, celled, turbinate, glabrous, placentation axile; ovules<br />
J. Bot. 50: 241-242, tab. 520. 1912; Standley, Publ.<br />
many in each locule, horizontally inserted; immature<br />
Field Columbian Mus., Bot. Ser. 7: 25, 44, 277, fruits green and semi-carose when fresh. Capsules<br />
370.1930, 1931; Sandwith, Kew Bull. 1949:258- obovoid, oblong to globose, truncate at apex, often<br />
259. 1949; Steyermark, Acta Bot. Venez. 6: 108- with white lenticels; dehiscing loculicidally, disk sep-<br />
110. 1971; Steyermark in Lasser & Steyermark, ticidal dehiscence present in old capsules. Seeds<br />
Fl. Venez. 9:235-241, fig. 36. 1974; Dwyer, Ann.<br />
many, horizontal, irregularly shaped, 3-5-angled,<br />
Missouri Bot. Gard. 67: 329-333, fig. 69. 1980; compressed.<br />
Burger & Taylor in Burger, Fl. Costaricensis,<br />
Fieldiana, But. n.s. 33: 218, fig. 16. 1993. Type Pogonopus is commonly encountered as beautiful<br />
species. Pogonopus ottonis Klotzsch [= P. speci- multi-stemmed shrubs (rarely single-stemmed trees),<br />
osus (Jacquin) K. Schumann var. speciosus]. with highly ornamental red-purple calycophylls. It is<br />
Carmenocania Wemham, J. Bot. 50:241. 1912. Type spe- easily recognizable because of its long-tubular pinkcies.<br />
Carmenocaniaporphyrantha Wemham [= Pogon- red flowers, small deltoid stipules with a tuft of hairs<br />
opus speciosus (Jacquin) K. Schumann var. speciosus]. between them, and small capsules with horizontally<br />
Chrysoxylon Weddell, nom. poster. homonym., Monogr. inserted seeds. Its seeds are minute, non-winged, trun-<br />
Cinch. 100. 1849, species typica Chrysoxylonfebri- cate at the apex, and posses a reticulate and minutely<br />
fugum Weddell, nom. inval. [= Pogonopus tubulosus tuberculate exotesta (within the primary reticulation).<br />
(A. Richard in A. P. de Candolle) K. Schumann]; Its bark is often reported to be a good remedy for<br />
non Chrysoxylon Casaretto, Nov. Stirp. Dec., decas<br />
malaria and intermittent fevers, and more<br />
7: recently for<br />
59. 1843 (= Plathymenia Bentham, Mimosaceae).<br />
its antitumor<br />
Howardia Weddell, Ann. Sci. Nat. 4(1): 66, fig. 10. 1854;<br />
properties (for additional information see<br />
non Howardia Klotzsch, nom. under<br />
poster. homonym., Uses).<br />
Monatsber. Deutsch. Akad. Wiss. Berlin 1859: 584, Pogonopus (see Figs. 53, 55, 56, 57) is closely re-<br />
607. 1860 [= Aristolochia Linnaeus, Aristolochi- lated to Pinckneya, from southeastern North America.<br />
aceae]. Type species. Howardiafebrifuga (Weddell) The latter differs from the former in having corolla<br />
Weddell [= P. tubulosus (A. Richard) K. Schumann]. cream-white, its throat without a ring of hairs (present
122 Flora Neotropica<br />
in Pogonopus), corolla lobes long-narrow pubescent var. speciosus); and H. richardi (based on Macroinside,<br />
and coiling outward at anthesis (vs. spreading), cnemum tubulosum A. Rich.) and H. febrifuga (based<br />
seeds arranged in two vertical ranks, with orbicular on Chrysoxylonfebrifugum), both using Bolivian colwing<br />
(vs. many minute unwinged seeds), and exotesta lections (and both = P. tubulosus). Weddell (1854)<br />
secondarily reticulate (within the main reticulation). also commented that Calycophyllum tubulosum (A.<br />
The distribution of Pogonopus is an interesting Rich. in DC.) DC. shared many affinities with his Howcase<br />
of geographical vicariance (see Fig. 54). Re- ardia and, instead of making a new combination, he<br />
stricted to tropical seasonal forests, this genus occurs reduced this species under synonymy of his H. richardi.<br />
in two areas of distribution that mirror to each other Hooker (1873) established the tribe Condamiwith<br />
respect to the equator. In the northern hemisphere neeae, and in his third subtribe Pinckneyinae he placed<br />
Pogonopus ranges from Chiapas (southern Mexico) Pogonopus and Pinckneya.<br />
to northern Colombia, from 6?N to 16?N; while in the Baillon (1880) included Pogonopus under Pincksouthern<br />
hemisphere it occurs from central Peru to neya, simply stating (translated from French): "In<br />
northern Argentina, from 9?S to 22?S. Pogonopus also those named Pogonopus, the corolla, instead of beoccurs<br />
along elevational gradients and is usually en- ing tomentose internally, is glabrous. The fruit is<br />
countered at medium elevations of(50-)500-1400 m. ovoid, whilst in the true Pinckneya [as Eupinckneya]<br />
I attribute the occurrence of Pogonopus in the low- it is more globular and subdidymous. They are shrubs<br />
lands to its introduction into cultivation for its orna- of the two Americas, with rather large and showy<br />
mental and medicinal properties. Pogonopus is here flowers in terminal or axillary clusters of cymes."<br />
treated as a genus of three species and two varieties. Wernham (1912) described Carmenocania, a genus<br />
that he included in the Mussaendeae. The type<br />
Taxonomic History<br />
collection, Schlim 755, is from the vicinity of the city<br />
of Carmen, in the province of Ocafia (hence the ge-<br />
Pogonopus was established by Klotzsch in 1853; neric name), Venezuela [and not Colombia as reported<br />
its name was derived from the Greek words iwoycov<br />
by Wernham (1912), Standley (1930a), and Sandwith<br />
[pogon = beard] and oOG [pous = feet], because of<br />
(1949)]. Wernham apparently overlooked that this was<br />
its basally bearded filaments. The first species dethe<br />
same collection previously used by Weddell<br />
scribed under this genus was P. ottonis [= P. speciosus<br />
(1854) to establish Howardia grandiflora.<br />
(Jacq.) K. Schum. var. speciosus], from material col-<br />
Standley (1918) reduced Chrysoxylon and<br />
lected in Venezuela, with the specific name dedicated<br />
Howardia to synonymy under Pogonopus, but subto<br />
its collector [Otto 901]. Klotzsch (1853) also noted<br />
sequently he (Standley, 1930a) listed Pogonopus (in<br />
that another species (from Costa Rica) should be inthe<br />
Condamineeae) and Carmenocania (in the<br />
cluded in Pogonopus, namely Macrocnemum exsert-<br />
Mussaendeae) without noting their similarities.<br />
um Orsted (Orsted, 1852), but he did not make the of-<br />
Sandwith (1949) recognized the mistakes made both<br />
ficial transfer. When Klotzsch (1853) described P.<br />
by Wernham (1912) and Standley (1930a) and synottonis,<br />
he was perhaps unaware that this species was<br />
onymized Carmenocania under Pogonopus.<br />
already described by Jacquin (1797) as Macrocnemum<br />
Steyermark (1971) divided Pogonopus speciosus<br />
speciosus. The latter was transferred to Pogonopus into three subspecies: exsertus, speciosus, and sandmuch<br />
later by Schumann (1889).<br />
withianus. He later reduced subsp. sandwithianus to<br />
Augustin Pyramus de Candolle (1830) transferred<br />
a variety under P. speciosus subsp. speciosus<br />
Macrocnemum tubulosum A. Rich. in DC. [= Pogon-<br />
(Steyermark, 1974). In the present treatment P. exopus<br />
tubulosus (A. Rich. in DC.) K. Schumann] to<br />
sertus is returned to specific rank (Orsted, 1863) and<br />
Calycophyllum, which in turn was placed next to<br />
the subspecific divisions are no longer maintained. As<br />
Pinckneya [as "Pinkneya"], within his Cinchoneae.<br />
here treated, Pogonopus consists of three species, one<br />
Weddell (1849) described Chrysoxylon, an invalid<br />
with two varieties, collectively ranging from southname<br />
because this generic epithet was previously used<br />
ern Mexico to northern Argentina.<br />
by Casaretto (1843) for a mimosoid genus. Realizing<br />
the invalidity of his Chrysoxylon, Weddell renamed<br />
this genus Howardia, ignoring (purposely or otherwise)<br />
the fact that Klotzsch (1853) had meanwhile<br />
described it as Pogonopus (the valid name for this<br />
genus). Under Howardia, Weddell (1854) described<br />
four species: H. caracasensis and H. grandiflora, both<br />
using Venezuelan collections (and both = P. speciosus<br />
Key to the species of Pogonopus<br />
1. Corolla lobes erect to narrowly spreading;<br />
corolla tube narrowly cylindrical, not expanded<br />
or ampliate at the orifice, mem branaceous<br />
when fresh, glabrous to minutely<br />
puberulent outside.
Systematic Treatment of the Genera Studied 123<br />
2. Corollas 25-30 mm long, with erect lobes; two halves. Leaves<br />
tube cylindrical for the whole<br />
9-19(-25) x (3-)5-7(-10) cm,<br />
length;<br />
lobes '/s of corolla length; blades of lanceolate,<br />
calycoelliptic<br />
to elliptic-obovate, acute at base,<br />
phylls
124 Flora Neotropica<br />
IP<br />
F I<br />
FIG. 53. Pogonopus exsertus (A-D from Nee 9125, MO; E from Croat 8959, MO). A. Habit of inflorescence. B. Open<br />
flower with exserted anthers. C. Close up of calyx . D. Detail of external pubescence of corolla lobe. E. Detail of intemal<br />
glandular pubescence of corolla lobe. F. Mature capsule.
Systematic Treatment of the Genera Studied 125<br />
....... _ --[-.-..... .....--.---.. ...<br />
dp<br />
~ or- 1lil 1- - #- A- ---I-- __ 1<br />
FIG. 54. Distribution of Pogonopus exsertus (circles), P. speciosus var. speciosus (diamonds), P. speciosus var.<br />
sandwithianus (squares) and P. tubulosus (triangles).<br />
callous zone, at pilose ring); filaments 20-28 mm<br />
long, with a basal tuft of hairs; anthers elliptic-oblong,<br />
1.9-2.3 x ca. 0.8 mm, dorsifixed at medial zone, with<br />
a narrow space between the thecae, base and apex<br />
rounded. Pollen exine reticulate. Style exserted, 27-<br />
35 mm long, microscopically (40x) papillose, green;<br />
style branches linear to oblong, 2-2.5 x 0.5 mm, re-<br />
versed, stigmatic surface microscopically (40x) glan-<br />
dular. Capsules obovoid, base acute, apex shallowly<br />
hemispherical, 5-8 x 4-6 mm, with sparse lenticels,<br />
dark brown, minutely puberulent below the disk; disk<br />
glabrous, black, with lenticels, with a white tubercule<br />
at the point of attachment of the style. Seeds 0.38-<br />
0.58 x 0.23-0.35 mm.<br />
Distribution (Fig. 54) and ecology. Seasonal ev-<br />
ergreen and deciduous forests, secondary tropical<br />
moist forest, and relative remnants, in steep wooded
126 Flora Neotropica<br />
slopes and thickets, usually near small creeks or in<br />
river banks, 50-1000 m, from S Mexico to NW Colombia.<br />
Flowering specimens were collected in January,<br />
February, and July through December, but especially<br />
from September through January. Fruiting<br />
specimens were collected in January, March, September,<br />
November, and December.<br />
Representative specimens examined. MEXICO.<br />
Shattuck 688 (MO); N of Paraiso, near Gaillard, 22 Nov 1970<br />
(fl), Croat 12652 (MO, NY); Rio Pedro Miguel, near E<br />
Paraiso, 7 Jan 1924 (fl), Standley 29972 (A, GH, US); between<br />
Juan Mina and El Vigia, 11 Jan 1911 (fr), Pittier 2396<br />
(NY); tributary of Rio Chagres, 3 km SW of Cerro Brewster,<br />
14 Dec 1967 (fl), Lewis et al. 3392 (COL, UC, VEN);<br />
Serrania de Maje, trail along Rio Ipeti Grande, 300 m,<br />
08?55'N, 78?31'W, 28 Jan 1984 (fl), Churchill & Nevers<br />
4446 (MO[2]); Forest Preserve, jet. Boyd-Roosevelt Hwy.<br />
with Madden Rd. 1 km from Madden Dam, 1967, Mori 421<br />
CHIAPAS: Rio Testecapa, 10 km SE of Mapastepec, 180 m,<br />
24 Dec 1972 (fr), Breedlove & Thorne 30715 (NY); Mal<br />
(MO). UNCERTAIN LOCALITY: Maume and Gorgone, Jan 1858<br />
(fr), Wagner 3495 (M), 3496 (M).<br />
Paso, Carrillo Puerto, 250 m, 6 Oct 1984 (fl), Ventura & COLOMBIA. ANTIOQUIA: Uraba, Dabeiba, Riosucio,<br />
L6pez 424 (MO); Escuintla, 15 Nov 1947 (fl), Matuda 16811<br />
(NY); Huixtla, Rio Huixtla, 4 Dec 1985 (fl), Ventura &<br />
350 m, 11 Jan 1947 (fl), Uribe-Uribe 1441 (COL, F, NY);<br />
rd. Antioquia-Anza, Rio Cauca, 06?25'N, 75?52'W, 2 Nov<br />
L6pez 2856 (NY); Azulejo, Margarita, 250 m, 7 Jan 1950<br />
(fr), Matuda 18748 (MO).<br />
GUATEMALA. ALTA VERAPAZ: Finca Mocca, 750 m,<br />
25 Nov 1919 (fl), Johnson 57 (NY, US); Tucuru, 18 Aug<br />
1979 (fl), Boeke & Utzischneider 2925 (MO); Pancajche, 18<br />
1988 (fl), Zarucchi et al. 7044 (F, NY). CHOC6: Mun.<br />
Riosucio, Parque Natural Los Katios, Camino Peye-Tilupo,<br />
60 m, 22 Nov 1976 (fl), Le6n 455 (COL, MO); Parque<br />
Nacional Los Katios, Quebrada de La Tigra, 18 Jan 1983<br />
(fr), Zuluaga 1184 (COL).<br />
Nov 1920 (fl-fr), Johnson 1022 (US). SAN MARCOS: Finca<br />
Armenia, San Rafael, at the base of Cuesta to Carrizal (cul-<br />
CULTIVATED. UNITED STATES: FLORIDA: Miami,<br />
Fairchild Tropical Garden, 24 Jan 1969 (fl), Gillis 7535 (A),<br />
tivated), 1300-1600 m, 9 Aug 1980 (fr), Dwyer 15265 (MO);<br />
near Malacatan, 400 m, Nov-Dec 1940 (fl), Grant 630 (A,<br />
24 Nov 1971 (fl), Gillis 11168 (A). HAWAII: Oahu, U.S.<br />
Nursery, 4 Jan 1937 (fl), E.L.C. s.n. (A). GUATEMALA.<br />
GH). SANTA ROSA: Plains N of Los Cerritos, rd.<br />
Chiquimulilla-El Ahumado, 75 m, 7 Dec 1940 (fl), Standley<br />
79549 (US). SUCHITEPtQUEZ: Cocales (cultivated), 215 m,<br />
ALTA VERAPAZ: Near Tucuri, 500 m, Oct 1917 (fl), Popenoe<br />
802 (A, GH). HONDURAS. Lancetilla Experimental Station,<br />
11 Jul 1934 (fl), Yuncker 4529 (NY). COSTA RICA.<br />
5 Jan 1939 (fl), Standley 62055 (A).<br />
PUNTARENAS: Golfito nursery, 23 Aug 1950 (fl), Allen 5621<br />
EL SALVADOR. AHUACHAPAN: Near Salto de Atehue- (DS). CUBA. SANTA CLARA: Cienfuegos, Soledad, Harvard<br />
cia, 600 m, 22 Jan 1947 (fr), Standley & Padilla 2885 (MO). Tropical Garden, 1 Feb 1932 (fr),Jack 8430 (A[2], NY, US),<br />
SAN MIGUEL: La Libertad, vic. of La Laguna, 830 m, Cruz 11 Mar 1931 (fr), Jack 8173 (A). PUERTO RICO. Rio<br />
WB-00482 (MO). SAN SALVADOR: La Cebadilla, 1922 (fl), Piedras, Experimental Station, 1 Dec 1964 (fl), Wagner 711<br />
Calder6n 1225 (GH, MO, NY, US); on Rt. 382, W of Sanso- (A), 30 Aug 1965 (fr), Wagner 897 (A); Humacao Playa, 22<br />
natejct., 6 Sep 1964 (fl), Elias et al. 583 (MO). SAN VICENTE: Jan 1966 (fl), Howard 16044 (A); Pennock's Nursery, Hato<br />
Vic. of San Vicente, 2-11 Mar 1922 (fr), Standley 21672 Rey, Howard & Nevling 16910 (A).<br />
(GH, US). SANTA ANA: Trail to Cerro Verde, 1000 m, 30 Sep<br />
1959 (fr), Montalvo 3849 (MO); E side of Lake Coatepeque, Local names and uses. El Salvador: chorcha de<br />
790 m, 2 Dec 1970 (fl), Harmon & Fuentes 4965 (MO, NY).<br />
COSTA RICA. GUANACASTE: Colonia Carmina, 100 m,<br />
Feb 1912 (fl), Jimenez 541 (US). PUNTARENAS: Cant6n de<br />
Osa, Rio T6rralba, E of Palmar del Norte, 100 m, 08?56'N,<br />
83?20'W, 3 Jan 1990 (fl), Hammel et al. 17726 (MO). SAN<br />
gallo (Calderon 1922). Guatemala: quina (Standley<br />
79549). Panama: chibigui (Bristan 1425), madrofio.<br />
Reportedly a local remedy for malaria in Guatemala<br />
(Standley 79549).<br />
JosE: Vic. of El General, cliff above Rio Pacuan, 610 m, Jan<br />
1939 (fl), Skutch 3950 (A, MO, NY, US[2]); Puriscal, San<br />
Martin, 09?44'N, 84024'W, 800 m, 29 Nov 1989 (fl), Jimenez<br />
et al. 751 (MO); trail San Miguel de Turricanes-Piedras<br />
Negras de Mora, Rio Virilla, 5 Dec 1985 (fl), Gdmez et al.<br />
Pogonopus exsertus is a very elegant, glabrescent<br />
shrub with slender branches, easily distinguishable<br />
from the other two species in having thin, delicate,<br />
lax inflorescences. It has been found in cultivation<br />
24504 (MO).<br />
PANAMA. COLON: Around Porto Bello, 18 Jan 1911 (flfr),<br />
Pittier 2476 (GH[2], NY, US); ca. 2 km E of Transisthmian<br />
Hwy to Salamanca, 100 m, 19 Dec 1972 (fr), Gentry<br />
6720 (MO, NY); near Salamanca, hydrografic station, Rio<br />
Pequeni, 70-80 m, 15 Dec 1934 (fl), Dodge etal. 16977(BR,<br />
G, GH, NY, US). DARIEN: Banks of Rio Piedras, 26 Jun 1959<br />
(fr), Stern et al. 724 (GH, MO, US); Rio Aruza, 24 Oct 1967<br />
(fl), Bristan 1346 (MO[2]); Parque Nacional del Darien,<br />
ridge between Rio Topalisa and Rio Pucuro, 08?03'N,<br />
77?17'W, 750-850 m, 18 Oct 1987 (fl), Cuadros etal. 3884<br />
(MO). PANAMA: Barro Colorado Island, 6 Jan 1932 (fr),<br />
throughout its natural distribution and elsewhere (see<br />
above). Pogonopus exsertus is well deserving of specific<br />
rank because it differs in many floral and vegetative<br />
morphological characters from its South American<br />
counterparts (P. speciosus and P. tubulosus).<br />
Pogonopus exsertus closely resembles P. tubulosus,<br />
from Peru and southward (rather than P. speciosus,<br />
from the Caribbean coast of Colombia and Venezuela),<br />
in having glabrescent, narrowly cylindrical,<br />
membranous corollas; the former differing from the<br />
latter in having corolla lobes erect (vs. narrowly
Systematic Treatment of the Genera Studied 127<br />
spreading), shorter corollas (to 3 cm long), and smaller bescent hairs; flowers on distal branches in lax<br />
calycophylls that are basally and apically acute. cymules of 9-50 flowers; distal bracts usually leaf-<br />
Pogonopus exsertus is easily distinguished from P. like, petiolate, elliptic-obovate, to 22 x 8 cm, decreasspeciosus<br />
in the latter having thick and fleshy corol- ing in size distally, often reduced to deltoid scales 20las<br />
that are gradually expanded toward the orifice, and 30 x 5-6 mm; bracteoles subtending flowers to 3 x<br />
having reproductive and vegetative parts densely yel- ca. 0.5 mm, linear, densely golden-puberulent. Flowlow<br />
pubescent.<br />
ers pedicellate, pedicels 2-10 mm long, puberulent<br />
At least two specimens of this species [Wagners.n. and pubescent intermixed, to hirsutulous to pilose;<br />
(M), Maume and Gorgone, Jan 1858 (fr); and Hayes hypanthium obconical to obovoid, 2.5-4 x 2-2.8 mm,<br />
164 (BR), Isthmus of Panama, Oct 1861 (fl)] were yellow appressed puberulent, with lenticels; young<br />
annotated by Schumann as "Pogonopus speciosus flower buds minute glandular-pilose (var. speciosus)<br />
(tubulosus) var. panamensis K. Schumann," but this or golden-hirsutulous (var. sandwithianus). Calyx<br />
name was never published.<br />
extremely reduced, 0.3-0.7 x 2.5-4 mm, densely yellow<br />
pubescent; lobes 5, barely distinguishable, narrowly<br />
triangular to linear, 1-3 x 0.5-1 mm; in some<br />
flowers 1-5 calyx lobes expanded into calycophylls.<br />
2. Pogonopus speciosus (Jacquin) K. Schumann in Calycophylls (3-)4.5-6.5<br />
Martius, Fl. Bras. 6(6): 265. 1889.<br />
Figs. 3K,L, 4G, 8C,D, 16C, 54, 55A-E, 56A-F<br />
Shrubs 4-8 m tall, rarely small trees to 10 m tall,<br />
to 30 cm dbh, much-branched, with spreading crown;<br />
main trunk (when present) short, terete; central<br />
branches erect, outer branches often scandent; bark<br />
brown. Leafy branchlets sparsely to densely antrorse<br />
short-pubescent to hirsutulous to hirsute; older<br />
branches sparsely short pubescent or sparsely hirsutulous;<br />
lenticels cream-white, punctiform to linear, to 5<br />
mm long. Stipules deltoid to shallowly triangular,<br />
acuminate, sparsely long pilose (var. speciosus) or<br />
hirsutulous to hirsute (var. sandwithianus), with elongated<br />
colleters (to 0.5 mm long) inside; 3.5-6.5 x 3.5-<br />
5 mm, persistent. Leaves 11-25 x 5-10.5 cm, elliptic<br />
to obovate, cuneate to decurrent at base, acute to<br />
obtuse and acuminate at apex, acumen to 3 cm long;<br />
dark green above, pale green below, stiff-foliaceous;<br />
drying olive-green, stiff-chartaceous; glabrous above,<br />
glabrous to minutely puberulent to hirtellous below;<br />
primary and secondary veins sparsely short appressedpubescent;<br />
secondary veins 9-14 each side, tertiary<br />
veins starting subparallel and subreticulate in the<br />
middle; petioles 0.8-2.2 mm long, 1-1.5 mm thick,<br />
adaxially concave to flattened, minutely puberulent;<br />
domatia absent (var. sandwithianus) or a tuft of sparse<br />
erect to curled hairs, rarely pocket domatia. Inflorescences<br />
laxly paniculate, pyramidal, with opposite<br />
decussate branches, each lateral branch subtended by<br />
leaf-like bracts, terminating in a small cyme; 11-30 cm<br />
long, basal branches 4-20 cm long, lateral branches<br />
2-5 pairs, basal portion of axis not branched 5-13 cm<br />
long; rachis terete, with punctiform to linear lenticels,<br />
rachis and branches sparsely short puberulent to<br />
densely golden-pubescent (to hirtellous), branchlets<br />
of terminal cymes densely puberulent, with few pu-<br />
x (1-)2.5-4.5 cm, blades<br />
ovate to widely ovate, base obtuse to rounded (rarely<br />
cordate), apex acute (with the very tip rounded), palmately<br />
veined (3-5 veins each side), glabrous to glabrescent;<br />
primary and secondary veins sparsely puberulent;<br />
pink, lavender to crimson red; stalks 1.5-2<br />
cm long. Corolla tubular gradually expanded (var.<br />
speciosus, Fig. 55F) or abruptly ampliate (var.<br />
sandwithianus, Fig. 55B) at orifice, 25-34 mm long,<br />
aestivation valvate-reduplicate with contact zone,<br />
thick and fleshy when fresh, rose-pink to pink-red,<br />
sometimes with white-pink lobes; tube subcylindrical<br />
with distal expansion, 20-28 mm long, 2.5-4 mm<br />
wide at base and 3-7 mm wide at orifice, sparsely (at<br />
base) to densely (distally) antrorse puberulent-pubescent<br />
(var. speciosus, Fig. 55G), or golden-hirsute (var.<br />
sandwithianus, Fig. 55C) outside; with a white callous<br />
basal area 5-7.5 mm long, topped by a dense ring<br />
of hispid or golden ascending hairs (ca. 1 mm long),<br />
the medial and superior zones of throat glabrous to<br />
pubescent; lobes 4-5, spreading, 2.9-6.5 x 2.2-5 mm,<br />
narrowly to widely ovate, antrorse yellow pubescent<br />
outside (var. speciosus), or golden hirtellous (var.<br />
sandwithianus), with sparse glandular hairs inside.<br />
Stamens 5, exserted, slightly unequal, attached 6-7 mm<br />
from the base of the tube (just above callous zone, at<br />
pilose ring); filaments 27-37 mm long, white, with a<br />
basal tuft of hairs; anthers elliptic, 2-3 x 0.8-1.1 mm,<br />
dorsifixed at medial zone, base rounded or elliptic,<br />
apex acute or round, with no space between the thecae,<br />
purple to blue when fresh, drying black. Pollen<br />
exine reticulate. Style exserted, 32-40 mm long, glabrous,<br />
green; style branches oblong to oblate, 1.25-<br />
1.5 x 0.5-0.75 mm, reversed, stigmatic surface microscopically<br />
(40x) papillose or with glandular hairs.<br />
Capsules ellipsoid-ovoid (var. speciosus, Fig. 551) or<br />
oblong to subglobose (var. sandwithianus, Fig. 55E),<br />
base rounded, apex round to shallowly hemispheri-
128 Flora Neotropica<br />
FIG. 55. A-E. Pogonopus speciosus var. sandwithianus (from fresh material and Delprete & Apreza 6359, TEX).<br />
A. Habit of inflorescence. B. Flower (note corolla tube abruptly ampliate). C. Detail of external vestiture of corolla.<br />
D. Detail of internal glandular pubescence of corolla lobe. E. Portion mature capsule (note hirsute pubescence). F-I. Pogonopus<br />
speciosus var. speciosus (Williams 12408, F). F. Flower (note corolla tube gradually ampliate). G. Detail of external vestiture of<br />
corolla. H. Detail of internal glandular pubescence of corolla lobe. I. Portion mature capsule (note appressed pubescence).<br />
re<br />
tnn
Systematic Treatment of the Genera Studied<br />
...<br />
129<br />
?~~~~ ? ::, ~' ..? :. .O j.,:.~ ..'.~..~, ? ., '131 1 :~...,:..., __<br />
. . 9'<br />
_@).. .o '-~ _:: ...'
130 Flora Neotropica<br />
glabrous to minute puberulent below, veins<br />
sparsely short appressed-pubescent, secondary veins<br />
sparsely appressed-pubescent below; capsules 9-14 each<br />
ellipsoid-ovoid, 7-8 mm side;<br />
long, golden<br />
petioles 0.8-2.2 mm long, 1-1.5 mm<br />
puberulent;<br />
disk shallowly hemispherical (Venezuela: thick, minutely puberulent; tufts domatia, with sparse<br />
Caribbean coast) ............................. 2a. var. speciosus<br />
erect to curled hairs (rarely pocket domatia). Inflo-<br />
1. Corollas golden-hirsutulous outside; corolla tube rescences pyramidal, laxly paniculate; 11-30 cm long,<br />
abruptly ampliate at orifice, 3-4 mm at base, basal branches 6-20 cm long, lateral branches 2-5<br />
5-7 mm at orifice; lobes 4-6.5 mm long, ca. '/7 pairs, basal portion of axis not branched 5-12 cm long;<br />
of corolla length; leaf blades sparsely to densely rachis and branches sparsely short puberulent, branchgolden-hirtellous<br />
below, veins sparsely to densely lets of terminal cymes densely puberulent, with sparse<br />
hirsutulous to hirsute below; capsules oblong to<br />
subglobose, 6-7 mm long, short pubescent; disk<br />
pubescent hairs intermixed; distal branches lax<br />
with outer elevated ring (do-nut shaped) and cymules of 20-50 flowers; distal leaf-like bracts to<br />
concave (Colombia: Sierra Nevada de Santa Marta 22 x 8 cm, decreasing in size distally, often reduced<br />
and Serrania de Perija) ........... 2b. var. sandwithianus to deltoid scales 20-30 x 5-6 mm; bracteoles subtending<br />
flowers to 3 x ca. 0.5 mm, linear, densely goldenpuberulent.<br />
Flower pedicels 3-10 mm long, puberulent<br />
and pubescent intermixed; young flower buds<br />
2a. Pogonopus speciosus (Jacquin) K. Schumann var. minute glandular-pilose throughout (hairs to 0.5 mm<br />
speciosus. Pogonopus speciosus subsp. speciosus long). Calyx 0.3-0.7 x 2.5-4 mm, densely yellow<br />
var. speciosus Steyermark in Lasser & Steyermark, pubescent; lobes barely distinguishable, narrowly tri-<br />
Flora de Venezuela 9(1): 239. 1974. Chrysoxylon angular, 1-2 x 0.5-1 mm; in some flowers 1-3 calyx<br />
speciosum (Jacquin) O. Kuntze, Revis. Gen. PI. lobes expanded into calycophylls. Calycophylls (3-<br />
278. 1891. Mussaenda speciosa (Jacquin) Poiret )4-6.5 x (1-)2.5-4.5 cm, blades ovate to widely ovate,<br />
in Lam. Encyc. Suppl. 4: 37. 1816. Macrocnemum base obtuse to rounded (rarely cordate), apex acute<br />
speciosus Jacquin, PI. hort. schoenbr. 1: 19, tab. (with the very tip rounded), palmately veined (3-5),<br />
43. 1797. Type. Cultivated in Sch6nbrunn Botani- glabrous to glabrescent; primary and secondary veins<br />
cal Gardens, Vienna, brought from Venezuela, sparsely puberulent; pink, lavender to crimson red;<br />
Caracas, 1787-1788, Bredemeyer s.n. (lectotype, stalks 1.5-2 cm long. Corolla tubular, gradually ex-<br />
W, here selected; isolectotypes, W[3]). Figs. 8C, panded at orifice, 25-32 mm long, rose-pink to pink-<br />
54, 55F-I<br />
red, sometimes with whitish pink lobes; tube cylin-<br />
Pogonopus caracasensis (Weddell) Nichols, Diet. Gard. drical with gradual expansion, 20-28 mm long,<br />
3:175. 1886. Howardia caracasensis Weddell, Ann. 2.5-3.5 mm wide at base and 4-6 mm wide at orifice,<br />
Sci. Nat. 4(1): 70, pl. 10 (figs. 4-7). 1854. Type. sparsely (at base) to densely (distally) antrorse puberu-<br />
Venezuela. Distrito Federal: La Guayra, Oct 1843 lent-pubescent outside; with a white callous basal area<br />
(fl), Funck 463 (holotype, P).<br />
6-7.5 mm long, with a dense tuft of golden ascend-<br />
Pogonopus ottonis Klotzsch, Monastber. Koenigl. ing hairs (ca. 1 mm long), the medial and superior<br />
Preuss. Acad. Wiss. Berlin 1853: 500. 1853. Type. zones of throat glabrous; lobes 2.9-4.2 x 2.2-3.4 mm,<br />
Venezuela. Caracas, 1838, E. Otto 901 (B*; isotypes,<br />
narrowly ovate, antrorse yellow pubescent outside,<br />
BR?, K?, W?).<br />
with<br />
Howardia grandiflora Weddell, Ann. Sci. Nat. 4(1): 70. sparse short glandular hairs (0.2-0.4 mm long)<br />
1854. Type. Venezuela. Ocafia: Carmen, ca. 3000 inside. Stamens attached 6-7 mm from the base of<br />
m, Aug 1846-1852 (fl), Schlim 755 (holotype, P; the tube; filaments 31-37 mm long, subequal, with a<br />
isotypes, BR[2], G[2], K, frag-F).<br />
basal tuft of hairs; anthers elliptic, 2-2.6 x 0.8-1.1<br />
Carmenocania porphyrantha Werham, J. Bot. 50: 241, mm, base rounded, apex acute. Style 32-38 mm long,<br />
fig. 520. 1912. Type. Venezuela. Ocafia: Carmen, microscopically (40x) multicostate; style branches<br />
ca. 3000 m, Aug 1846-1852 (fl), Schlim 755 (holo- oblong, 1.25-1.5 x 0.5 mm, stigmatic surface microtype,<br />
K; isotypes, BR[2], G[2], P, frag-F). scopically (40x) papillose. Capsules ellipsoid-ovoid,<br />
apex round to shallowly hemispherical, 7-8 x 5-6<br />
Leafy branchlets sparsely to densely antrorse mm, sparsely golden-puberulent below the disk; disk<br />
short-pubescent; older branches sparsely short pubes- shallowly hemispherical, with a white tubercule at the<br />
cent. Stipules deltoid, acuminate, sparsely long-pi- point of attachment of the style, with some lenticels.<br />
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.<br />
x 5-10.5 cm, elliptic to obovate, cuneate to decurrent<br />
at base, acute to obtuse and acuminate at apex, acu- Distribution (Fig. 54) and ecology. Steep, dry<br />
men to 3 cm long; glabrous above, glabrous to mi- forested slopes, semi-deciduous and transitional fornutely<br />
puberulent below; primary and secondary veins ests, disturbed forest margins, and shady ravines, 300-
Systematic Treatment of the Genera Studied 131<br />
1200 m, in the coastal cordillera of Venezuela. Flow- Guarenas, 700 m, 22 Dec 1939 (fl-fr), Williams 12408 (A,<br />
ering specimens were collected during rainy season ECON, F, UC); Parque Nacional El Avila, southern slope,<br />
in January, August, September, October, and Decem- NNE of Caucagiiita, Qda. Tacamahaca, 10?30'N, 44?45'W,<br />
ber. Fruiting specimens were collected in<br />
850-1000 m, 3<br />
January,<br />
Aug 1977 (fl), Steyermark et al. 114008<br />
March, and November.<br />
(VEN). YARACUY: San Felipe, Rio Cocorotico, Camino del<br />
Isleflo, 18 Oct 1950 (fl-fr), Trujillo 1413 (MY).<br />
CULTIVATED. BRAZIL. Rio de Janeiro: Rio de<br />
Representative specimens examined. VENEZUELA.<br />
Janeiro, 3 Feb 1891 (fl), Glaziou 18305 (G[2], BR), 18309<br />
ARAGUA: Slopes of La Fila Maestra of Interior Coastal Cor-<br />
(P), and 18905 (P).<br />
dillera, NE of San Casimiro, 650 m, 26 Oct 1963 (fl),<br />
Steyermark 91807 (NY, P); Parque Nacional, below Rancho<br />
Grande, toward Maracay, 9 Dec 1943 (fl), Steyermark 54921<br />
Local names. Venezuela: coralito (Machado 11-<br />
(A, VEN); near Aragua-Miranda border, between Hda. 1984), naranjillo montafiero (Saer 642, Steyermark et<br />
Negro and Hda. Negrito, S of La Fila Maestra of Interior al. 114008).<br />
Coastal Cordillera, NE of San Casimiro, 650 m, 26 Oct 1963<br />
(fl), Steyermark 65741 (VEN); Parque Nacional Aragua, 600 This variety differs from Pogonopus speciosus var.<br />
m, 25 Aug 1938 (fl), Williams 10299 (F[2], G, MO, UC, US); sandwithianus in having corollas gradually expanded<br />
Carmen, 300 m, s.n. (fl), Williams 10415 (A, F[2], US); Mun.<br />
(Fig. 55F) at the orifice (vs. abruptly expanded, see<br />
Ocumare de la Costa, Parque Nacional Pittier, 800 m,<br />
Fig. 55B) with ascending pubescence, leaves puberu-<br />
Castillo 1966 (MO); Rancho Grande, 19 Sep 1975 (fl), lent<br />
Rodriguez 266 (MY); Maracay, Parque Ecol6gico, Fac.<br />
(vs. hirsute) below, and capsules ellipsoid-ovoid<br />
Agron6mica, 2 Jun 1957 (fl), Trujillo 3416 (MY). DISTRITO (vs. oblong-subglobose).<br />
FEDERAL: Between Caracas and La Guaira, Oct 1916 The<br />
(fl), complex nomenclatural history of this species<br />
Rose & Rose 21691 (GH, US); Valley of Tacagua, rd. is explained in the section dedicated to the taxonomic<br />
Caracas-La Guaira, near El Pauji, 1 Feb 1925 (fr), Pittier history of Pogonopus (see above). Schlim 755-P is the<br />
11672 (A, VEN); vic. of Caracas, 2 Nov 1916 (fr), Rose & holotype of Howardia grandiflora; Schlim 755-K is<br />
Rose s.n. (US[2]); vic. of Caracas, 1842, Lansberge 214 (A, the holotype of Carmenocania porphyrantha; and<br />
MO, NY, US); Caracas, La Guaira, 950 m, Nov 1938 (fl), Schlim 755-BR was annotated (unknown handwriting)<br />
Williams 10646 (UC); Quebrada Las Adjuntas, 10?32'N, as Pinckneya ionantha. Schlim 755 is the isotype of<br />
66?52'W, 11 Nov 1984 (fl-fr), Machado 11-1984 (VEN); rd.<br />
both Howardia grandiflora (Weddell, 1854), and<br />
to electricity plant behind Naiquetia, 12 Mar 1974 (fr), Gentry<br />
& Morillo 10339 (MO, NY); Parque Nacional El Carmenocaniaporphyranta (Wernham, 1912), which<br />
Avila,<br />
are both<br />
1200 m, 19 Jul 1978 (fl), Maroto & Salas 123 (MO); vic. of synonymous with Pogonopus speciosus var.<br />
Colonia Tovar, 1854-1855, Fendler 584 (MO, NY); near speciosus.<br />
Zigzag, between Caracas and Puerto Cabello, 18 Oct 1921 In the herbarium of the Museum of Natural His-<br />
(fl), Pittier 58 (F, G[2], GH, NY, US); Maracay (St. Ottilien), tory of Vienna (W), are preserved four specimens,<br />
450 m, Aug 1925 (fl), Zehnter 38 (M[2]); El Valle, Hda. once part of the Jacquin herbarium, all labeled as<br />
Sosa, 11 Sep 1949 (fl), Trujillo & Ferncndez 293 (MY); "Macrocnemum/Calycophyllum speciosum Jacq.<br />
Colonia Tovar, 1852 (fl), Moritz 840 (B*, F, G, P). GUARICO:<br />
Hwy. to Caucagua, 10 km NWN of Altagracia de Orituco,<br />
440 m, 18 Nov 1973 (fl-fr), Davidse 4174 (MO, VEN); Dto.<br />
Libertador, Rio Chichiriviche, 1-2 km S of Chichiriviche,<br />
20-50 m, 10?32'N, 67?14'W, 9 Oct 1976 (fl), Steyermark &<br />
Carrerio-Espinoza 112706 (F, MO, NY, VEN). LARA: Dto.<br />
Jim6nez, Paso de Angostura, represa de Yacambu, confluence<br />
of Qda. Honda y Rio Yacambu, 09?4 1'N, 69?3 'W, 35<br />
km E of Sanare, 500 m, 27-28 Dec 1973 (fr), Steyermark &<br />
Carreno-Espinoza 108790 (VEN); El Altar, near La Miel, 4<br />
Jan 1967 (fr), Smith VI-280 (VEN); Serrania de Terepaima,<br />
S of Barquisimento, 800-1000 m, Aug 1930 (fl), Scer<br />
d'Heguert 642 (M, VEN). MIRANDA: Rio Guarita, Morro de<br />
La Gaurita, 10?27'N, 66?48'W, 900-950 m, 17 Aug 1975,<br />
Steyermark & Berry 112141 (VEN); Morro de La Guarita,<br />
880 m, 11 Jan 1976 (fr), Berry 1847 (VEN); near El Encantado,<br />
7 Oct 1917 (fr), Pittier 7468(VEN); Qda. Las Comadres,<br />
near Las Mostazas, 1100 m, Nov 1924 (fl),Allart 236 (VEN);<br />
Parque Nacional Guatopo, 16 Jan 1967 (fr), L6pez-Palacios<br />
1939 (VEN), 17 Jan 1967 (fr), 1940 (NY); vic. of El Petare,<br />
11 Sep 1921 (fl),Pittier9799(F, GH[2], K, NY, US, VEN);<br />
-<br />
Caracas - Bredemeyer." Bredemeyer collected in<br />
Venezuela in 1787-1788 (Lanjouw & Stafleu, 1954),<br />
and part of his living collections were transported to<br />
the greenhouses of the Schonbrunn gardens. In turn,<br />
Jacquin described Macrocnemum speciosus from living<br />
material cultivated in the Schonbrunn greenhouses.<br />
Of these four specimens, one bears the annotation<br />
(in Schumann's handwriting): "Pogonopus<br />
speciosus - det. Schumann in Fl. Bras.," and I select<br />
this specimen as lectotype ofM. speciosus. The three<br />
other specimens are treated as isolectotypes.<br />
Specimens of Moritz 840 (B*, F, P; photo-B at G,<br />
GH, MO, NY, VEN; photo-P at F, MO, VEN), collected<br />
in Colonia Tovar (Venezuela) were annotated<br />
by Klotzsch as "Pogonopus moritzii Klotzsch" and by<br />
Schumann as "Pogonopus speciosus var. ottonis K.<br />
Schumann." Because both names remained unpublished,<br />
Moritz 840 should not be considered type<br />
material.
132 Flora Neotropica<br />
2b. Pogonopus speciosus (Jacquin) K. Schumann var.<br />
sandwithianus Steyermark. Pogonopus speciosus<br />
(Jacquin) K. Schumann subsp. speciosus var. sandwithianus<br />
(Steyermark) Steyermark in Lasser &<br />
Steyermark, Flora de Venezuela 9(1): 240. 1974.<br />
Pogonopus speciosus (Jacquin) K. Schumann<br />
subsp. sandwithianus Steyermark, Acta Bot. Venez.<br />
6: 110. 1971. Type. Colombia. Magdalena: Santa<br />
Marta, near Bonda, 50 m, 20 Aug 1898 (fl), H. H.<br />
Smith 100 (holotype, NY; isotypes, A, BR, CM,<br />
K, LL, MO, GH, COL, F, US, VEN).<br />
Figs. 3K,L, 4G, 8D, 16D, 54, 55A-E, 56A-F<br />
Pinckneya ionantha Planchon & Linden, Bot. Zeit. 12:<br />
365. 1854. Type: Colombia. Magdalena: Valley of<br />
Rio Magdalena, Momp6s, 1847, Schlim s.n. (lectotype,<br />
BR, here selected).<br />
Leafy branchlets hirsutulous to hirsute, with<br />
golden-yellow hairs; older branches sparsely hirsutulous.<br />
Stipules shallowly triangular, strongly acuminate;<br />
hirsutulous to hirtellous; 3.5-6.5 x 3.5-5 mm.<br />
Leaves 10-29 x 5.5-10 cm, elliptic to obovate, cuneate<br />
to decurrent at base, acute to obtuse and acuminate<br />
at apex, acumen to 3 cm long, sparsely puberulent<br />
(at least on midrib) above, sparsely to densely<br />
golden-hirtellous below; primary and secondary veins<br />
sparsely to densely hirsutulous to hirsute, secondary<br />
veins 9-16 each side; petioles 4-36 mm long, 1-2.5<br />
mm thick, densely golden-hirsutulous, giving a velvety<br />
appearance; domatia absent. Inflorescences<br />
subpyramidal, laxly paniculate; 14-19(-22) cm long,<br />
basal branches 4-8 cm long, lateral branches 2-3<br />
pairs, basal portion of axis not branched 6-8(-13) cm<br />
long; rachis and branches densely golden-pubescent<br />
(to hirtellous), becoming sparsely pubescent in older<br />
parts, branchlets of terminal cymes densely goldenhirtellous;<br />
distal branches congested cymules of 9-<br />
20 flowers; distal bracts leaf-like, to 12.5 x 5.5 cm,<br />
decreasing in size distally, down to 3 x 1.5 cm;<br />
bracteoles subtending flowers 1.5-1.5 x ca. 0.5 mm,<br />
linear, densely golden-pubescent. Flower pedicels 2-<br />
8 mm long, densely hirsutulous to pilose; hypanthium<br />
densely hirsutulous to hirtellous; young flower buds<br />
golden-hirsutulous throughout (hairs to 0.7-1.1 mm<br />
long). Calyx 0.3-0.7 x 3.5-4 mm, densely yellow<br />
pubescent; lobes usually linear, 1.5-3 x 0.5-1 mm;<br />
in some flowers 1-5 calyx lobes expanded into<br />
calycophylls. Calycophylls 4.5-5(-9) x 2.5-4.5(-5.5)<br />
cm, blades ovate to widely ovate, base obtuse to<br />
rounded (rarely subcordate), apex acute (with the very<br />
tip rounded), palmately 3-5-veined, glabrous to glabrescent;<br />
primary and secondary veins sparsely puberulent,<br />
pink, dark pink, to pink-red; stalks 1-2 cm<br />
long. Corolla tubular, abruptly ampliate at orifice, 26-<br />
34 mm long, rose-pink, lilac to pink-red; tube subcylindrical<br />
with abrupt expansion, 23-25 mm long, 3-4<br />
mm wide at base and 5-7 mm wide at orifice, densely<br />
erect golden-hirsutulous outside; with a white callous<br />
basal area 5-7 mm long, with a dense tuft of whitehispid<br />
hairs (ca. 1 mm long), medial zone of throat<br />
glabrous, superior zone sparsely long-pubescent; lobes<br />
4-6.5 x 3.8-5 mm, widely ovate, golden hirtellous-<br />
pubescent outside, with sparse elongate glandular<br />
hairs (0.5-0.7 mm long) inside. Stamens attached ca.<br />
7 mm from the base of the tube; filaments 27-34 mm<br />
long; anthers narrowly elliptic, 2.6-3 x 0.8-1 mm,<br />
base narrowly elliptic, apex round, short-ligulate.<br />
Style 32-40 mm long, microscopically (40x) papil-<br />
lose; style branches oblate, ca. 1.5 x ca. 0.75 mm,<br />
stigmatic surface with microscopic (40x) glandular<br />
hairs. Capsules oblong to subglobose, apex sub-<br />
truncate to shallowly hemispherical, 6-7 x 5-6 mm,<br />
short-pubescent below the disk; disk an elevated ring<br />
(do-nut shaped), concave, with a brown tubercule at<br />
the point of attachment of the style. Seeds 0.44-0.64 x<br />
0.35-0.42 mm.<br />
Local names and uses. Colombia: quina (Kernan<br />
167), quina morada (Delprete & Apreza 6359).<br />
Distribution (Fig. 54) and ecology. Seasonal de-<br />
ciduous forested slopes, often in limestone soil, 50-<br />
1200 m, endemic to Sierra Nevada de Santa Marta<br />
(Magdalena Department, Colombia) and Serrania de<br />
Perija (Cesar Department, Colombia). Flowering<br />
specimens were collected in July, August, September,<br />
November, and December. Fruiting specimens were<br />
collected in February, August, and December.<br />
Reproductive biology. The flowers of this vari-<br />
ety are visited by both butterflies and bees of uniden-<br />
tified species. Some flower buds are initially robbed<br />
by wasps, which make lateral incisions at the height<br />
of the nectaries, and are secondarily robbed by ants<br />
that enter through these incisions (Fig. 56E).<br />
Specimens examined. COLOMBIA. CESAR: Mun.<br />
Valledupar, Pueblo Bello, 1200 m, 13 Jul 1983 (fl), Cuadros<br />
1677 (COL, US); Serrania de Perija, 8 km NE of [Augustin]<br />
Codazzi, 275 m, 10 Feb 1945 (fr), M. L. Grant 10891 (NY);<br />
near Codazzi, 150 m, 30 Sep 1943 (fl), Haught 3705 (F, NY,<br />
US); Cerro Chimichagua, 35 km NE of El Banco, 11 Aug<br />
1938 (fl), Haught 2236 (COL, F, NY, US). MAGDALENA:<br />
Parque Nacional Tayrona, 50-150 m, I 11 8'N, 74005'W, 50-<br />
150 m, 2 Nov 1972 (fl), Kirkbride 2637 (COL, MO, NY,<br />
US, VEN); Parque Nacional Tayrona, quebrada Cinto, 50<br />
m, Sep 1979 (fl), Moreno 193 (COL); Parque Nacional
Systematic Treatment of the Genera Studied 133<br />
Tayrona, 74?02'W, 11?20'N, 23 Aug 1986 (fr), Gentry &<br />
Cuadros 55525 (MO); Sierra Nevada de Santa Marta, rd.<br />
from San Pablo to San Pedro de la Sierra, 2 km before El<br />
Mico, 10?53'N, 74?09'W, 300 m, 29 Dec 1992 (fl), Delprete<br />
& Apreza 6359 (AAU, COL, F, GB, MO, NY, TEX[3]);<br />
Bonda, near Santa Marta, 26 Aug 1944 (fr), Romero-<br />
Castaneda 132 (COL, F[2]); Mun. Fundaci6n, N of Santa<br />
Rosa, 400-800 m, 6 Aug 1971 (fl), Romero-Castaneda<br />
11198 (COL, MO, NY); Sierra Nevada de Santa Marta, Rio<br />
Frio, Jul 1925 (fl), Walker 1277 (F, GH, MO, US); trail<br />
Cenizo-Santa Rosa, Rio Tucurinca, 31 Jul 1944 (fl), 300 m,<br />
Kernan 167 (US[2]); Magdalena Valley, La Jagua, 8 Sep<br />
1924 (fl), Allen 622 (F, MO); Linden s.n. (BR).<br />
This variety is readily distinguishable from Pogonopus<br />
speciosus var. speciosus in having corollas<br />
abruptly ampliate (Fig. 55B) at the orifice (vs. gradually<br />
ampliate, Fig. 55F) and golden hirsutulous (Fig.<br />
55C) outside (vs. appressed-pubescent, Fig. 55G), leaf<br />
blades sparsely to densely golden-hirtellous below,<br />
secondary veins and capsules golden-hirsute, and capsular<br />
disk with an outer elevated ring [doughnut<br />
shaped] (vs. hemispherical). Also, this variety has<br />
ard in A. P. de Candolle) A. P. de Candolle, Prodr.<br />
4: 367. Sep 1830. Macrocnemum tubulosum A.<br />
Richard in A. P. de Candolle, Prodr. 4: 403. Sep<br />
1830. Macrocnemum tubulosum A. Richard, nom.<br />
superfl., Mem. Fam. Rubiacees. Dec 1830 (reimp.<br />
Mem. Soc. Hist. Nat. Paris. ser. 3, 5: 279. 1834).<br />
Howardia richardi Weddell, nom. superfl., Ann.<br />
Sci. Nat. 4(1): 69, pl. 10 (figs. 8-10). 1854. Type.<br />
Brazil. Locality, date, and collector unknown (holotype,<br />
P-Richard Herbarium; photo-P at F).<br />
Figs. 8E, 16D, 54, 57A-E<br />
Pogonopus febrifugus (Weddell) J. D. Hooker in<br />
Bentham, G. & J. D. Hooker, Gen. P1. 2: 47. 1873.<br />
Howardiafebrifuga Weddell, Ann. Sci. Nat. 4(1):<br />
67, pl. 10 (figs. 1-3). 1854. Chrysoxylonfebrifugum<br />
Weddell, nom. inval., Monogr. Cinch. 100. 1849.<br />
Type. Bolivia. Enquisivi, Dec 1846 (fl), Weddell<br />
4190 (lectotype, P, here selected; isolectotype, US).<br />
Pogonopus febrifugus (Wedd.) J. D. Hooker var.<br />
macrosema Hutchinson, Kew Bull. 1910: 200. 1910.<br />
Type. Bolivia. "Forest of eastern Bolivia," 6 Mar<br />
1910 (fl), Gosling s.n. (lectotype, K, here selected).<br />
been found only in the Sierra Nevada de Santa Marta<br />
and the Serrania de Perija (Colombia, Figs. 54, 56A),<br />
whereas var. speciosus is endemic to the Venezuelan<br />
Caribbean coast (Fig. 54).<br />
Sandwith (1949) helped to clarify the complex<br />
taxonomic history of Pogonopus speciosus (see<br />
above) and pointed out the differences between the<br />
specimens collected in Venezuela (Schlim 755, Funk<br />
463) and those collected in Sierra Nevada de Santa<br />
Marta (Smith 100, Dawe 698). Steyermark, following<br />
Sandwith's (1949) observations, described this<br />
taxon first as a subspecies (Steyermark, 1971) of P.<br />
speciosus and later as a variety (Steyermark, 1974)<br />
of P. speciosus subsp. speciosus, dedicating its epithet<br />
to its first careful observer, Sandwith himself.<br />
The type of Pinckneya ionantha was cited in the<br />
original description (Linden & Planchon, 1854) as:<br />
"von Schlim [s.n.] in Neu-Granada [Colombia], an<br />
den brenden Ufern des Magalenenflusse [valley of<br />
Magdalena River] in der Prov. Mompox [Mompos]<br />
gesammelt." In the herbarium of BR there is a specimen<br />
simply labeled "Pinckneia ionantha [orth. var.,<br />
handwriting unknown], Neu-Granada, Schlim." This<br />
specimen is here selected as the lectotype of this taxon.<br />
Shrubs 2.5-8 m tall, exceptionally singlestemmed<br />
small trees to 10 m tall, to 30 cm dbh, muchbranched<br />
shrubs to treelets, with central branches or<br />
central stem erect, outer branches usually scandent;<br />
bark smooth, brownish gray. Leafy branchlets<br />
antrorse minute-puberulent; older branches sparsely<br />
puberulent to glabrate; lenticels cream-white, punctiform<br />
to linear, to 5-6 mm long. Stipules very shallowly<br />
triangular, not acuminate to short acuminate,<br />
glabrescent to puberulent outside, with elongated<br />
colleters (to 0.5 mm long) inside, 3-4 x 4.5-6 mm,<br />
persistent; older stipules often breaking into two<br />
halves. Leaves (8-)13.5-25<br />
3. Pogonopus tubulosus (A. Richard in A. P. de<br />
Candolle) K. Schumann in Martius, Fl. Bras. 6(6):<br />
265. 1889. Chrysoxylon tubulosum (A. Richard in<br />
A. P. de Candolle) O. Kuntze, Revis. Gen. P1. 1:<br />
278. 1891. Calycophyllum tubulosum (A. Rich-<br />
x 5-12.5 cm, narrowly<br />
to widely elliptic to elliptic-obovate, cuneate to longdecurrent<br />
at base, acute to obtuse and short-acuminate<br />
at apex, acumen to 3 cm long; dark green above, pale<br />
green below, stiff-foliaceous; drying olive-green to<br />
yellow-green, stiff-chartaceous; glabrous above, glabrous<br />
to puberulent to white-pilose below; primary<br />
and secondary veins glabrous to puberulent to sparsely<br />
pilose, secondary veins (5-)7-12 each side, tertiary<br />
veins subparallel, quaternary veinlets reticulate; petioles<br />
4-24 mm long, 1-2.5 mm thick, adaxially concave<br />
to flattened, glabrous to minute puberulent;<br />
domatia absent or tuft-domatia of sparse pubescent<br />
hairs. Inflorescences short-pyramidal, paniculate with<br />
opposite decussate branches, each lateral branch subtended<br />
by leaf-like bracts, terminating in a lax cymule;<br />
(7.5-)14-29 cm long, basal branches (4-)6-15 cm<br />
long, lateral branches to 4 pairs, basal portion of axis<br />
not branched 3.5-8.5 cm long; rachis terete, with puncti-
134 Flora Neotropica<br />
..~~~~~~~~~~~~.<br />
OOF ~ ~ ~<br />
-Ac<br />
I~~~~~~J<br />
FIG. 5'7. Pogonopus tubulosus (A-E from Alfaro 35 9, M O; F firom Lourteig 714, F). A. Habit of inflorescence. B. Open<br />
glandular pubescence of corolla lobe. F. Mature capsule.
Systematic Treatment of the Genera Studied 135<br />
form to linear lenticels (to 3 mm long), rachis and<br />
branches glabrous to puberulent; flowers on distal<br />
branches alternate or in lax cymules of 4-8 flowers;<br />
distal bracts leaf-like, petiolate, elliptic-obovate, to 16 x<br />
7 mm, decreasing in size distally down to 3-9 x 1-3 mm;<br />
bracteoles subtending flowers 0.5-1.2 x ca. 0.5 mm,<br />
narrowly triangular. Flowers pedicellate, pedicels 6-<br />
20 mm long, minute puberulent; hypanthium<br />
obconical to narrowly oblong to obovoid, 3-4.5 x 2-<br />
3.5 mm, glabrous to minute puberulent; young flower<br />
buds minutely puberulent at apex. Calyx tube ex-<br />
tremely reduced, 0.3-0.6(-1) x 2.8-5.2 mm, glabrous<br />
to minute puberulent; lobes 5, deltoid to acuminate,<br />
0.5-1.5 x 0.3-0.5 mm; in some flowers 1-5 lobes<br />
expanding into foliose expansions of different sizes,<br />
from pale green to pink elongate lobes, to dark-pink<br />
calycophylls. Calycophylls (3.2-)4.5-15.5 x (2-)4-<br />
13 cm, blades elliptic-acuminate to ovate to bilobate<br />
with or without a small mucron in the middle, base<br />
obtuse to cordate (rarely acute), apex round to obtuse<br />
(rarely acute), palmately veined, blade glabrous above<br />
and below, primary veins and/or blade margins sparsely<br />
puberulent, bright pink, pink-red, wine-red to crimson;<br />
stalks 0.7-2.5 cm long. Corolla narrowly tubular,<br />
with a moderate constriction near the base, 30-50 mm<br />
Distribution (Fig. 54) and ecology. Fairly com-<br />
mon in primary and secondary forests and their rem-<br />
nants, in seasonal-dry forests (500-1400 m), and up<br />
to cloud forests (1600-2800 m), sometimes in clay-<br />
soils, of Peru, Bolivia, western Brazil, and northern<br />
Argentina. Flowering specimens were collected in<br />
January, February, March (most collections), April,<br />
May, June, July, and December. Fruiting specimens<br />
were collected in February, March, May, June, July,<br />
and November.<br />
Representative specimens examined. PERU. Cuzco:<br />
Prov. Calca, Chanchamayo, Lares Valley, 12 Mar 1929 (flfr),<br />
Weberbauer 7940 (F, GH); Prov. La Convenci6n, Sambray<br />
valley, W affluent of Vilcanota, 1000-1500 m, 10 Mar<br />
1936 (fr), Mexia 8035 (F, G, GH, K, MO, US). HUANCAVE-<br />
LICA: Vuelo-Pata, above Virgen-Pampa, SE of Tintay, 1300<br />
m, 13 Apr 1964 (fl), Tovar 4629 (US[2]). JUNIN: E of La<br />
Merced, 4 Nov 1962 (fr), Schunke 6214 (F); rd. La Merced-<br />
Villa Rica, 75?17'W, 10?54'S, 6 Jan 1984 (fl), Smith et al.<br />
5605 (F, MO, NY); La Merced, May-Jun 1929 (fl), Killip<br />
& Smith 23811 (F, NY); Prov. Tarma, San Ram6n, 800 m,<br />
16 Mar 1976 (fl), Plowman & Kennedy 5649 (F, GH, K).<br />
BRAZIL. ACRE: Rio Acre, Str. Seringal Sao Francisco,<br />
Mar 1911 (fl), Ule 9848 (B, G, US); Cobija, Jan 1912, Ule<br />
9848b (K, MG, UC); Cruzeiro do Sul, sub-base do Projecto<br />
RADAM, 15 Mar 1976 (fi), Ramos & Mota 368 (INPA,<br />
MG). MATO GROSSO: Jauru, km 16 W on rd. MT-248, 7 May<br />
1995 (fl), Hatschbach et al. 62451 (US). MATO GROSSO DO<br />
long, membranaceous (not fleshy) when fresh, pale SUL: Mun. Aquidauana, Piraputanga, 18 Feb 1970 (fl),<br />
purple, lilac to dark red outside, cream-white inside; Hatschbach 23784 (MO, NY, US); Mato Grosso, Dec 1827<br />
tube subcylindrical to narrowly obconical, abruptly (fl), Riedel 1229 (BR); Campos dos Urupas, Feb 1919 (fl),<br />
ampliate at orifice, 28-46 mm long, 2.5-6 mm wide Kuhlmann 2353 (R, RB). ROND6NIA: Vilhena, 12?58'S,<br />
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<br />
puberulent outside; with a white callous basal area 10- Preto do Oeste, Reserva Ecol6gica do CNPq-INPA, 10? 1 'S,<br />
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,<br />
the medial and superior zones of throat glabrous; lobes MG, NY, US); Ouro Preto do Oeste, rd. Cuiaba-Porto Velho,<br />
km<br />
5, narrowly spreading, 3-4.3 x 382, 10?11'S, 62?63'W, 1 Jul 1984<br />
2.3-4 mm, narrowly to<br />
(fr), Cid et al. 4939<br />
(INPA, MG, NY, US); Mun. Jam, rd. Cuiaba-Porto Velho,<br />
widely ovate, glabrous to sparsely puberulent outside, km 428, 3 Jul 1984 (fl-fr), Cid et al. 5022 (K, MG, NY, US);<br />
microscopically (40x) glandular (0.02-0.15 mm long) Mun. Jiparana, Gleba km 20, 9 Apr 1983 (fl), Paulino Filho<br />
inside. Stamens exserted, slightly unequal, attached & Silva 83-96 (INPA), Silva 6125 (INPA, MG).<br />
10-15 mm from the base of the tube (ust above cal- BOLIVIA. LA PAZ: Tipuani-Guanay, Dec 1892 (fl),<br />
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]),<br />
white, with a basal tuft of sericeous hairs; anthers nar- Apr-Jun 1892, 1344 (A, F, G, GH, K, M, MO, NY[3],<br />
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,<br />
dial zone, base rounded, apex acute, purple-blue when 67?43'W, 1200 m, 25 Mar 1982 (fl-fr), Solomon 7321 (MO,<br />
fresh, drying black. Pollen exine reticulate. Style ex- NY[2]); 41.5 km N of Loyosa, 16?00'S, 67036'W, 1000 m,<br />
26 Mar 1982, Solomon 7359<br />
serted, 35-54 mm long, glabrous, white; style branches<br />
(INPA); Prov. Nor-Yungas,<br />
Milluguaya, 900 m, Dec 1917 (fl), Buchtien 761 (F, GH, MO,<br />
membranaceous, linear-oblong, 1.5-2.5 x 0.5-0.7 mm,<br />
NY, US); Prov. Sur-Yungas, basin of Rio Bopi, San<br />
reversed, stigmatic surface microscopically (40x) papil- Bartolom6, 750-900 m, 1-22 Jul 1939 (fr), Krukoff 10054,<br />
lose. Capsules subglobose to ellipsoid, base rounded, 10167 (A, F, G[2], K, MO, NY, UC); Coroico, 1000 m, 16<br />
apex subtruncate, 7-10 x 5-9 mm, with circular white Mar 1928 (fl), Troll 1514 (B). SANTA CRUZ: Lagunillas, 900<br />
lenticels, dark brown, glabrous to puberulent below m, May 1949 (fl), Cdrdenas 4261 (GH, US); Las Tejitas,<br />
the disk; disk sometimes exceeding the calyx, gla- 420 m, 2 Mar 1945 (fl), Peredo 525 (A, NY); Prov. Florida,<br />
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,<br />
63?33'W, 16 Feb 1988 (fl-fr), Nee & Saldias 36320 (NY);<br />
Prov. Cordillera, Alto Parapeti, 800 m, 25 Jan 1980 (fl), de<br />
Michel 41 (UB); Prov. Nuflo de Chavez, San Ram6n, 21 Feb
136 Flora Neotropica<br />
1991 (fr), Quevedo 316 (NY); Prov. Nuflo de Chavez, study Weddell (1849) also named this species Chrysoarea<br />
of BOLFOR project, Las Trancas-95, 1631 'S, 61 ?50'W, xylonfebrifugum nom. inval., using material he col-<br />
450 m, 20 Mar 1995 (fl), Mamani 436 (NY); Prov. Chiquitos, lected in Bolivia, and later (Weddell, 1854) validated<br />
Serrania de Sunsas, Tucavaca, 4 Jul 1995 (fr), Jardim &<br />
it by transferring it to Howardia. In the Paris her-<br />
Cadden 2153 (NY). TARIJA: Prov. O'Connor, San Sim6n,<br />
barium there is a<br />
20 km E of Entre Rios, 1300 m, 7 Feb 1937 (fl), West 8261<br />
specimen (Weddell 4190), collected<br />
(GH, MO, UC); rd. Tarija-Villa Montes, San Sim6n, 22<br />
in Bolivia<br />
May<br />
(Prov. Enquisivi), that bears on the attached<br />
1971 (fr), Krapovickas et al. 19143 (F, GH); Prov. Gran label (in Weddell's handwriting) the names "Chryso-<br />
Chaco, Sierra de Aguarague, 1200 m, 8 Jun 1977 (fl), xylonfebrifugum<br />
Krapovickas & Schinini 31024 (F[2], MO); Prov. Gran<br />
Chaco, Chaco, rd. Villa Montes-Jomiba, 500 m, Feb 1980<br />
(st), Rojas 1546 (UB); Villamontes, 1923, Pflanz 2053 (B).<br />
ARGENTINA. JUJUY: Depto. Ledesma, El Sauzal, 5 Jul<br />
1911 (fr), Lillo 10785 (F, GH, NY[2], UC, US); Depto.<br />
Ledesma, El Quemado, 4 Jun 1973 (fr), Turpe et al. 4690 (K,<br />
NY[2], US); Calilegua, 2 Jun 1943 (fr), Bartlett 20361 (GH,<br />
NY[2]); Depto. Ledesma, Parque Nacional Calilegua, 28 Jun<br />
1989 (fr), Kiesling 7109 (NY). SALTA: Urundel, 400 m, 18<br />
Jun 1941 (fr), Zabala 165 (F, GH, US); Depto. Orin, Campamento<br />
Rio Pescado, 16 Jan 1951 (fl), Scolnik 1678 (F[2]);<br />
Depto. Oran, Vespucio, 13 Jul 1937 (fr), Cabrera 4159 (F,<br />
NY[2]); Depto. Oran, Rio Pescado, 400 m, 1 Jun 1951 (fr),<br />
Sleumers.n. (B). TUCUMAN: Depto. Capital, Instituto Miguel<br />
Lillo, 450 m, 1 Apr 1974 (fl-fr), Borsini s.n. (CAS).<br />
Local names and uses. Peru: bouganvillea (Cuzco,<br />
Mexia 8035), flor de hoja (Cuzco, Mexia 8035). Bolivia:<br />
quina (Cardenas 2593; Peredo 525; Fosberg 28653; Nee<br />
& Coimbra 35199), sacha quina (Krapovickas et. al.<br />
19143). Argentina: cascarilla (Schultz 978), palo marfil<br />
(Lillo 10785), sacha quina (Borsini s.n.; Meyer 4811).<br />
Pogonopus tubulosus has been used by native<br />
people and European settlers as a secondary source<br />
of quinine to cure malaria and intermittent fevers, and<br />
reported by many as: "Bark used for quinine" (Bolivia,<br />
Gosling s.n., 3 Jun 1910), "Corteza febrifuga"<br />
(Argentina, Schultz 978), and "used as Cinchona bark"<br />
(Bolivia, Cdrdenas 2593). This species is also widely<br />
cultivated as an ornamental shrub, because of its bright<br />
red calycophylls: "Muy decorative por las grandes<br />
bracteas rosado-vivo" (Schultz 978).<br />
- Howardiafebrifuga Wedd," which<br />
is here selected as the lectotype of Howardiafebrifuga.<br />
Another specimen of Weddell 4190 is present<br />
at US, which is here treated as isolectotype. Another<br />
specimen (ex Paris Herbarium) collected by Weddell<br />
in Prov. Enquisivi is preserved at US, and this is a<br />
probably an isolectotype (its label lacks Weddell's<br />
collection number). Two other Weddell collections (ex<br />
Paris Herbarium) from "Prov. de la Cordillera, Bolivia"<br />
are at BR and K, and one (ex Paris Herbarium)<br />
from "Prov. de Santa-Cruz de la Sierra, Bolivia" is at<br />
F. These last three specimens were cited among the<br />
syntypes by Weddell (1849, 1854) and therefore are<br />
paratypes of H. febrifuga.<br />
Hooker (1873) transferred Howardiafebrifuga to<br />
Pogonopus. Hutchinson (1910) published P.febrifugus<br />
var. macrosema using specimens collected by Cecil<br />
Gosling in eastern Bolivia. One of the latter (Gosling<br />
s.n., 6 Mar 1910) was located at K, on which has been<br />
penciled (unknown handwriting): "P. febrifugus var.<br />
macrosema." This specimen is here selected as the lectotype<br />
of this variety. The same specimen also bears<br />
a label that has the annotation (in Hutchinson's handwriting):<br />
"Pogonopus amarus, Hutchinson sp. nov.,"<br />
but this name was never published.<br />
Some herbarium specimens (Rusby 2090, G, P)<br />
were annotated by Britton as "Pogonopus bolivianus<br />
Britton, sp. nov.," but this specific name has never<br />
been published. Another specimen (Bridges s.n., BR-<br />
Martius Herbarium) was annotated by Schumann as<br />
Pogonopus tubulosus var. boliviensis, a varietal name<br />
that also remained unpublished.<br />
In the Richard Herbarium (P) a specimen was<br />
found with a label annotated (in A. Richard's handwriting):<br />
"Macrocnemum tubulosum A. Rich. - Bresil,"<br />
which I consider to be the holotype of this species.<br />
Weddell (1854) renamed this species as Howardia<br />
richardi, a superfluous name based on the type of M.<br />
tubulosum. The holotype of M tubulosum annotated<br />
by A. Richard has two additional labels: one with the<br />
annotation (in Weddell's handwriting) "Howardia<br />
Richardi Wedd.," the other with morphological pencil-drawings<br />
and lightly penciled the name "Calycophyllum<br />
tubulosum." Because of the additional labels,<br />
this specimen preserved in the Richard herbarium<br />
is here selected also as the holotype for both names.<br />
Rejected Species<br />
Pogonopus erythroxylon (Standley) J. H. Kirkbride,<br />
Acta Amaz. 15: 54. 1985. Capirona erythroxylon<br />
Standley, Publ. Field Mus. Nat. Hist., Bot. Ser.<br />
22: 48. 1940, non Sickingia erythroxylon<br />
Willdenow, Gesellsch. Naturf. Freunde Neue Schr.<br />
3: 446. 1801. Non Simira erythroxylon<br />
(Willdenow) Bremekamp, Acta Bot. Neerl. 3:153.<br />
1954. Type. Ecuador. Surroundings of Guayaquil,<br />
"Vulgo Palo Colorado, F.H.D. 139.L495, Afio de<br />
1800," Tafalla s.n. (holotype, [F.H.D. 139.L495]<br />
MA; isotype, MA; photo-MA at F, TEX) = Simira<br />
ecuadorensis (Standley) Steyermark.
Systematic Treatment of the Genera Studied 137<br />
This species was originally described<br />
&<br />
by Standley<br />
Hooker, Gen. PI. 2: 45. 1873; Baillon, Hist. PI.<br />
(1940) as Capirona erythroxylon, citing two<br />
7: 397, 493.<br />
speci-<br />
1880; Poiret in Lamark, Encycl. Suppl.<br />
mens collected by Ruiz and Pavon in 1800 and 9:<br />
pre-<br />
230; Schumann in Martius, Fl. bras. 6(6): 259served<br />
in the Herbarium Peruvianum (MA). Both 260. 1889; Schumann in Engler & Prantl, Nat.<br />
"putative collectors" returned from South America to Pflanzenfam. 4(4): 18, fig. 6A-E. 1891; Urban,<br />
Madrid in 1788, leaving their collecting efforts under Symb. Ant. 410-411. 1900; Standley, N. Amer.<br />
the responsibility of Juan Jose Tafalla. Between 1799 Fl. 32: 5-6. 1918; Rizzini, Rev. Bras. Biol. 7(2):<br />
and 1803, Tafalla lead a collecting expedition in the 275-277. 1947; Bremekamp, Rec. Trav. Bot.<br />
forests Neerl. 31: 260-261.<br />
(now mostly destroyed) surrounding<br />
1934; Standley, Publ. Field<br />
Guayaquil, southern Ecuador (Estrella, 1991). It is Columbian Mus., Bot. Ser. 7: 203,370-371. 1931;<br />
most likely that the type specimens of C. erythroxylon Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 192.<br />
were collected in this area. Tafalla periodically sent 1936; Steyermark, Ceiba 3: 18-19. 1952;<br />
his own collections to Pavon, who filed them in the Steyermark, Mem. New York Bot. Gard. 12(3):<br />
Herbarium Peruvianum (MA). The type specimens of 178-186. 1965; Steyermark, Acta Bot. Venez. 8:<br />
C. erythroxylon should, therefore, be cited as collected 248-249. 1973; Steyermark in Lasser &<br />
by Tafalla in the vicinity of Guayaquil (Ecuador), and Steyermark, Fl. Venez. 9(3): 244-253. 1974;<br />
not collected by Ruiz and Pav6n in Peru (as reported<br />
Howard, Fl. Lesser Antilles 397. 1989; Dwyer,<br />
Ann. Missouri Bot. Gard. 67:<br />
by Kirkbride,<br />
82-84,<br />
1985a).<br />
fig. 18. 1980;<br />
The type specimens were annotated by Krause as Burger & Taylor in Burger, Fl. Costaric.,<br />
Capirona and later described as C. erythroxylon by<br />
Fieldiana, Bot. n.s. 33: 94-95, fig. 37. 1993;<br />
Standley (1940). Kirkbride (1985a) transferred this Delprete, Brittonia 48: 35-44, fig 1. 1996; Delprete<br />
species to Pogonopus because of its "valvate aestiva- in Harling & Andersson, Fl. Ecuador (in press).<br />
tion and many small, ovules in each locule ... with 1999. Type species. Chimarrhis cymosa Jacquin.<br />
one calyx lobe sometimes expanded into a large col- Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio Janeiro<br />
ored foliar organ." I studied a detailed set of photos 4:177, pl. 23. 1925. Type species. Pseudochimarrhis<br />
of the two Madrid types taken by Kirkbride and con- turbinata (A. P. de Candolle) Ducke [= Chimarrhis<br />
cluded that C. erythroxylon matches the types of subgen. Pseudochimarrhis (Ducke) Delprete<br />
Sickingia ecuadorensis [Mille 884 (F)] (= Simira turbinata A. P. de Candolle].<br />
ecuadorensis), which was collected in the forests of<br />
Guayaquil, and with which it is here reduced to syn- Trees, often tall canopy trees, bole irregular, with<br />
onymy. The flowers of the types have imbricate aes- or without buttresses; buttresses sometimes very large,<br />
tivation, but one photo of the set taken by Kirkbride especially in the Amazonian species; bark smooth to<br />
shows a few loose flowers much bigger than those on deeply fissured; wood yellow, very hard. Stipules<br />
the specimens, and with valvate aestivation. These interpetiolar, above leaf attachment, free or connate<br />
dubious flowers, certainly fragments of a mixed col- at base, adnate to the petioles, persistent or caducous,<br />
lection, were the ones that probably directed Kirkbride leaving a scar encircling the stem above the node.<br />
(1985a) to transfer this species to Pogonopus. The Leaves elliptic to ovate to obovate, usually acuminate<br />
common name "palo colorado" (= red wood), anno- at apex; not pellucid punctate; petioles often thickened<br />
tated on the Tafalla's specimen, also supports the at base but not pulvinate; domatia a tuft of hairs (tufttransfer<br />
of this species into Simira, which typically pit domatia in C. jamaicensis) or absent. Infloreshas<br />
red wood.<br />
cences subterminal (axillary on the terminal nodes),<br />
corymbose, with opposite to subopposite lateral<br />
branches terminating in cymules. Flowers<br />
protogynous, sessile to pedicellate, very fragrant;<br />
CHIMARRHIS<br />
hypanthium obconical to turbinate. Calyx extremely<br />
6. Chimarrhis Jacquin, Select. stirp. amer. hist. 61. reduced, truncate or with barely distinguishable lobes,<br />
1763; Chimarhis Rafinesque, orth. var., Ann. Gen. persistent; in few species (e.g., C. brevipes and C.<br />
Sci. Phys. 6: 84. 1820. Jacquin, Select. stirp. amer. gentryana) in some of the flowers one calyx lobe exhist.<br />
77-78. 1788; Jussieu, Mem. Mus. 6: 381. panded into a white to greenish white calycophyll.<br />
1820; de Candolle, Prodr. 4: 403. Sep 1830; A. Corolla funnelform, deeply lobed, with reflexed<br />
Richard, Mem. Rubiac. 278-280. Dec 1830; lobes, white, greenish white or cream-white; tube fun-<br />
Endlicher, Gen. P1. 553 (N. 3260). 1838; Meisner, nelform to ciatiform; with a ring of pubescent hairs<br />
PI. vase. gen. 158 (N. 114). 1838; Grisebach, Fl. inside on the tube or on the base of the lobes; lobes<br />
Br. W.-Ind. 323. 1861; J. D. Hooker in Bentham (4-)5(-6); aestivation narrowly imbricate, superfi-
138 Flora Neotropica<br />
cially resembling valvate aestivation. Stamens 5, al- rolla, followed by the opening of the stigmas (resemternate<br />
to the petals, totally or partially exserted; fila- bling two lips exserted beyond the corolla, Fig. 4J),<br />
ments attached on the upper part of the tube, slender, which then become receptive. Shortly after the corolla<br />
basally flattened, basally pilose to villous; anthers opens, the style is abscissed, the filaments elongate,<br />
elliptic, dorsifixed, dehiscing by lateral slits. Pollen and the anthers are exserted well above the corolla<br />
tricolporate, exine densely thick-reticulate. Style ex- (Fig. 4K), becoming functional and opening by latserted;<br />
style branches protruding above corolla before eral slits. The flowers of this genus are minute (a few<br />
anthesis, very short, rounded to ovate, reflexed at millimeters long), cream-white, and sweet-fragrant.<br />
maturity. Ovary 2-celled, turbinate to obovate, pla- The corollas of Chimarrhis have been described<br />
centation peltate on the septum; ovules many in each as having valvate aestivation, and for this reason this<br />
locule, vertically inserted; immature fruits green and genus was placed in the Condamineeae sensu Hooker.<br />
semi-carnose when fresh. Capsules woody, globose, Close observation of the corollas of Chimarrhis reoblong<br />
or obovoid; dehiscing septicidally, disk-locu- vealed that the aestivation is instead narrowly imbrilicidal<br />
dehiscence present in old capsules of some cate, superficially resembling valvate aestivation.<br />
species. Seeds suborbicular, with central hilum and More specifically, the tips of the lobes do not overlap<br />
narrow concentric wing, margin undulate. (valvate), but the lateral margins of the petals obviously<br />
do overlap (imbricate) in bud and in early stages<br />
Chimarrhis is unique in the <strong>Rubiaceae</strong> in its axil- of anthesis.<br />
lary-subterminal inflorescences, capsules with peltate Chimarrhis is often confused with Calycophyllum<br />
placentation, and minute seeds umbilically attached (Calycophylleae), but the latter has terminal infloresand<br />
with minute concentric wings. The seeds of at cences (vs. lateral subterminal), seeds with strongly<br />
least one species (C. barbata) have a tendency to de- bipolar wings, smooth thin bark exfoliating in long<br />
velop bipolar wings, resembling those of the tribes vertical strips, and white wood (vs. yellow in Chimar-<br />
Cinchoneae and Calicophylleae. All the species of rhis). Chimarrhis is also often confused with Bathysa<br />
Chimarrhis are trees, varying from mid-canopy to (<strong>Rondeletieae</strong>) and Simira (<strong>Rondeletieae</strong>), but the latupper-canopy<br />
levels. Mature individuals of the genus ter two differ from the previous in having terminal<br />
usually develop buttresses, especially those growing inflorescences and different seed morphology. Simira<br />
in the Amazon basin (see Figs. 68A, 71A). The bark differs from Chimarrhis in having larger seeds, horiof<br />
these trees is smooth and grayish in young indi- zontally inserted, laterally attached, with well-develviduals;<br />
and fibrous, deeply fissured, reddish, and oped lateral-orbicular wings, and reddish wood.<br />
falling off in small irregular pieces in mature individu- Bathysa is similar to Chimarrhis in having small<br />
als (see Figs. 67C,D, 69C,D, 76B).<br />
cream-white fragrant flowers, small fruits, minute<br />
The tree architecture of this genus has been stud- seeds, and yellowish wood; the previous differing<br />
ied in only a few species. Chimarrhis microcarpa from the latter in having floral aestivation strongly<br />
(Halle et al., 1978), C. glabriflora (pers. obs.), and imbricate, roundish petals with irregular margins (vs.<br />
C. hookeri (pers. obs.; Fig. 68A) display the narrowly triangular with entire margins in Chimar-<br />
"Aubreville Model (Terminalia branding)," whereas rhis), and horizontal seeds, laterally attached, and<br />
C. cymosa (Halle et al., 1978) develops the "Petit's minute lateral wing (vs. vertical seeds, ventrally attached,<br />
Model," according to the architectural classification minute concentric wing peltate to a central hilum).<br />
of Halle et al. (1978).<br />
After careful observation, I concluded that the<br />
Leafdomatia were found to be a reliable taxonomic species of chimarrhis can be divided into two easily<br />
character, variable within certain species or consis- recognizable and probably monophyletic groups, ustent<br />
within others (Fig. 58A-L; see Domatia, above). ing the persistence and general morphology of their<br />
Chimarrhisjamaicensis and C. cubensis can be rec- stipules, and their geographic distribution: subgen.<br />
ognized solely by their characteristic domatia (Fig. Chimarrhis, with readily caducous stipules; and<br />
58B,C), while in the species with glabrous blades (C. subgen. Pseudochimarrhis (Ducke) Delprete, with<br />
latifolia, C. brevipes, C. gentryana, and C. barbata) persistent stipules. The second subgenus is a new stadomatia<br />
were not found.<br />
tus for the genus Pseudochimarrhis Ducke (1925),<br />
Several species have been reported to be either which he separated from Chimarrhis and transferred<br />
heterostylous or dioecious (Grisebach, 1862; Urban, to the Cinchoneae. To the two species already included<br />
1899), but all Chimarrhis species have strongly in subgen. Pseudochimarrhis (C. turbinata and C.<br />
protogynous flowers, which are neither dioecious nor barbata) I add C. duckeana (described below), C.<br />
heterostylous. Flower anthesis in this genus starts with brevipes, and C. gentryana, all with persistent deltoid<br />
the protrusion of the style above the still-closed co- stipules. Subgenus Pseudochimarrhis occurs mostly
II ~~~~~AIPJ.~~~~~~~~~~~~~~~~~~ .....<br />
~? .~,<br />
:<br />
x\x,qj~<br />
t<br />
I rI J<br />
I<br />
Z~ ~ ~ ,F~<br />
~~~~~~~~~~~~~~~~~~~~~'"'"'<br />
'Ii<br />
" .$ '-....':'<br />
!'." . ~<br />
\ ?<br />
~~~~~~~~~~~~~~ 11X1<br />
"' '/<br />
~ I~<br />
:'.:.A<br />
~<br />
' ': . : :<br />
~'.*" ')?'::<br />
: :-;,.. " .<br />
'0.c<br />
-%~,.. :..? ..<br />
:'.":"- ~~~~?? :):~., ?? ??" ? :<br />
rr~ ~ ~ ~~~~~~~ ..r<br />
'I, ?I*<br />
'"~ ~ ~ ~ ~ ~ ~ ~ ~ ~<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~"<br />
;t? J ~~~ ~ ,,.~~<br />
r ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?I<br />
=<br />
C hooker (topright Nel & Zaruma 7021 NY bottomleft Spruce 4930 K isotype) H. C latfolia J... (Hallen 6302 MO) C brev<br />
~~~~~~~~~~~~~~'-.<br />
??<br />
'"""" ... ~~~'"-..'..;."': " :<br />
II ~<br />
~<br />
.'. ': : .' : '~:. .~. . " - : I;:.~.--:..:'".-,'..-<br />
': .... ;- . ' : :'.~....~ . : ... ; . , - '. .'.': - '<br />
'""'" ":' "'<br />
'":~ . ;.:; , .. '<br />
FIG. 58. Leaf domatia in Chimarrhis, abaxial views and blade cross sections. A. C. parviflora (Harber & Bello 1 735, F). B. C. cu<br />
(Proctor 22591, DAV). D. C cymosa (Webster et a! 9205, DAV). E. C. microcarpa (Berti 241, K). F. C glabriflora (top-right Du<br />
(Vtisquez & Arevalo 8969, TEX). K. C barbata (Kuhlmann RB1 7384, US, isotype). L. C. turbinata (top-right Boom et a! 8756, N
140 Flora Neotropica<br />
in the lowlands of the Amazon basin, while the duced Sprucea (= Simira) and Sickingia (= Simira)<br />
subgen. Chimarrhis is distributed in the Antilles, in to synonymy under Chimarrhis.<br />
Central America, the Andean slopes, and western Schumann (1889, 1891) reported Chimarrhis to be<br />
Amazonian rain forests.<br />
a genus of three species, recognizing C. cymosa, C. tur-<br />
Chimarrhis is here recognized as a genus of 14 binata DC., and describing C. hookeri (Schumann,<br />
species (9 in subgen. Chimarrhis, and 5 in subgen. 1889) from Spruce 4930, previously pointed out by<br />
Pseudochimarrhis).<br />
Hooker (1873). Schumann (1889, 1891) placed<br />
Chimarrhis in the Condamineeae.<br />
Taxonomic History<br />
Urban (1899) subdivided C. cymosa into three<br />
subspecies: subsp. genuina from the Lesser Antilles;<br />
Chimarrhis was founded by Jacquin (1763) with subsp.jamaicensis from material collected in Jamaica;<br />
the description of C. cymosa from Martinique. Jacquin<br />
reported it to be locally called "Bois de riviere," and<br />
and subsp. microcarpa from material collected in<br />
Cuba (Wright 1262).<br />
derived its Latin name from the Greek Xtjqrappy<br />
(chimarro = torrent), because it was originally encoun-<br />
Schumann and Krause (1908a) (four years after<br />
Schumann's death) described Chimarrhis dioica as a<br />
tered in the proximity of torrents. He also reported species exceptional to Chimarrhis in having dioecious<br />
Chimarrhis as having one seed per locule, an error that flowers. This species was later chosen by Steyermark<br />
was reproduced by Poiret (1811).<br />
(1963) as the type species of his new genus Dio-<br />
Achille Richard (1830) placed Chimarrhis cymosa icodendron.<br />
(as "Chimarrhis corymbosa [hort. var.] Jacquin") in Standley (1918) did not recognized Urban's (1899)<br />
synonymy under Macrocnemum, but admitted that the subspecies and reduced them to synonymy (along with<br />
genera were poorly known to him. He correctly noted Macrocnemum longifolium) under C. cymosa. In adthat<br />
the capsules of Chimarrhis are polyspermous and dition, he transferred <strong>Rustia</strong> ferruginea Standl. (=<br />
septicidal, as in M. candidissimum (= Calycophyllum Bathysa sp.) to Chimarrhis. Later, Standley (1926)<br />
candidissimum), M. jamaicense, and M. tetrandrum. described C. microcarpa based on specimens collected<br />
Because he erroneously reported C. cymosa as C. cor- in Trinidad, and C. pittieri (Standley, 1930b) based<br />
ymbosa, when including this species into Macrocnemum, on specimens collected in Venezuela. Standley<br />
the combination M. corymbosum (= Condaminea (1931c, 1936) merely listed C. dioica (= Dioicocorymbosa)<br />
already existed, and he had to establish dendron dioicum) and C. pittieri (= Bathysa pittieri),<br />
the new name M. longifolium (based on Jacquin's maintaining Chimarrhis in the Condamineeae (sensu<br />
collection from Martinique).<br />
Hooker).<br />
De Candolle (1830) recognized Chimarrhis as a Adolpho Ducke (1876-1958) was very influential<br />
good genus, placing Macrocnemum longifolium (and<br />
Chimarrhis corymbosa) under C. cymosa, and dein<br />
understanding the phenology and clarifying the<br />
taxonomy of the Amazonian species of Chimarrhis,<br />
scribing C. turbinata on the basis of material collected<br />
by Patris in Cayenne (French Guiana). He placed<br />
primarily due to his 50 years of accurate fieldwork in<br />
the Brazilian Amazon. He often collected one tree in<br />
Chimarrhis in the tribe Hedyotideae, subtribe flowering stage and returned to the same individual<br />
Rondeletiinae (as "<strong>Rondeletieae</strong>").<br />
Joseph Dalton Hooker (1873) attributed four speseveral<br />
months later to collect its fruiting material,<br />
mounting both collections on the same herbarium<br />
cies to Chimarrhis: two from the West Indies (but only<br />
one species was described at that time!), the third<br />
species C. turbinata, and the fourth from a Spruce's<br />
collection (Spruce 4930) from Tarapoto, Peru, but<br />
without giving it a name. He placed Chimarrhis in his<br />
newly founded tribe Condamineeae, subtribe Condamineinae<br />
(as "Eucondamineae").<br />
Baillon (1880) placed Chimarrhis in his "Cinchona<br />
Series" (or Cinchoneae). He reported it "exceptional<br />
sheet. Ducke (1922) separated Pseudochimarrhis<br />
(with C. turbinata as type species) from Chimarrhis,<br />
stating that the latter genus differed from the former<br />
in having narrower, longer flowers, and vertically<br />
arranged seeds, characters that would place it in the<br />
Cinchoneae. Three years later, Ducke (1925) added a<br />
second species to Pseudochimarrhis (P. barbata).<br />
Benoist (1920) described Bathysa difformis (= C.<br />
turbinata), which he later transferred to Pseudoin<br />
this group as it would be in any other in which it<br />
might be placed," later adding that its "stamens are<br />
of two kinds according as we examine this or that<br />
flower with filaments short or sometimes long" (probably<br />
suggesting dimorphic flowers, which do not exist<br />
in any Chimarrhis species), and erroneously rechimarrhis<br />
(Benoist, 1933).<br />
Bremekamp (1934) stated that the separation of<br />
Pseudochimarrhis from Chimarrhis, proposed by<br />
Ducke (1922, 1925) was unacceptable, reporting:<br />
"The seeds of C. cymosa Jacq., the type species of the<br />
genus, however are arranged in exactly the same way
Systematic Treatment of the Genera Studied 141<br />
as those of C. turbinata, and the longer and narrower duced Pseudochimarrhis to Chimarrhis and synonyflower<br />
is a character of so little importance that it is mized the other species accordingly. He maintained<br />
in itself not sufficient to justify the creation of a new Chimarrhis barbata and C. turbinata under Chimargenus.<br />
Pseudochimarrhis turbinata (DC.) Ducke is rhis; described C. brevipes and C. bathysoides [= Bathysa<br />
therefore referred back to Chimarrhis and P. barbata bathysoides ((Steyerm.) Delprete]; raised C. cymosa<br />
Ducke became Chimarrhis barbata (Ducke) Brem. subsp. jamaicensis Urb. and subsp. microcarpa to<br />
comb. n." In the same work Bremekamp described C. specific rank; maintained C. williamsii, C. hookeri,<br />
longistipulata, which is here treated as synonymous C. glabriflora, and C. parviflora as previously dewith<br />
C. microcarpa (see discussion under this spe- scribed; and divided C. microcarpa into var. microcies).<br />
Pseudochimarrhis difformis has never been carpa and var. speciosa (which are here raised to spepositioned<br />
in Chimarrhis, but this species is synony- cific rank). Finally, in the list of excluded species,<br />
mous with C. turbinata.<br />
Steyermark (1965) reduced C. dioica and C. venezuel-<br />
Rizzini (1947) treated Chimarrhis turbinata as a ensis to synonymy under Dioicodendron dioicum;<br />
dubious taxon, proposing C. duckei Rizz. (nom. treated C. decurrens as synonymous with Allenanthus<br />
superfl.) as a substitute name for Pseudochimarrhis erythrocarpus Standl.; transferred C. goudotii, C. pisoniturbinata<br />
(DC.) Ducke.<br />
aeformis, and C. pittieri to Sickingia [= Simira]; and<br />
Steyermark (1965) published the only monograph returned C. ferruginea to <strong>Rustia</strong>.<br />
of Chimarrhis, where he recognized 12 species, and Dwyer (1980) and Burger and Taylor (1993) recwith<br />
only the new species fully described. Following ognized Chimarrhis as a genus of 14 species ranging<br />
Bremekamp's (1934) observations, Steyermark re- from Costa Rica to South America and the West Indies.<br />
Key to the subgenera and species of Chimarrhis<br />
1. Stipules readily caducous before adjacent leaves expansion (Central America, Antilles, and South<br />
American Andes) ....................................................................................... subgen. Chimarrhis<br />
2. Capsular disks glabrous.<br />
3. Anthers 1-2 mm long (Antilles).<br />
4. Domatia absent or few hairs present; young leafy branchlets 3-4 mm thick; leaf blades<br />
7-11 x 3-5 cm, with 7-8 secondary veins each side; inflorescences 7-10 cm long;<br />
capsules 2-2.5 x 2 mm (Cuba) ............................................................ .................... 3. C. cubensis<br />
4. Domatia well-developed as tufts in most vein axils; young leafy branchlets 4-15 mm<br />
thick; leaf blades 10-47 x 5-20 cm, with 8-15 secondary veins each side; inflorescences<br />
>10 cm long; capsules 3-5 x 2.5-3.5 mm.<br />
5. Tuft-domatia without pit; leafy branchlets 10-15 mm thick, semi-succulent, costate;<br />
leaf blades 21-47 x 10-20 cm, with 11-15 secondary veins each side; inflorescences<br />
15-24 cm long; capsules 3-4 x 2.5-3 mm (Lesser Antilles) ............................................ 5. C. cymosa<br />
5. Tuft-domatia with pit; leafy branchlets 4-7 mm thick, woody, not costate; leaf blades<br />
(8-)10-16 x 5.5-8 cm, with 8-9 secondary veins each side; inflorescences (10-)12-22<br />
cm long; capsules 4-5 x 3-3.5 mm (Jamaica and Haiti) ............................................ 4. C. jamaicensis<br />
3. Anthers 0.5-0.7 mm long (Central and South America).<br />
6. Leafy branchlets 9-12 mm thick, succulent to semi-succulent, smooth (Colombia, Ecuador,<br />
Peru, and B razil) ................................................................................................................... 6. C. glabriflora<br />
6. Leafy branchlets 1.5-8 mm thick, woody, exfoliating in small longitudinal strips.<br />
7. Domatia absent; stipules 6-25(-30) mm long, 2-4 mm wide at base; inflorescences<br />
4-12 cm long; leaf blades 5-15(-20) x 4-7.5 cm, with length:width ratio 2:1 to 2.5:1;<br />
capsules obovoid to turbinate, acute at base (Nicaragua, Costa Rica, Panama) ......... 1. C. parviflora<br />
7. Tuft-domatia present at least in some axils; stipules 25-35 mm long, 5-7 mm wide at<br />
base; inflorescences 10-22 cm long; leaf blades (9-)14-24 x 6.5-12 cm, with length:width<br />
ratio 1.5:1 to 2:1; capsules subglobose, rounded at base (Venezuela, Trinidad and Tobago,<br />
the Guianas, Brazil).<br />
8. Corolla funnelform, 4-5 mm long; inflorescences very compactly branched (L/A =<br />
3:1); leafy branchlets 2.5-5(-6) mm thick (Venezuela, Trinidad and Tobago, the<br />
Guianas and Brazil) ........................................ ....................... 8. C. microcarpa<br />
8. Corolla campanulate, 1.6-2.4 mm long; inflorescences laxly branched (L/A = 2:1);<br />
leafy branchlets (4-)5-8 mm thick (Venezuela) ....................................................... 9. C. speciosa<br />
2. Capsular disks puberulent to pubescent.
142 Flora Neotropica<br />
9. Capsules 2-2.5 mm long; corollas 2.5-3.5 mm long (Ecuadorand Peru) ................................... 7. C. hookeri<br />
9. Capsules 4-5 mm long; corollas 4-5 mm long (Costa Rica) ........................................................ 2. C. latifolia<br />
1. Stipules persistent after adjacent leaves abscission (mostly Amazon basin) .......................... subgen. Pseudochimarrhis<br />
10. Calycophylls present at least in some of the flowers; leaf blades pandurate.<br />
11. Leaf blades with 5-6 secondary veins each side, length:width ratio 1:1 to 1.5:1; calycophylls<br />
obovate; inflorescence L/A = 2.5-3:1; basal portion of axis not branched ["peduncle"] (3.5-)<br />
6-9.5; capsules obovoid (Venezuela) ............ ..................................................................... 13. C. brevipes<br />
11. Leaf blades with 7-10 secondary veins each side, length:width ratio 1.5:1 to 1.7:1; calycophylls<br />
ovate-elliptic; inflorescence L/A = 1.5-2:1; basal portion of axis notbranched 4.5-7.5; capsules<br />
narrowly obconical to turbinate (Amazonian Ecuador and Peru, and W Brazil). ................... 14. C. gentryana<br />
10. Calycophylls absent (rarely present in 1-2 flowers per inflorescence); leaf blades elliptic, ovate, or<br />
obovate (not pandurate).<br />
12. Disk minutely puberulent, obviously exceeding the calyx (Brazil: Amazonas).<br />
13. Leaf blades 5.5-10(-14) x 4-7 cm; leafy branchlets thin (2-4 mm wide), woody; capsules<br />
w ith oblong-ovoid disks .......................................................................................................... 11. C. barbata<br />
13. Leaf blades >12.5 x 7 cm; leafy branchlet thick (4-6 mm wide), succulent; capsules with<br />
obtuse to rounded disks . ....................................... .................... 12. C. duckeana<br />
12. Disk pubescent to pilose, not exceeding the calyx (Surinam, Brazil: Amapa, Para, Amazonas).<br />
........................................................................................................................................................ 10 . C . tu rb in a ta<br />
CHIMARRHIS JACQUIN SUBGEN.<br />
CHIMARRHIS<br />
1. Chimarrhis parviflora Standley, Trop. Woods 11:<br />
26.1927. Type. Panama. Bocas del Toro: Changui-<br />
nola Valley, 1927 (fl), Cooper & Slater 120 (ho-<br />
lotype, US, N.1315890; isotypes, A, F, G, GH,<br />
NY). Figs. 9A, 17A,B, 58A, 59A-G, 61<br />
Trees to 30 m tall, 40-50(-60) cm dbh, with clear<br />
bole to 10 m, with small buttresses or flutes; bark soft,<br />
pale brown; wood yellow to golden-yellow, very brittle.<br />
Leafy branchlets 1.5-4 mm thick, terete to compressed,<br />
ochraceous, glabrous to appressed puberulent; older<br />
branches glabrate, terete, grayish to pale brown.<br />
Stipules free at base, contorted in bud, narrowly triangular,<br />
acuminate, appressed puberulent to gray-sericeous<br />
outside, glabrous with basal colleters inside, 6-<br />
branchlet, 4.5-12 x 3-7 cm; lateral branches 1-2 pairs,<br />
basal portion of axis not branched 2-7 cm long; rachis<br />
decussately compressed to subterete, reddish<br />
brown; L/A = 1.6-2.1; rachis and branches densely<br />
curled to appressed gray-puberulent; distal bracts<br />
(when present) deltoid, ca. 0.5 mm long; bracteoles<br />
subtending flowers absent. Flowers pedicellate;<br />
pedicels 0.5-2 mm long, glabrous to minute puberulent;<br />
hypanthium turbinate, 1-1.5 mm long, glabrous.<br />
Calyx extremely reduced with wavy margin or barely<br />
distinguishable lobes, to 0.5 mm long; lobes to 0.4 mm<br />
long, ovate to deltoid, ciliolate at apex. Corolla funnelform,<br />
deeply lobed, 2-4 mm long, white to creamwhite;<br />
tube short cylindrical, 1-1.5 x 1-1.5 mm, glabrous<br />
outside and inside; lobes (4-)5, 1.5-2 x 1-1.5<br />
mm, narrowly ovate, truncate at apex, glabrous outside,<br />
densely villous at medio-basal zone. Stamens<br />
(4-)5, exserted well above the corolla, equal, attached<br />
1-1.5 mm from the base of the tube; filaments 3-4<br />
mm long, terete, basally flattened, connate to throat,<br />
densely villous at base; anthers elliptic, 0.6-0.7 x 0.2-<br />
0.4 mm, dorsifixed at medial zone, reversed at matu-<br />
25(-30) x 2-4 mm, reddish with a grayish tinge, readily<br />
caducous, leaving a white scar encircling the stem, 0.5-<br />
1 mm wide. Leaves 6-17(-22) x 4-7.5 cm, L/W = 2:1<br />
to 2.5:1, elliptic to oblong-obovate, acute-decurrent<br />
(ca. 25?) at base, obtuse and mucronate at apex, the<br />
deltoid acumen 0.7-1 cm long; dark green above, pale rity, base rounded. Pollen exine irregularly foveolategreen<br />
below, foliaceous; drying olive-green to brown, reticulate. Style exserted, 2-4 mm long, as long as the<br />
stiff-chartaceous; glabrous above, glabrous to sparsely corolla, glabrous, minutely costate; style branches<br />
appressed puberulent below, hairs whitish to yellow- extruding as two lips above corolla before anthesis,<br />
ish; primary, secondary, and tertiary veins appressed widely ovate to widely oblong, 0.3-0.4 mm long, stigpuberulent<br />
below; secondary veins (5-)8-14 each side; matic surface microscopically (40x) papillose; ovatertiary<br />
veins starting subparallel and reticulate in the ries often infested by insects, developing into globose<br />
center; petioles 10-23 mm long, 1-2 mm thick, ter- galls containing minute yellow larvae. Capsules oboete,<br />
glabrescent to appressed minute-puberulent; void to turbinate, acute at base, apex rounded, 1.5domatia<br />
absent. Inflorescences usually 2 (rarely 3-4) per 2.5 x ca. 1.5 mm, dark brown without lenticels, glanode,<br />
corymbose with opposite to subopposite decus- brous throughout; disk loculicidal dehiscence present<br />
sate branches, densely cymose at distal end of each in old capsules. Seeds 0.5-0.88 x 0.38-0.5 mm.
Systematic Treatment of the Genera Studied 143<br />
C<br />
iH^HBRl IIf I<br />
I<br />
FIG. 59. Chimarrhis parviflora (A-F from Cooper & Slater 120, US, holotype; G from Jimenez 3418, F). A. Habit of<br />
inflorescences. B. Flower with exserted anthers. C. Variation in calyx lobes. D. Detail of corolla, inside view. E. Stamen.<br />
F. Apical portion of style. G. Mature capsules.
144 Flora Neotropica<br />
Distribution (see Fig. 61) and ecology. Primary<br />
and secondary rain forests, and respective remnants,<br />
from low to medium elevations (Bosque pluvial<br />
premontano), 100-900 m, of the Caribbean slopes<br />
(rarely of the Pacific slopes), usually in proximity of<br />
creeks and small streams, in alluvial soil or volcanic<br />
loam of Nicaragua, Costa Rica, and Panama. Flower-<br />
ing specimens were collected in March, April, May,<br />
June, and July. Fruiting specimens were collected in<br />
May, July, August, and September.<br />
Colorado Island, trail 30 m S of Creek 7, 13 Jul 1970 (fr),<br />
Croat 11244 (MO[2], VEN). VERAGUAS: Valley of Rio Dos<br />
Bocas, between Escuela Agricola Alto Piedra and Calovebora,<br />
NW of Santa Fe, 08?30'N, 81 ?10'W, 400-450 m, 31<br />
Aug 1974 (fr), Croat 27755 (MO).<br />
Local names and uses. Costa Rica: higueron<br />
(Hartshorn 1211), pejiballito (Dayton & Barbour<br />
3007). Panama: yema de huevo (Cooper 401), llema<br />
de huevo (Cooper & Slater 3/10708), man wood (Dunlap<br />
33). As in most species of this genus, C. parviflora is<br />
reported to be good timber wood.<br />
Representative specimens examined. NICARAGUA.<br />
ZELAYA: Cerro El Escobin, 11?33'N, 84?21,W, 30 Jul 1982<br />
(fr), Sandino 3412 (MO); Colonia Serrano, on trail to Colonia This species is easily distinguished from the other<br />
Yolaina, 100 m, 1 1?35'N, 84?22'W, 28 Jul 1982 (fl), Sandino Central American taxa by its minute capsules (Fig.<br />
3294 (MO); Kurimwuasito, 50 km SE of Rio Blanco, 15 Mar 59G) with glabrous disk, and its slender-woody leafy<br />
1984 (fl), Sandino 4842 (MO); Kurinwacito, 13?09'N, branchlets. It is commonly encountered as a tree with<br />
84?55'W, 100 m, 23 Mar 1984 (fl), Moreno 23868 (MO). soft flaky white bark, with wood quickly turning rust-<br />
Rio SAN JUAN: Buenos Aires, 1 km N of Sabalo, above Rio colored when cut (probably due to its alkaloid con-<br />
Sabalo, 1 102'N, 84?28'W, 70 m, 10 Sep 1985 (fr), Moreno<br />
tent); leaves thickly membranous, waxy dark green<br />
26317 (MO).<br />
and faintly shiny above; and<br />
COSTA RICA. ALAJUELA: 2 km N of Bijagua, trail<br />
vegetative buds lightto<br />
San Miguel, 400-500 m, 8 Jul 1976 (fr), Utley & Utley 5337 green to whitish, with readily caducous stipules. The<br />
(CAS, F); Reserva Biol6gica Monteverde, Rio Peias young ovaries are often infected by insects which<br />
Blancas, 10?20'N, 84?43'W, 850 m, 28 Mar 1987 (fl), Haber cause them to enlarge abnormally into globose galls<br />
& Bello 6837 (F, GH, MO, TEX); Upala, Colonia Libertad, (sometimes mistaken for mature fruits).<br />
Valley of Rio Cucaracho, 450 m, 6 May 1988 (fl), Herrera Chimarrhis parviflora is most similar to C.<br />
1976 (F, MO); rd. Canas-Upala, 13.8 km N ofBijagua, 100- microcarpa (from Venezuela and the Guianas), the<br />
150 m, 26 Jun 1976 (fl), Croat 36431 (MO); Cton. San former differing from the latter in having the termi-<br />
Carlos, Villa Guisada, 725 m, 14 Apr 1939 (fl), Smith 1890 nal nodes of the branchlets appressed-puberulent; leaf<br />
(A, F, MO, NY). CARTAGO: Turrialba, Instituto, 600 m, 1 veins ascending-pubescent (vs. glabrous), domatia<br />
Sep 1950 (fr), L&on 2143 (F); Turrialba, E of Pavones, 700<br />
absent (vs.<br />
m, 8 May 1972<br />
present;<br />
(fr), Holdridge 6723 (F). HEREDIA: Finca La<br />
Figs. 58A, 58E); corollas 2-4 mm<br />
Selva, Sabalo-Esquina trail, 10?26'N, 84001'W, 30 long (vs. 4-5 mm long; Figs. 59B-D, 71C,D); corolla<br />
May<br />
1974 (fl), Hartshorn 1500 (F, MO); Finca La Selva, OTS<br />
lobes acute at apex (vs. obtuse to truncate); infloresfield<br />
station, South Boundary trail, 100 m, 21 May 1985 cences with L/A =<br />
(fl),<br />
3:1 (vs. L/A = 2:1; Figs. 59A, 71B),<br />
Wilbur 37205 (CAS, F, K, MO, US) vic. of Puerto Viejo, and basal portion of axis not branched 2-7 cm long<br />
canal entering Rio Sarapiqui, 100-150 m, 11 Aug 1965 (fr), (vs. 7 cm or more).<br />
Jimenez 3418 (F, MO, NY, US); 1 km E of Cariblanco, 850<br />
m, 21 May 1973 (fl), Hartshorn 1211 (F). LIM6N: Cerro<br />
Coronel, E of Laguna Danto, 10?4 'N, 83?38'W, 60-100 m,<br />
15 Mar 1987 (st), Stevens et al. 24859 (MO); La 2. Chimarrhis latifolia<br />
Florida,<br />
Standley, Publ. Field<br />
250 m, 7 May 1943 (fl), Dayton & Barbour 3007 (MO, US).<br />
Columbian Mus., Bot. Ser. 4: 265. 1929. Type.<br />
PUNTARENAS: Osa Peninsula, Reserva Forestal Golfo Dulce, Panama. Chiriqui: Progreso, Jul-Aug 1927 (fr),<br />
Rancho quemado, 08?43'N, 83?35'W, 200-400 m, 28 May Cooper & Slater 260 (holotype, F, N.573,160;<br />
1988 (fl), Hammel et al. 16858 (F, MO). SAN JOSE: Parque photo-F at F, G, TEX, VEN; isotype, F).<br />
Nacional Braulio Carrillo, Estacion Carrillo, 500 m, 25 Jul<br />
Figs. 9B, 17C,D, 58H, 60A-F, 61<br />
1984 (fr), G6mez et al. 22944 (F, MO); Parque Nacional<br />
Braulio Carrillo, path beyond Rio Sucio, 400-500 m, May Trees to 30 m tall, 30 cm or more dbh, canopy trees<br />
1984 (fl), G6mez et al. 22770 (MO, NY, VEN).<br />
with long buttresses; wood yellow, heavy, very brittle.<br />
PANAMA. BocAS DEL TORO: Region of Almirante, Daytona<br />
Farm, Jan-Mar 1928 (fr), Cooper & Slater 401<br />
Leafy branchlets 4-9 mm thick, subterete to com-<br />
(A, F,<br />
K, US); Changuinola Valley, 1927, Cooper & Slater 3 pressed, ochraceous, glabrous; older branches terete,<br />
(F,<br />
US); Changuinola Valley, 1 Aug 1923 (fr), Dunlap 33 (F,<br />
grayish to pale brown. Stipules free at base, contorted<br />
G, US). COL6N: Along Rio Guanche, S of Portobelo in<br />
Hwy., bud, narrowly triangular, acuminate, glabrous with<br />
09030'N, 79?40'W, 50 m, I 1 Apr 1982 (fl), Knapp et al. 4586 ciliate margins outside, glabrous with basal colleters<br />
(LL). PANAMA: Barro Colorado Island, slope 300 m S of Bat inside, 20-35 x 5-13 mm, reddish brown, readily<br />
Cove, 16 Jul 1978 (fr), Foster 2877 (K, MO, US); Barro caducous, leaving a dark-brown scar encircling the
Systematic Treatment of the Genera Studied 145<br />
tr q<br />
,aW~~~~~~~~~~~~~~~~~~~~~,<br />
?- .I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
FIG. 60. Chimarrhis latifolia (A-E from Allen 6302, MO; F from Allen 5762, F). A. Habit of inflorescences. B. Flower<br />
with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style. F. Mature capsule.
146 Flora Neotropica<br />
FIG. 61. Distribution of Chimarrhis parvifolia (triangles) and C. latifolia (circles).<br />
stem, 0.5-1 mm wide. Leaves (13-)16-28 x (5-)7-<br />
11 cm, L/W = 1.5:1 to 2:1; broadly elliptic to ellipticobovate,<br />
acute-decurrent at base, obtuse and mucronate<br />
at apex, the deltoid acumen 0.5-1 cm long; dark<br />
green above, pale green below, foliaceous; drying<br />
brown above, olive-green below, chartaceous; glabrous<br />
above and below; primary and secondary veins<br />
glabrous, secondary veins 7-12 each side; tertiary<br />
veins starting subparallel and reticulate in the center;<br />
petioles 20-45 mm long, 1.3-1.8 mm thick, terete to<br />
adaxially concave, glabrous; domatia absent or rarely<br />
a few hairs at the axil. Inflorescences 2 per node,<br />
corymbose with opposite to subopposite decussate<br />
branches, densely cymose at distal end of each<br />
branchlet; 8-15 x 8-13 cm, lateral branches 1(-2)<br />
pairs, basal portion of axis not branched 2.5-7 cm<br />
long; rachis decussately compressed to subterete, reddish<br />
brown, rachis and branches glabrous to sparsely<br />
minute puberulent; distal bracts usually absent or deltoid,<br />
1-2 mm long, ciliolate; bracteoles subtending<br />
flowers absent. Flowers sessile to short-pedicellate;<br />
pedicels (when present) to 3 mm long, minute puberulent;<br />
hypanthium turbinate, 1.5-3 mm long, glabrous,<br />
reddish brown. Calyx extremely reduced with wavy<br />
margin or barely distinguishable lobes, 0.5-1 x 2-2.5<br />
mm; lobes (when present) to 2 mm long, shallowovate<br />
to shallow-triangular. Corolla funnelform,<br />
deeply lobed, 4-5 mm long, white to greenish white;<br />
tube short cylindrical, 1-2 x ca. 1.5 mm, glabrous<br />
outside and inside; lobes 5, 2.5-3.5 x 1-1.5 mm, oblong<br />
and rounded to truncate at apex, glabrous outside,<br />
densely villous at medio-basal zone. Stamens<br />
5, exserted well above the corolla, equal, attached 2.5-<br />
4 mm from the base of the tube; filaments 4-5 mm<br />
long, terete, basally flattened, connate to throat,<br />
densely white-villous at base; anthers elliptic, 0.7-1<br />
x 0.3-0.4 mm; dorsifixed at medial zone, reversed at<br />
maturity, base rounded. Pollen exine reticulate-foveolate.<br />
Style exserted, 3.5-5 mm long, as long as the<br />
corolla, glabrous, minutely costate; style branches<br />
extruding as two lips above corolla before anthesis,<br />
rounded, ca. 1 mm long, stigmatic surface microscopically<br />
(40x) papillose; ovary-galls not found. Capsules<br />
obovoid to oblong, acute at base, apex rounded to<br />
subtruncate, 4-5 x 3-3.5 mm, dark brown without<br />
lenticels, glabrous below the disk; disk black, densely<br />
minute-puberulent; disk loculicidal dehiscence present<br />
in old capsules. Seeds 0.49-0.78 x 0.33-0.58 mm.
Systematic Treatment of the Genera Studied 147<br />
Distribution (Fig. 61) and ecology. Primary and<br />
secondary forests, forming stands of canopy trees,<br />
sometimes in secondary forests in association with<br />
Ochroma and Schizolobium, or in remnant patches of<br />
mature forests at low elevations (0-300 m) of the<br />
Pacific coast of Costa Rica and Panama. Flowering<br />
specimens were collected in October, and one speci-<br />
men (Jimenez & Zuniga 760) in late blooming-early<br />
fruiting was collected in December. Mature fruiting<br />
specimens were collected only in January.<br />
Field Mus. Nat. Hist., Bot. Ser. 11: 192. 1936.<br />
Chimarrhis cymosa Jacquin subsp. microcarpa<br />
Urban, Symb. Ant. 1: 411. 1899. Type. Cuba.<br />
Granma: Monte Verde, Jan-Jul 1859 (fl), Wright<br />
1262 (lectotype, G, here selected; isolectotypes,<br />
BR, G, HAC[5], K, P).<br />
Figs. 9C, 17E, 58B, 62A-F, 63<br />
Trees to 10-20(-30) m tall, without or with very<br />
small buttresses; bark light brown, soft; wood yellow,<br />
very hard. Leafy branchlets 3-4 mm thick, terete,<br />
Reproductive biology. The white, fragrant flowglabrous;<br />
older branches grayish to pale brown; leners<br />
are reported to be visited by bees and butterflies<br />
ticels not found. Stipules free at base, contorted in<br />
(Jimenez & Zuniga 760).<br />
bud, narrowly triangular, acuminate, glabrous outside,<br />
Specimens examined. COSTA RICA. PUNTARENAS: glabrous, with two basal rows of colleters inside, 15-<br />
Vic. of Jalaca Station, 30 m, 8 Oct 1951 (fl), Allen 6302 (DS, 25 x 5-8 mm, reddish brown, readily caducous, leav-<br />
F[2], GH, MO, US); Esquinas forest preserve, 0 m, 10 Jan ing a whitish scar encircling the stem, 0.5-1 mm wide.<br />
1951 (fr), Allen 5762 (DS, F, GH, US); Reserva Biol6gica Leaves (5-)8-12.5<br />
Carara, Lomas Pizote, Bijagual trail, 09?47'N, 84?35'W, 300<br />
m, 8 Dec 1989 (fl-fr), Jimenez & Zuniga 760 (CR, F, K[2],<br />
MO); vic. of Palmar Norte de Osa, Rio Terralba, 20 Aug<br />
1950 (st), Allen 5613 (F); Cant6n de Osa, Fila Retinto, Quebrada<br />
Benjamin, 08?59'N, 83?28'W, 50-200 m, 3 Jan 1990<br />
(fr), Hammel et al. 17719 (CR, F, MO); Cant6n de Osa,<br />
Peninsula de Osa, trail to Rancho Quemado, 08?40'N,<br />
83?31'W, 100m, 19Nov 1992 (fr),Zamoraetal. 1901 (NY);<br />
Corcovado National Park, Sirena Field Station, 08?30'N,<br />
83?35'W, 14 Aug 1984 (st), Gentry & OTS class 48572<br />
(MO). SAN Jost: Cerros de Turrubales, Lagunas, 09?50'N,<br />
84?31'W, 200 m, 15 Feb 1993 (fr), Morales et al. 1124 (NY).<br />
Local names and uses: Costa Rica: yema de huevo<br />
(Allen 6302). Panama: jagua amarilla (Cooper & Slater<br />
260). As in most other species, the wood of this species<br />
is used in construction. One label (Cooper & Slater<br />
260, holotype) bears the following comments: "Tree<br />
25 m, with high buttresses and yellow wood. Wood<br />
heavier and harder to split than 'Jagua blanca', grain<br />
very crooked and crossed, making it hard to split."<br />
This species is easily distinguishable from the other<br />
species present in Central America (C. parviflora) by<br />
its glabrous condition, larger leaves, larger capsules,<br />
and absence ofdomatia. It has been suggested (Burger<br />
& Taylor, 1993) that this species may be synonymous<br />
with C. cymosa (of the Lesser Antilles), but the latter<br />
differs in having succulent (with large spongy pith)<br />
costate (Fig. 65A), leafy branchlets (1-1.5 cm thick!),<br />
characteristic tuft-domatia (Fig. 58D), and much<br />
larger leaves.<br />
3. Chimarrhis cubensis Steyermark, Mem. New<br />
York Bot. Gard. 12(3): 184. 1965. Chimarrhis<br />
cymosa Jacquin var. microcarpa (Urban) Standley,<br />
x 3-5 cm, L/W = (2:1) 2.5:1; elliptic<br />
to elliptic-obovate, acute-decurrent at base, acute<br />
at apex, with a deltoid acumen ca. 7 mm long; dark<br />
green above, pale green below, semi-coriaceous; drying<br />
brown above, olive-green below, semi-coriaceous;<br />
glabrous above, sparsely yellow-pubescent below;<br />
primary and secondary veins erect pubescent to hirsutulous,<br />
secondary veins 7-8 each side; tertiary veins<br />
mainly reticulate (rarely subparallel); petioles 6-10 mm<br />
long, 1-1.5 mm thick, adaxially flattened, minutely<br />
winged, glabrous; domatia absent or rarely a few hairs<br />
at the axil. Inflorescences 2(-3) per node, corymbose<br />
with opposite decussate branches, densely cymose at<br />
distal end of each branchlet; 7-10 x 2.5-5 cm, one<br />
pair of lateral branches, basal portion of axis not<br />
branched 5.5-7.5 cm long; rachis decussately compressed<br />
to subterete, reddish brown, rachis and<br />
branches yellow short-puberulent; distal bracts absent<br />
or reduced to deltoid scales 0.5-1 mm long; bracteoles<br />
subtending flowers absent. Flowers sessile to shortpedicellate;<br />
pedicels (when present) to 1.5 mm long,<br />
minute puberulent; hypanthium turbinate, 1-1.5 mm<br />
long, glabrous, reddish brown. Calyx extremely reduced<br />
with wavy margin (rarely with 5 minute teeth),<br />
0.5 x 1.5 mm. Corolla funnelform, deeply lobed, 4-<br />
5 mm long, cream-white; tube short cylindrical, 1.5-<br />
2 x ca. 1.5 mm, glabrous outside and inside; lobes 5,<br />
2-3 x ca. 1 mm, oblong-ovate, rounded to truncate at<br />
apex, glabrous outside, white-villous at medial zone,<br />
the remaining basal and distal zones glabrous inside.<br />
Stamens 5, exserted well above the corolla, equal,<br />
attached ca. 2 mm from the base of the tube; filaments<br />
3-4 mm long, terete, basally flattened, connate to<br />
throat, densely white-villous at base; anthers narrowly<br />
elliptic, rectilinear, 1.5-2 x ca. 0.3 mm, dorsifixed<br />
near the base, reversed at maturity, base rounded, truncate<br />
apiculate at apex. Pollen exine foveolate-reticu-
148 Flora Neotropica<br />
FIG. 62. Chimarrhis cubensis (A-E from Wright 1262, BR, isotype; F from Wright 1622, BR, paratype). A. Habit of<br />
inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style.<br />
F. Mature capsule.
Systematic Treatment of the Genera Studied 149<br />
s0 70 so<br />
FIG. 63. Distribution of Chimarrhis cubensis (circles),<br />
* 0^0<br />
. . -cymosa (triangles).<br />
/ Tc.r.c, V? ^i<br />
FIG. 63. Distribution of Chimarrhis cubensis (circles), C. jamaicensis (squares) and C. cymosa (triangles).<br />
late. Style exserted, 3.5-5 mm long, as long as the<br />
corolla, glabrous, minutely costate; style branches<br />
extruding as two lips above the corolla before anthe-<br />
sis, rounded, ca. 0.5 mm wide, stigmatic surface mi-<br />
croscopically (40x) papillose; ovary-galls not found.<br />
Capsules globose to subglobose, obtuse at base, apex<br />
shallowly rounded, 2-2.5 x ca. 2 mm, dark brown<br />
without lenticels, glabrous throughout; disk black;<br />
disk loculicidal dehiscence present in old capsules.<br />
Seeds 0.58-0.87 x 0.48-0.77 mm, more or less un-<br />
gulate, yellow.<br />
Distribution (Fig. 63) and ecology. Primary and<br />
secondary seasonal forests (pluvisilvas de montafia),<br />
in acid soils, on low mountain ranges, 700-1000 m.<br />
Endemic to Cuba. Flowering specimens were col-<br />
lected in January, February, and May. Fruiting speci-<br />
mens were collected in July and August.<br />
Conservation status. This species is known from<br />
relatively few collections and appears to be endan-<br />
gered. Some attempt should be made to protect the<br />
localities where it occurs.<br />
Specimens examined. CUBA. GRANMA: Nuevo Mundo,<br />
vic. of Rio Macaguanigua, 13 Jan 1972 (fl), Del Risco 27343<br />
(HAC); Rio Jaguani, 3 Feb 1973 (fl), Del Risco 27395<br />
(HAC); Sierra Maestra, Salida Cala, 17 May 1948, Acuna<br />
s.n. (HAC[2]); Toa, 19 Mar 1952, Smith 396 (HAC); Sierra<br />
Maestra, vic. of Sevilla, 800-900 m, 18 Jul 1918 (st), Ekman<br />
9417 (G). GUANTANAMO: Finca La Prenda, Jul 1922,<br />
Hioram & Maurel 6066 (HAC, NY); Camino La Melba, km<br />
26,31 Mar 1972, Del Risco 27458 (HAC). HOLGUIN: Sierra<br />
de Nipe, Loma Colorado, vic. of Rio Piloto, 10 Jul 1919 (fl),<br />
Ekman 9692 (G, NY). PINAR DEL Rio: Banks of Rio Santa<br />
Cruz, 1861, Wright 1622 (BR, G[2], HAC[2]); San Crist6bal,<br />
6 Dec 1915, Roig 6419 (HAC); San Diego, Finca Cortina,<br />
Mar 1910, Fors 142 (HAC); Valley of Taco-taco River,<br />
Rangel, Aug 1926, Le6n 12813 (HAC, NY). SANTIAGO DE<br />
CUBA: Pico Turquino, Loma del Sabici, 1000 m, Jul 1922,<br />
Le6n 11011 (HAC, NY); Loma del Gato, Cobre Range of<br />
Sierra Maestra, 900 m, Jul-Aug 1921, Le6n et al. 9941<br />
(HAC, NY).<br />
Local names and uses: Cuba: jilacho (Ekman<br />
9692), hilacho (see below), penda (pers. comm.), roble<br />
amarillo (pers. comm.), cera (see below). Roig and<br />
Mesa (1928) report the following (free translation<br />
from the original): "Cera: so it is called a tree that
150 Flora Neotropica<br />
grows in the mountains of San Cristobal, from which dark green above, pale green below, semi-coriaceous;<br />
is extracted a hard and valuable wood, of the same drying brown above, olive-green below, semi-coriaintense<br />
yellow as wax [cera]" and "Hilacho: a name ceous; glabrous above and below; primary and secgiven<br />
to a tree of the Sierra Maestra and of the family ondary veins glabrous, secondary veins 8-9 each side;<br />
<strong>Rubiaceae</strong> [C. cubensis]."<br />
tertiary veins starting subparallel and reticulate in the<br />
center; petioles 10-25 mm long, 1.5-2 mm thick,<br />
Of the three species of Chimarrhis endemic to the subterete to adaxially concave, minutely winged, gla-<br />
Caribbean Islands, this is distinct in having smaller brous; tuft-pit domatia often present, with a tuft of<br />
leaves, which are yellow erect-pubescent below. The densely yellowish-sericeous hairs obscuring the pit<br />
petioles and central veins of this species are minutely just below surface, randomly present at axils of secwinged,<br />
forming "pocket-domatia" in the axils of sec- ondary and tertiary veins (Fig. 58C). Inflorescences<br />
ondary veins, never forming tufted domatia. Because 2(-3) per node, corymbose with opposite decussate<br />
of this I treat this taxon as separate, as did Steyermark branches, tertiary branchlets opposite, densely cymose<br />
(1965).<br />
at distal end of each branchlet; (10-)12-22<br />
Grisebach (1862) identified Wright 1262 and 1622<br />
(both from Monte Verde, Cuba) as C. cymosa Jacq.;<br />
the former collection is here selected as the lectotype<br />
of C. cubensis. Grisebach (1862) reported the flowers<br />
of these specimens to be "dioecious-dimorphic,<br />
either with exserted stamens and included stigma, or<br />
exserted style and included anthers." Barely reproducing<br />
Wright's specimen labels, Urban (1899) reported<br />
the flowers of this species as "floribus heterostylis (ex<br />
Wr.)." What were reported as dioecious-dimorphic<br />
(Grisebach, 1862) or heterostylous (Urban, 1899)<br />
flowers are instead two successive stages of anthesis.<br />
4. Chimarrhis jamaicensis (Urban) Steyermark,<br />
Mem. New York Bot. Gard. 12: 184. 1965.<br />
Chimarrhis cymosa Jacquin var.jamaicensis (Urban)<br />
Standley, Field Mus. Nat. Hist., Bot. Ser. 11:<br />
192. 1936. Chimarrhis cymosa Jacquin subsp.<br />
jamaicensis Urban, Symb. Ant. 1: 411. 1899.<br />
Type. Jamaica. Saint James: Mount James, 330<br />
m, 25 Jul 1895 (fr), Harris 5810 (lectotype G, here<br />
selected). Figs. 17F, 58C, 63, 64A-F<br />
Trees to 18 m tall, single-stemmed trees profusely<br />
branching, with small buttresses; bark pale brown.<br />
Leafy branchlets 4-7 mm thick, woody (not succulent),<br />
terete (not costate), pale brown, glabrous; older<br />
branches grayish to pale brown; lenticels not found.<br />
Stipules free at base, contorted in bud, narrowly triangular,<br />
with many parallel veins departing from the<br />
base, glabrous outside and inside, with sparse basal<br />
colleters inside, 20-25 x 8-9 mm, reddish brown,<br />
readily caducous, leaving a pale brown scar encircling<br />
the stem, ca. 0.5 mm wide and very evident in older<br />
branches. Leaves (9-)1 1-18.5 x 5.5-8 cm, L/W = 2:1,<br />
mostly obovate, acute-decurrent at base, obtuse and<br />
mucronate at apex, deltoid acumen ca. 5 mm long;<br />
x (6-)9-<br />
11 cm, 1-2 pairs of lateral branches, basal portion of<br />
axis not branched (5-)6.5-11 cm long; rachis<br />
decussately compressed to subterete, reddish brown,<br />
rachis and branches yellow short-puberulent; distal<br />
bracts absent, reduced to a line just below the branching<br />
point, or minute deltoid scales to 1 mm long;<br />
bracteoles subtending flowers absent. Flowers sessile<br />
to short-pedicellate; pedicels to 2 mm long, glabrous<br />
to sparsely minute-puberulent; hypanthium turbinate,<br />
1-1.5 x ca. 1.5 mm, glabrous to sparsely puberulent.<br />
Calyx reduced, truncate or with wavy margin, 0.5-1<br />
x 1.5-2.5 mm, glabrous. Corolla funnelform, deeply<br />
lobed, 4-5(-6) mm long, white; tube short cylindrical,<br />
ca. 1 x 2-2.5 mm, glabrous outside and inside;<br />
lobes 5, 2-3.5 x 1-1.5 mm, oblong-ovate, rounded to<br />
truncate at apex, glabrous outside, white-villous at<br />
medial zone, the remaining basal and distal portions<br />
glabrous inside. Stamens 5, exserted well above the<br />
corolla, equal, attached 1.5-2 mm from the base of<br />
the tube; filaments 4-4.5 mm long, terete, basally flattened,<br />
connate to throat, densely white-villous at base;<br />
anthers narrowly elliptic, rectilinear, 1.5-2 x ca. 0.3 mm,<br />
dorsifixed near the base, reversed at maturity, base<br />
rounded, with a membranaceous truncate ligula at<br />
apex. Pollen exine irregularly reticulate-foveolate.<br />
Style exserted, 4-5 mm long, as long as the corolla,<br />
glabrous, minutely costate (to winged) toward distal<br />
portion; style branches extruding as two lips above<br />
the corolla before anthesis, rounded to widely elliptic,<br />
ca. 0.5 mm wide, stigmatic surface microscopically<br />
(40x) papillose; ovary-galls not found. Capsules<br />
obovoid to short-obovoid, obtuse at base, shallowly<br />
obtuse at apex, 4-5 x 3-3.5 mm, dark brown without<br />
lenticels, glabrous, brown below the disk; disk glabrous,<br />
black; disk loculicidal dehiscence present in old<br />
capsules. Seeds not seen.<br />
Distribution (Fig. 63) and ecology. Rain forests<br />
on limestone or bauxitic soils, on mountain slopes and
Systematic Treatment of the Genera Studied 151<br />
FIG. 64. Chimarrhisjamaicensis (A-E from Proctor 19818, LL; F from Harris 5810, G). A. Habit of inflorescences.<br />
B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Apical portion of style. F. Mature capsule.
152 Flora Neotropica<br />
low hills, 300-1400 m, of Jamaica and Hispaniola<br />
(Haiti). Flowering specimens were collected in March,<br />
June, July, August, and October. The single fruiting<br />
specimen known has been collected in July.<br />
Conservation status. This species is in danger of<br />
extinction, since it is known only from the heavily<br />
disturbed Massif de la Hotte (Haiti) and on the much<br />
decimated mountain slopes of Jamaica. Its habitat is<br />
in urgent need of protection.<br />
Champ Flore, on mtns. at low elev., Nov 1867 (fl-<br />
fr), Hahn 429 (neotype, P, here selected; isoneo-<br />
types, BR, G[5], K[2]).<br />
Figs. 30, 9D, 17G,H, 58D, 63, 65A-F<br />
Macrocnemum longifolium A. Richard, Mem. Fam.<br />
Rubiacees 199. Dec 1830 (reimp. Mem. Soc. Hist.<br />
Nat. Paris. ser. 3, 5: 279. 1834). Chimarrhis corymbosa<br />
A. Richard, orth. var. [of C. cymosa], Mem.<br />
Fam. Rubiace6s 199. Dec 1830. Type. Martinique.<br />
Locality unknown, 1820, A. Plee s.n. (lectotype, P,<br />
here selected).<br />
Specimens examined. JAMAICA. CLARENDON: Along<br />
trail between Sandy River and John Vale Forest Reserve, 700 Trees to 10 m tall, to 30 cm dbh; single-stemmed<br />
m, 12 Oct 1959 (fl), Proctor 19818 (DAV, LL, NY). PORT- trees, with slender trunk; bark pale brown; hardwood<br />
LAND: John Crow Mtns., 1.6 km S of Ecclesdown, 400 m, yellowish. Leafy branchlets 10-15 mm thick, mark-<br />
17 Jun 1959 (fl), Webster et al. 8341 (DAV[2], G, RB), edly costate, herbaceous, semi-succulent, green to pale<br />
Proctor 19742 (HAC, NY); Youthanside, 13 Jul 1908 (fl),<br />
brown, with a wide<br />
Moore 10648 (NY); near Troy, 650 m, Aug 1904 (fr) 8783<br />
pith, glabrous; older branches<br />
(NY), 16 Sep 1906 (fl) Harris 9449 (NY); Troy Valley, 13grayish<br />
to pale brown; lenticels not found. Stipules<br />
18 Sep 1906 (fr), Britton 610 (NY); Silver Hill, 1260 m, 5<br />
free at base, contorted in bud, narrowly triangular,<br />
Aug 1895 (fl-fr), Harris 5828 (NY); Bachelor's Hall, near acuminate, with many parallel veins departing from<br />
Bath, 300 m, 17 Sep 1908 (fr), Harris & Britton 105 76 (NY); base, glabrous outside, with a band of sericeous hairs<br />
Rodney Hall, 350 m, 28 Mar 1961 (fl), Adams 9289 (M). at the margins, glabrous and with sparse basal colleters<br />
SAINT ANN: Chippenham Park, rd. Moneague-Brown's inside, 40-60 x 12-15 mm, reddish brown, readily<br />
Town, 26 Jul 1850 (fl), Lemann 473 (K). SAINT JAMES: Gar- caducous, leaving a pale brown scar encircling the<br />
lands distr., 4-5 km SSW of Flamstead Gardens, 700 m, 7 stem, ca. 0.5 mm wide and very evident in older<br />
Aug 1962 (fl), Proctor 22591 (DAV, LL). WEST MORELAND: branches. Leaves 22-49 x 10.5-20 cm, L/W = 2:1,<br />
Copse Mt. Woods, 1.6 km SW of Rat Trap, 400-450 m, 23<br />
obovate, acute-decurrent at base, obtuse and mucr-<br />
Oct 1960 (fr), Proctor 21467 (NY).<br />
onate at<br />
HAITI. Massif de la Hotte, western group, Jeremie,<br />
apex, deltoid acumen 5-15 mm long; dark<br />
More Pain-de-Sucre, 1400 m, 22 Jul 1928 (fl), Ekman green above, pale green below, thick-foliaceous; dry-<br />
10398 (G, K); Fairfield, 1849 (fl-fr), Wullschaegel 1344bis ing brown above, olive-green below, stiff-charta-<br />
(M[2]).<br />
ceous; glabrous above and below; primary and secondary<br />
veins glabrous, secondary veins (11-)12-15<br />
Local name and use. Jamaica: wild fiddle wood each side; tertiary veins starting subparallel and re-<br />
(Adams 9289). This species has been reported to be a ticulate in the center; petioles 7-12(-20) mm long, (2good<br />
construction wood.<br />
)3.5-5 mm thick, subterete to adaxially concave and<br />
minutely winged, glabrous; tuft-domatia (not pitted)<br />
This species is unique in the genus in having leaf often present, but only at axils of primary with secblades<br />
with very distinct tuft-pit domatia (Fig. 58C) ondary veins, the hairs of the domatia departing from<br />
in the axils of secondary and sometimes tertiary veins. the veins at the axil zone, not from the blade (Fig.<br />
Chimarrhis jamaicensis is similar to C. cubensis in 58D). Inflorescences 2 per node, corymbose with<br />
having narrowly elliptic anthers 1.5-2 mm long, but opposite decussate branches, laxly cymose at distal<br />
differs in having much larger inflorescence, larger leaf end of each branch, tertiary branchlets alternate to<br />
blades with tuft-pit domatia (vs. "pocket-domatia"), subopposite; 15-24 x (7-)11-17 cm, (1-)2 pairs of<br />
and obovoid-oblong capsules (vs. globose to sub- lateral branches, basal portion of axis not branched<br />
globose).<br />
(5-)8-14 cm long; rachis decussately compressed,<br />
brown, rachis and lateral branches glabrous, terminal<br />
branchlets yellow short-puberulent (curled hairs); distal<br />
bracts absent or reduced to deltoid scales up to 2<br />
5. Chimarrhis cymosa Jacquin, Select. stirp. amer. mm long; bracteoles subtending flowers absent. Flowhist.<br />
61. 1763. Chimarrhis cymosa Jacquin subsp. ers sessile to short-pedicellate; pedicels (when<br />
genuina Urban, nom. inval., Symb. Ant. 1: 410. present) to 1.5 mm long, sparsely minute-puberulent;<br />
1899. Chimarrhis cyrrhosa Stend., Nom. ed. 2. ii. hypanthium turbinate, 1-1.5 x 1.5-2 mm, glabrous to<br />
88, in syn., orth. var. [ofC. cymosa]. Type. Martinique. sparsely puberulent. Calyx reduced, truncate or with
Systematic Treatment of the Genera Studied 153<br />
r~~~~<br />
..<br />
" ?<br />
?' . .<br />
??<br />
?? o<br />
,<br />
\<br />
ii<br />
...<br />
?~~~~~~~ B<br />
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-<br />
men. E. Apical portion of style. F. Mature capsule.
154 Flora Neotropica<br />
wavy margin, ca. 0.5 x 2-2.5 mm, glabrous. Corolla<br />
funnelform, deeply lobed, 3.5-5 mm long, white; tube<br />
short-cylindrical, ca. 1 x 1.5-2 mm, glabrous outside<br />
and inside; lobes 5, 3-4 x 1-1.5 mm, narrowly ob-<br />
long, rounded at apex, glabrous outside; densely<br />
white-villous at basal-medial zone, the distal portion<br />
glabrous inside. Stamens 5, exserted well above the<br />
corolla, equal, attached ca. 1 mm from the base of the<br />
tube; filaments 4-6 mm long, terete, basally flattened,<br />
connate to throat, with the basal 2 mm densely white-<br />
villous; anthers elliptic, ca. 1 x 0.3-0.4 mm, dorsifixed<br />
near the base, base rounded, apex acute, microscopi-<br />
cally (40x) mucronate. Pollen exine reticulate<br />
(densely reticulate-foveolate at poles). Style exserted,<br />
3.5-5 mm long, as long as the corolla, glabrous, mi-<br />
nutely costate (to winged) toward distal portion; style<br />
branches extruding as two lips above the corolla be-<br />
fore anthesis, rounded to oblate, ca. 0.3 mm wide,<br />
stigmatic surface microscopically (40x) papillose;<br />
ovary-galls not found. Capsules globose, obtuse at<br />
base, apex shallowly obtuse, 3-4 x 2.5-3 mm, dark<br />
brown without lenticels, glabrous to sparsely short-<br />
puberulent below the disk; disk black, glabrous, ob-<br />
viously exceeding the calyx; disk loculicidal dehis-<br />
cence present in old capsules. Seeds 0.56-0.78 x<br />
0.56-0.67 mm.<br />
Distribution (Fig. 63) and ecology. Rain forests,<br />
cloud forests, and their disturbed edges, on mountain<br />
slopes, 450-650 m, in the islands of the Lesser Antilles<br />
(Antigua, Guadeloupe, Dominica, Martinique, St.<br />
Lucia, and St. Vincent). On one specimen label (Duss<br />
345) it was noted that the species in Guadeloupe flow-<br />
ers from March to July. Flowering specimens were<br />
collected in February, March, and June, A single<br />
specimen (from Martinique) was collected in late<br />
blooming-early fruiting at the end of July. Fruiting<br />
specimens were collected in January, August, and<br />
October.<br />
Representative specimens examined. WINDWARD<br />
ISLANDS. GUADELOUPE: Close to river, 28 Feb 1895 (fl),<br />
Duss 345 (P); Ermitage a Trois Rivieres, 450 m, 29 Sep 1936<br />
(fr), Stehle 3015 (NY, P); 1944-46, Bena 1066 (P); Camp-<br />
Jacob, Trois Rivieres, 1892 (fl), Duss 2548 (NY), 1896 (fl),<br />
3424 (NY); Basse Terre, Col des Mamelles, 600 m, 26 Oct<br />
1990 (fr), Billiet & Jadin 5010 (BR); 1824, Perrottet 283<br />
(G). DOMINICA: St. Paul, ca. 1.6 km NE of Pont Casse on rd.<br />
to Rosalie, 650 m, 23 Jun 1965 (fl), Webster 13478 (DAV);<br />
Morne Colla Anglais, Sylvania, 610-732 m, 10-23 Aug<br />
1938 (fl), Hodge 695 (NY[2]); St. Joseph Parish, 9 km N of<br />
St. Joseph area of owner of Castaways, 15?24'N, 61?26'W,<br />
521 m, 23 Aug 1992 (fl), Lee 49 (NY[2]); 1 km SW of Millet,<br />
on the Mittel River, 20 m, 22 May 1984 (St), Slane 140<br />
(NY); St. David Parish, Imperial Hwy., near Pont Cass6, vic.<br />
of Grete Palmiste, ca. 560 m, 14 Mar 1993 (fr), Hill 24770<br />
(NY); St. David Parish, Bois Diable, 3 km E of Point Casse,<br />
on rd. to Rosalie, 15?22'N, 61 ?18'W, 24 Apr 1992 (fl), Tuxill<br />
et al. 63 (NY); 3 km from Pont Casse on rd. to Rosalie, 1<br />
Aug 1964 (fr), Wilbur et al. 8137 (CAS, LL); 30 Mar 1916<br />
(fl), Fishlock H338-16 (K); St. Joseph, Roche d'Or Estate,<br />
NE of Salisbury, 650 m, 1 Aug 1966 (fl), Stern & Wasshausen<br />
2582 (B). MARTINIQUE: Camp Flore area, 13-18 Mar<br />
1979 (fr), Howard & Howard 18846(NY, US); Borde de Miere,<br />
trail to St. Pierre, 150 m, 13 Jul 1936 (fl), Stehle 987 (NY);<br />
Gros Morne, 480 m, 11 Sep 1937, Stehle 2298 (US); Le Mont<br />
Vert, 21 Jul 1942 (fl), Stehle 5068 (US); Mome Calabasse<br />
area, 3 km NE of More Rouge toward Ajoupa-Bouillon,<br />
500-600 m, 23 Jul 1959 (fl-fr), Webster et al. 9205 (DAV[2]);<br />
1857, Belanger 195 (G), 196 (G); Martinica, Siebers.n. (M);<br />
Morne Rouge, Boullon, Duss 1452 (NY), 1880 (NY), 1883<br />
(NY); Camp-Jacol, Gompagne, Tres-Rivieres, Duss 2548<br />
(NY), 3424 (NY). ST. LUCIA: Matouba, 17-27 Apr 1979 (fl),<br />
Howard & Howard 19487; vic. of Mt. Gimie, 500 m, 2 Aug<br />
1959 (fr), Webster et al. 9403 (DAV); Canaries River approaching<br />
Mome Gimie, 28 Jan 1985 (fr), Howard et al.<br />
19930 (G, NY); St. Jacques, 8 Oct 1888 (fl), Ramage s.n.<br />
(K). ST. VINCENT: Between Three Rivers and Silver Spoon,<br />
1-7 Apr 1950 (fr), Howard 11143 (NY); Smith & Smith 889<br />
(NY[2]), Mar 1890 (fr), 1245 (K); Anderson s.n. (K).<br />
Local names and uses. Guadeloupe: r6solu de<br />
riviere (Duss 345), resolu de montagne, bois des bamsjaunes.<br />
Dominica: bois riviere (Fishlock H338-16),<br />
quinoyer. Martinique: bois de riviere (Hahn 429), bois<br />
riviere r6solu (Stehle 6626), river wood (Howard<br />
18846). St. Vincent: waterwood. On the specimen<br />
Duss 345 (Guadeloupe) it is reported that this species<br />
gives a much sought-after construction wood.<br />
This species is unique in the genus in having very<br />
thick, semi-succulent, strongly costate leafy branch-<br />
lets (Fig. 65A), with large spongy central pith, which<br />
makes the terminal branchlets very brittle (pers. obs.).<br />
The domatia in C. cymosa are commonly a sparse tuft<br />
of hairs attached on the central and secondary veins<br />
(Fig. 58D), and not on the blade itself (where mites<br />
and scale-insects are usually found).<br />
This is the type species of the genus, and is based<br />
on specimens collected by Jacquin in Martinique, in<br />
proximity of a river. No specimens collected by<br />
Jacquin, however, could be found (not at BM, not at<br />
P, not at W, LE?). The neotype should be selected<br />
from among the specimens listed by Urban (1899)<br />
under C. cymosa subsp. genuina (an invalid subspe-<br />
cific name, by which Urban referred to the "genuine"<br />
C. cymosa from the Lesser antilles). The best candi-<br />
date is Hahn 429 because it was collected in<br />
Martinique, fits the original description by Jacquin,<br />
and it has both flowering and fruiting material. The<br />
specimen of Hahn 429 preserved at P is here selected
Systematic Treatment of the Genera Studied 155<br />
as the neotype of C. cymosa and as lectotype of C.<br />
cymosa subsp. genuina until material from LE is examined<br />
or other possible type material is found.<br />
Jacquin (1763), in the original description, erroneously<br />
reported that the capsules of this species have<br />
one seed per locule. Achille Richard (1830) noted this<br />
error and treated Chimarrhis (which he erroneously<br />
reported as "Chimarrhis corymbosa") as synonymous<br />
with Macrocnemum. Richard (1830) established<br />
Macrocnemum longifolium (= C. cymosa) using material<br />
collected in Martinique (not Jacquin's types),<br />
but without accurate description and specimen citation.<br />
One specimen has been found at P annotated as:<br />
"Le Bois de Reviere - (Chimarrhis cymosa) -<br />
Martinique 1820 - A. Plee." Because of its collection<br />
locality, and the possibility that it was studied by A.<br />
Richard, I have selected this specimen as lectotype of<br />
M. longifolium.<br />
Urban (1899), the first worker to recognize the<br />
existence of three taxa on the Caribbean Islands, called<br />
this species C. cymosa subsp. genuina, because<br />
Jacquin's description was based on material collected<br />
in the Lesser Antilles.<br />
6. Chimarrhis glabriflora Ducke, Bol. Tecn. Inst.<br />
Agron. N. 4: 26. 1945. Type. Brazil. Amazonas:<br />
Esperanca, Rio Solim6es, at mouth of Rio Javari,<br />
"non rara in silva non inundabili," 15 Mar 1944 (fl),<br />
Ducke 1618 (holotype, R; isotypes, A, F, NY,<br />
US[2]). Figs. 3P, 4J,K, 9E,F, 58F, 66A-G,<br />
67A-D, 70, 77A<br />
Trees to 35 m (one individual reported 55 m tall,<br />
Bensman 331), to 50 cm dbh, much-branched at<br />
crown, with buttresses to 3 m high, extending several<br />
meters from trunk; bark red-brown, fibrous, soft, longitudinally<br />
fissured, brown-cream striate; inner bark<br />
pink, striate; wood yellow, very hard. Leafy branchlets<br />
decussately compressed, 3-4 mm thick, 9-12 mm<br />
wide', succulent to semi-succulent, subterete to<br />
decussately compressed, glabrous, youngest 2-3<br />
nodes sparsely to densely minute-puberulent; older<br />
branches glabrate, grayish; lenticels not found.<br />
Stipules free at base, contorted in bud, narrowly triangular,<br />
acuminate, sparsely minute-puberulent outside,<br />
glabrous with sparse small colleters at base inside,<br />
(18-)20-53 x 5-11 mm, with many parallel veins<br />
departing from base, reddish green, readily caducous,<br />
leaving a white scar encircling the stem 1-2 mm wide.<br />
Leaves (15-)18-28 x long; dark green and shiny above, pale green-yellowish<br />
below, semi-coriaceous; drying rust-brown above,<br />
olive-green below, stiff-chartaceous; glabrous above<br />
and below; primary and secondary veins glabrous (to<br />
minute-puberulent), prominent below, secondary<br />
veins (10-)12-16 each side; tertiary veins starting subparallel<br />
and reticulate in the middle; petioles (10-)18-<br />
55 mm long, (1-)2.5-4 mm thick, terete to adaxially<br />
flattened, glabrous; domatia present or absent, usually<br />
a tuft of sparse hairs (white or yellow), 0.1-0.3 mm<br />
long, or barbellate at axils, or a few hairs surrounding<br />
a round slightly depressed glabrous area and converging<br />
toward its center (Fig. 57F). Inflorescences<br />
2 per node; corymbose, laxly branched, with opposite<br />
to subopposite decussate branches, cymules fairly<br />
compacted at distal end of each branchlet, forming a<br />
3-lobed flowering plane, (10-)13.5-20<br />
(7.5-)8-12.5 cm, L/W = ca. 2:1,<br />
obovate (to elliptic-obovate), acute-decurrent at base,<br />
obtuse at apex, sometimes with a deltoid acumen 0.5 cm<br />
x (8-)9.5-13 cm;<br />
lateral branches 3-4 pairs, basal portion of axis not<br />
branched (5.5-)8.5-10 cm long; L/A = 2:1 to 2.5:1;<br />
rachis decussately compressed, rachis and branches<br />
glabrous to puberulent sparsely short-pubescent; distal<br />
bracts absent or reduced to deltoid scales up to 2 x<br />
0.7 mm; bracteoles subtending flowers absent. Flowers<br />
sessile to short-pedicellate; pedicels to 7 mm long,<br />
glabrous to sparsely minute-puberulent; hypanthium<br />
obconical, 0.4-0.8(-1) x 0.4-0.8 mm, glabrous (to microscopically<br />
puberulent). Calyx extremely reduced<br />
and short-lobed, 0.4-0.5 x ca. 1 mm, glabrous; lobes<br />
deltoid to shallowly triangular, 0.1-0.5 mm long, often<br />
ciliate. Corolla funnelform, deeply lobed, 2.5-3.5<br />
mm long, white to cream-white; tube short cylindrical,<br />
1.5-1.8 x ca. 1 mm, glabrous outside and inside;<br />
lobes 5, 1.5-2.6 x 0.7-1.2 mm, oblong, rounded at<br />
apex, glabrous outside, with a tuft of white shaggy appressed-pilose<br />
hairs at medio-basal zone, with distal zone<br />
glabrous inside. Stamens 5, exserted, equal, attached<br />
1.5-1.8 mm from the base of the tube; filaments 2.6-<br />
3.1 mm long, terete, basally flattened, connate to<br />
throat, with a basal tuft of white-pilose hairs for half<br />
the filament length, the hairs decreasing in length toward<br />
medial zone; anthers yellow, elliptic, 0.5-0.6 x<br />
ca. 0.4 mm, dorsifixed at medial zone, base rounded,<br />
apex shortly apiculate (rounded to truncate). Pollen<br />
exine foveolate-reticulate. Style exserted, 2.6-3.5 mm<br />
long, as long as the corolla, terete to square, not costate;<br />
style branches extruding as two lips above corolla<br />
before anthesis, rounded, 0.3-0.4 mm wide, stigmatic<br />
surface microscopically (40x) papillose; ovary-galls<br />
not found. Capsules obovoid (to subglobose), with<br />
very thick pedicels, acute at base, shallowly hemispherical<br />
at apex, 1.5-2.5 x ca. 1.5 mm, rust-brown,<br />
without lenticels; disk black, glabrous, not exceeding<br />
the calyx; disk loculicidal dehiscence present in old<br />
capsules. Seeds 0.48-0.68 x 0.33-0.45 mm.
156 Flora Neotropica<br />
-[--------------------I---------------------------------------.....<br />
+rrl _rD<br />
~II ~C? - -c~B~q<br />
'': ' "'!<br />
tJ: ii ..~~~~~~~~~~~~~~~~<br />
FIG. 66. Chimarrhis glabriflora (A-F from Ducke 1618, GH, isotype; G from Delprete & Verduga 6424, TEX). A.<br />
Habit of inflorescences. B. Flower with exserted anthers. C. Close up of glabrous disk. D. Detail of corolla, inside view. E.<br />
Stamen. F. Apical portion of style. G. Mature capsule.<br />
i<br />
~~ ntr~~~~'T<br />
II~~~ ~~~ dame~.
Systematic Treatment of the Genera Studied 157<br />
,'1<br />
4 :..- ^ ,,tli? ;i.... \?<br />
" ~<br />
:'::~? ~ e<br />
' .~<br />
~<br />
i~~~~~~ic?<br />
'' ~<br />
:'I''t"' ~'<br />
ifi'!i,i' i::,"<br />
'~]gii!11.~1"i<br />
:<br />
I<br />
FIG.~~~~~~~~~*<br />
7. Ciari lbiir.A Toyugidvda c.30c b) .Flaeadinlrsecs .Tuko<br />
oldr ndviua (c. 0 m bh. . Dtal f hetrnk.(Pots akn a Jtfn aca iolgialSttin.<br />
FIG. 67. Chimarrhis glabrflora. A. Two young individual (ca. 30cm dbh). B. Foliage and inflorescences C Trunk of<br />
older individual (ca. 50 cm dbh). D. Detail of the trunk. (Photos taken at Jatun Sacha Biological Station.)<br />
Distribution (Fig. 77A) and ecology. Primary and<br />
secondary lowland (non-inundated) to upland rain<br />
forests or respective remnants, usually growing on<br />
well-drained lateritic soils (rarely in clay soil), in proximity<br />
of streams, of Amazonian Colombia, Ecuador,<br />
Peru, and Brazilian v&rzea forests. Flowering specimens<br />
were collected in January, February, March,<br />
April, May, October, and December. Specimens at<br />
intermediate stage between flowering and early fruiting<br />
have been collected in January, February, March,<br />
April, May, and August. Fruiting specimens were<br />
collected in all months of the year except March,<br />
April, and December. The cream-white, fragrant flowers<br />
have been reported to be visited by bees (Ecuador,<br />
Neill et al. 6220).<br />
Reproductive biology. In my several visits to<br />
Jatun Sacha Biological Reserve (Amazonian Ecua-<br />
dor), I observed a very interesting seed dispersal strat-<br />
egy of both C. glabriflora and C. hookeri, which co-
158 Flora Neotropica<br />
exist in this lowland forest. A mature individual of C.<br />
glabrifora (as most other species) during the flowering<br />
period(2-3 weeks) produces thousands of inflorescences,<br />
and their minute fragrant flowers are visited and<br />
most probably pollinated by bees. Approximately one<br />
month after fertilization the infructescences reach<br />
maturity and some of the capsules open, releasing their<br />
minute seeds. Most infructescences of the individuals<br />
observed were instead dropped (almost simultaneously)<br />
on the forest floor with many capsules still<br />
closed, forming a dense circle of the same diameter<br />
of the individual's crown. Most of these capsules open<br />
shortly after, releasing their seeds, and in about two<br />
weeks the young seedlings start sprouting in large<br />
circles just below the crown of the individual (sometimes<br />
directly from their capsules). Since the inflorescences<br />
are axillary on the terminal nodes, the<br />
branchlets resume their vegetative growth shortly after<br />
the infructescences were abscissed.<br />
The minute seeds of Chimarrhis have been thought<br />
to be wind-dispersed, which is probably the main<br />
means of dispersal in most species, but at least in this<br />
species (and C. hookeri) the mature infructescences<br />
seem to be a primary unit of dispersal. This dispersal<br />
behavior could also explain why Chimarrhis in the<br />
Amazon basin has been reported to have a tendency<br />
of clumping (see ecology discussion of C. turbinata,<br />
and Boom & Campos, 1991).<br />
N bank of Rio Napo, 200 m, 75?52'W, 00?36'S, 7-20 Jan<br />
1991 (fl), Bensman 331 (MO); Taisha, 500 m, 15 Feb 1962<br />
(fl), Cazalet & Pennington 7789 (K, NY, US); Coca, Palm<br />
Oriente Concession, 9 Sep 1983 (fl), Lescure 2060 (CAY,<br />
QCA); Puerto Misahualli, Reserva Biologica Jatun Sacha,<br />
01?04'S, 77?37'W, 450 m, 20 Aug 1994 (fl-fr), Delprete &<br />
Verduga 6423 (AAU, COL, G, MO, NY[2], QCA, QCNE,<br />
TEX), 6424 (AAU, CAS, COL, DAV, F, G, K, MO, NY[2],<br />
P, QCA, QCNE, TEX[3], UPS, US); Puerto Misahualli,<br />
Reserva Biologica Jatin Sacha, 9 Sep 1996 (fr), Delprete<br />
6526 (NY, QCNE). PASTAZA: Via Auca, 110 km S of Coca,<br />
10 km from Rio Tigiiifio, Sector Cristal, 01 15'S, 76?55'W,<br />
320 m, 7 Jan 1989 (fl), Palacios et al. 3380 (AAU, MO, NY,<br />
QCNE, TEX); Cton. Pastaza, "Ramirez" oil well, 20 km S<br />
of Curaray, 76051'W, 01?32'S, 300 m, 21-28 Feb 1990 (flfr),<br />
Zak & Espinoza 5130 (MO, TEX); Lorocachi, Rio<br />
Curaray, 75?58', 01 38'S, 200 m, 29 May 1980 (fl), Jaramillo<br />
et al. 31361 (AAU, MO).<br />
PERU. AMAZONAS: Rio Cenepa, vic. of Huampami, ca.<br />
5 km E of Chavez Valdivia, 78030'W, 04?30'S, 200-250 m,<br />
10 Aug 1978 (fr), Ancuash 1372 (F, MO, NY). HuANuco:<br />
Prov. Pachitea, Dtto. Honoria, Bosque Nacional Iparia, Rio<br />
Pachitea, 300-400 m, 18 Jan 1967 (fl), Schunke 1527 (F, G,<br />
GH, K, MO, NY, US); Prov. Pachitea, Dtto. Puerto Inca,<br />
Bosque Nacional de Iparia, 400-500 m, 19 Dec 1968 (fl),<br />
Schunke 2901 (F, NY, VEN); Prov. Pachitea, Dtto. Puerto<br />
Inca, 14 km from Rio Pachitea, 74?58'W, 093 I'S, 350 m,<br />
14 Apr 1982 (fl), Smith 1311 (MO[2]); Prov. Pachitea, Dtto.<br />
Puerto Inca, Yuyapichis, Unidad de Modelo y Producci6n<br />
Forestal DANTAS, 09?40'S, 75?02'W, 270 m, 1-15 Oct 1990<br />
(fr) Tello 313 (G), 16-31 May 1991 (fr) Tello 1995 (G).<br />
Representative specimens examined. COLOMBIA. JUNIN: Prov. Jauja, Reserva Forestal Granja, Satipo, 750 m,<br />
AMAZONAS: Mun. Leticia, Parque Nacional Amacayacu, trail 7 May 1963 (fr), Vdsquez 14-CBV(MO); Hda. Genova, 1600<br />
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.<br />
Rudas et al. 3016 (MO). CAQUETA: Venecia, margins of Rio Mazan, Gamitanacocha, Rio Mazan, 100-125 m, 1 Mar 1935<br />
Ortegueza, 400 m, 31 Mar 1940 (fl-fr), Cuatrecasas 8947 (fl), Schunke 331 (A, F, NY, US); Dtto. Mazan, Rio Mazan,<br />
(F, NY, US). PUTUMAYO: Rio Putumayo, Puerto 28<br />
Espina, Apr 1977 (fr), Rimachi 2973 (F, NY); Prov. Alto Ama-<br />
00?10'N, 75?50'W, 23-26 Mar 1953 (fl), Schultes & Cabrera zonas, Shucushuyacu (Rio Huallanga), 75?50'W, 06?02'S,<br />
18929 (U, US).<br />
250 m, 12 Sep 1981 (fr), Vdsquez & Jaramillo 2431 (MO,<br />
ECUADOR. EL ORO: Lim6n-Playa, near Rio Dumari, NY, TEX); Prov. Maynas, Maniti, Recreo, 115 m, 72?50'W,<br />
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.<br />
(GUAY, QCA). MORONA-SANTIAGO: Centro Shuar Yukutais, Maynas, Quistococha, 9 May 1977 (fr), Revilla & Froehner<br />
W of Pedro Kunkumas house, 02?30'S, 78?08'W, 900 m, 31 2449 (F); Prov. Maynas, Quebrada Yanomomo, Rio<br />
Mar 1989 (fl), Bennett & Gomez-Andrade 3638, (QCNE); Amazonas above mouth or Rio Napo, 130 m, 15 Nov 1979<br />
Centro Shuar Yukutais, 11 Mar 1990 (fl), Bennett et al. 4028 (fr), Gentry & Jaramillo 28093 (MO). PASCO: Rio Palcazu<br />
(QCNE); Bomboiza, vic. of Salesian Mission in Shuar camp, Valley, second demonstration strip, 09?50'-10?45'S, 68?00'-<br />
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<br />
Arguello 451 (K, NY, QCA, QCNE, TEX); Santiago, Rio (MO, NY, TEX); Rio Palcazu Valley, Samuel Ponce's pas-<br />
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<br />
Little et al. 763 (MO). NAPO: Estaci6n Esperimental de (NY, TEX), and 20 Jun 1986 (fr), 2632 (MO); Prov.<br />
INIAP, San Carlos, 6 km E of Los Sachas, 250 m, 4 Apr 1985 Oxapampa, Rio Palcazu Valley, Rio San Jose, 10?09'S,<br />
(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],<br />
Jose Payamino, 40 km W of Coca, 00?30'S, 77?20'W, 300- NY). SAN MARTIN: Prov. Mariscal Caceres, Dtto. Tocache<br />
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<br />
4.8 km E of Rio Conejo on rd. to Lago Agrio, 340 m, 1 Apr 10653 (F, K, MO, NY).<br />
1972 (fl), MacBride & Dwyer 1408 (MO, QCA); La Joya BRAZIL. ACRE: Cruzeiro do Sul, Rio Jurua & Rio Moa,<br />
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.<br />
Gudino 1707 (NY); Cton. Aguarico, Chiro Isla community, 12462 (GH, MO, NY[2], U[2], US).
Systematic Treatment of the Genera Studied 159<br />
Local names and uses. Ecuador: mincha caspi<br />
(Quichua, = wick tree) (Irvine 634, Napo, Irvine 585),<br />
mecha caspi (Quichua, Bensman 331), jatun mincha<br />
caspi (= large wick tree) (Napo, Irvine 204). Peru:<br />
itauba (Tello 1995), itauba amarilla (Tello 313), cascarilla<br />
masha (Quichua, Schunke 1527, 2901; 10653),<br />
palo palillo (Junin, C. B. Vasquez 14-CBV), pablo<br />
manchana, pampa remocaspi (Quichua, =<br />
that detached easily in irregular pieces. The tree architecture<br />
of this species follows the Aubreville Model<br />
(cf. Halle et al., 1978).<br />
7. Chimarrhis hookeri K. Schumann in Martius, Fl.<br />
Bras. 6(6): 259. 1889. Type. Peru. San Martin:<br />
paddlewood), and yero prueba (the last three names Tarapoto, in tall forest, tree ca. 25 m tall, Jan 1857<br />
reported by Duke & Vasquez, 1994).<br />
(fl), Spruce 4930 (B*; lectotype, K, selected by<br />
This species is reported by many foresters and in- Delprete, 1999b; isolectotypes, BR, C-n.v., F[2],<br />
digenous tribes to have very hard wood G, GH, K, NY, P;<br />
(with copiphoto-C<br />
at F, GH, MO, NY,<br />
ous watery orange juice when cut), providing excel- VEN). Spruce annotated these specimens as "Conlent<br />
timber for construction of houses (Peru, Schunke<br />
daminea glabrata DC. [unpubl.] aff., a curious<br />
331). Duke and Vasquez (1994) reported that the wood thing!" Figs. 9G, 18C, 58G, 68A-H, 69A-D, 70, 77B<br />
of this species is used for rural construction, firewood, Chimarrhis williamsii Standley, Publ. Field Columbian<br />
and living fences. Bensman 331 bears the following Mus., Bot. Ser. 8: 162. 1930. Type. Peru. Loreto:<br />
notes: "tree 55 mm high and over 50 cm at DBH, with Rio Nanay, 23 May 1929 (fl), LI. Williams 409 (holarge<br />
buttresses, yellowish wood with thin bark, used lotype, F, N. 601986; frag-F at G).<br />
as construction wood."<br />
Trees to 35 m tall, 50 cm or more dbh (exception-<br />
Chimarrhis glabriflora is sympatric with C. ally 50 m tall, 1 m dbh, Baker et al. 6000), much<br />
hookeri (Fig. 77A-B), with which it is commonly branched at crown, with tall buttresses; bark fissured,<br />
confused, but from which it differs in having larger white-grayish, peeling off in long fibrous strips exand<br />
rather fleshy leaves, which are deep green and posing the reddish bark underneath; wood yellow,<br />
shiny above, semi-succulent leafy branchlets (Figs. very hard. Leafy branchlets 3-4(-5) mm thick,<br />
66A, 68B, 70), a larger inflorescence, glabrous cap- woody (not succulent), subterete to decussately comsules,<br />
and glabrous disks. The leaf veins in C. pressed, youngest 2-3 nodes densely appressed-canesglabriflora<br />
are always glabrous and the domatia are cent; older branches glabrate, terete, grayish to pale<br />
rather variable in either being represented by a small brown; lenticels not found. Stipules free at base, contuft<br />
of hairs, or a peculiar area of hairs attached on torted in bud, narrowly triangular, acuminate, densely<br />
the veins' axil and perpendicular to them (resembling canescent (rarely puberulent or rarely glabrous) outa<br />
small brush), or completely absent.<br />
side, glabrous with sparse small colleters at base in-<br />
When Ducke (1945) described this species, he side, 12-18(-30)<br />
added the following notes: "This well defined species<br />
is easily distinguished from the two remaining species<br />
of Brazilian Amazonia (Ch. turbinata and Ch.<br />
barbata), by various botanical characters, chiefly by<br />
the very small calyx and by the entirely glabrous corolla.<br />
It is a rather frequent tree of the upland forest<br />
of the western limit of Brazil, and will certainly be<br />
observed in the neighboring parts of Peru and Colombia.<br />
The fragrant flowers appear at the climax of the<br />
rainy season. The tree yield good fire wood."<br />
Strangely Ducke (1945) did not compare this species<br />
to its close relative C. hookeri, from western Amazon<br />
(and with which it is sympatric), which is well<br />
described in Flora brasiliensis (Schumann, 1889).<br />
The largest individuals of Chimarrhis that I have<br />
personally seen belong to this species (at Jatun Sacha<br />
Biological Station, Ecuador). One individual had large<br />
buttresses (4-5 m long), the trunk about 1.5 m in diameter<br />
at about 4 m above ground (above the buttresses)<br />
and ca. 45 m tall, and rust-brown fibrous bark<br />
x 3-5 mm, gray-canescent, with<br />
many parallel veins departing from base, readily caducous,<br />
leaving a white scar encircling the stem, 0.5-<br />
1 mm wide. Leaves 6-16(-20) x 3-6.5(-9) cm, L/W =<br />
2:1, elliptic to elliptic-obovate, acute at base, obtuse<br />
at apex, sometimes with a deltoid acumen 0.5 cm long;<br />
dark green above, pale green below, semi-coriaceous;<br />
drying dark brown above, olive-green below, stiff-chartaceous;<br />
juvenile leaves with blade, primary and secondary<br />
veins sparsely appressed pubescent above,<br />
mature leaves glabrous above, appressed pubescent<br />
below; primary and secondary veins ascending-pubescent<br />
to canescent (rarely glabrous), secondary veins<br />
8-9 each side; tertiary veins starting subparallel and<br />
reticulate in the middle; petioles 8-20 mm long, 1-<br />
1.5 mm thick, terete to adaxially flattened, glabrous<br />
to puberulent to canescent; domatia represented by a<br />
few hairs surrounding a round slightly depressed glabrous<br />
area and converging toward its center (Fig.<br />
57G), hairs 0.4-0.7 mm long. Inflorescences 2 (rarely<br />
4) per node; corymbose with opposite to subopposite
160 Flora Neotropica<br />
_ ..<br />
DI II -<br />
FIG. 68. Chimarrhis hookeri (B-G from Spruce 4930, K, lectotype; H from Delprete & Verduga 6421, TEX). A. Archi-<br />
tecture and dimensions of mature individual (from pers. obs.). B. Habit of inflorescence. C. Flower with exserted anthers.<br />
D. Close up of pubescent disk. E. Detail of corolla, inside view. F. Stamen. G. Apical portion of style. H. Mature capsule.
Systematic Treatment of the Genera Studied 161<br />
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,<br />
i<br />
:.. ??;::'":4i.:<br />
FIG. 69. Chimarrhis hookeri. A. Tree climber reaching the c:rown of..-- a:youn. indiidual. Inf'l-~e.c:en.es.w,'h,"<br />
;., a B.<br />
- t.re<br />
[i. I!<br />
-,<br />
,~!~<br />
fruits~~~' (se figrfrsae.C. Trun of anodenivda (ca~~..",:', bh.: D.Dt.ail of.the'trunk.-? (Pooae tJti<br />
c4<br />
:<br />
40 4<br />
i ~ .'?, ~~~~~ ~ "!~'. ~ ~~~~~~~~~~~~~;<br />
: ,~~ '; :i<br />
1 "?'? _ ~ ~ S ~ i ~ ~ l v.,,.' ~ : i, :r ':<br />
; \91. , ;<br />
r'~' ''<br />
?<br />
| [f; \ l<br />
F'~~~ ~~~ ~ ~ ~~~~~~~~~~~~~~~~~~~~~~".<br />
FIG. 69. Chimarrhis hooleri. A. Tree climber reaching the crown of a young individual. Infiorescences B.<br />
with mature<br />
Sacha Biological Station, Ecuador).
162 Flora Neotropica<br />
:.... .,. e . .. ....<br />
H? ?<br />
_s __?<br />
cal Station, Ecuador).<br />
FIG. 70. Terminal leafy branchlets of Chimarrhis glabriflora (left) and C hookeri (right). Note larger leaves and semi-<br />
cal Station, Ecuador).<br />
decussate branches, cymules fairly compacted at dis-<br />
tal end of each branchlet, forming a continuous flow-<br />
ering plane, 8-10(-12) x (4-)5.5-7.5 cm, lateral<br />
branches 2(-3) pairs, basal portion of axis not<br />
branched 4-5.5(-8) cm long; L/A = 2.5:1 to 2.7:1<br />
(rarely 3:1); rachis decussately compressed, rachis and<br />
branches sparsely yellow-puberulent; distal bracts<br />
reduced to scales or linear, 1-3 x 0.3-0.5 mm;<br />
bracteoles subtending flowers absent. Flowers sessile<br />
to short-pedicellate; pedicels to 1 mm long, glabrous<br />
to sparsely pubescent (to short-villous); hypanthium<br />
obovate, 0.9-1.2 x 0.8-1.1 mm, glabrous, reddish<br />
brown. Calyx extremely reduced, truncate to short-<br />
lobed, 0.3-0.5 x 1-1.5 mm, glabrous; lobes shallowly<br />
triangular, connected by a membrane, 0.1-0.6 mm<br />
long, often ciliate. Corolla funnelform, deeply lobed,<br />
2.5-3.5 mm long, cream-white to yellowish white<br />
(turning pale yellow at maturity); tube short cylindri-<br />
cal, 1-1.5 x 0.7-1 mm, glabrous outside and inside;<br />
lobes 5, 1.5-2 x 0.5-0.7 mm, oblong, rounded at apex,<br />
glabrous outside, with a tuft of white thin-sericeous<br />
hairs at basal zone with medial and distal zones gla-<br />
brous inside. Stamens 5, exserted well above the co-<br />
rolla, equal, attached 1-1.5 mm from the base of the<br />
tube; filaments 2-2.5 mm long, terete, basally flat-<br />
tened, connate to throat, with a dense tuft of white-<br />
sericeous hairs for half the length at base; anthers el-<br />
liptic, 0.5-0.7 x 0.4-0.5 mm, dorsifixed at medial<br />
zone, base rounded, apex rounded, not mucronate.<br />
Pollen exine foveolate-reticulate. Style exserted, 2.5-<br />
3.5 mm long, as long as the corolla, terete to square<br />
(not costate), often with few sparse hairs on distal<br />
portion; style branches extruding as two lips above<br />
corolla before anthesis, rounded, ca. 0.5 mm wide,<br />
stigmatic surface microscopically (40x) papillose;<br />
ovary-galls not found. Capsules subglobose to obo-<br />
void, obtuse at base, shallowly hemispherical at apex,<br />
2-2.5 x 1.5-2 mm, capsule dark brown, without len-<br />
ticels; disk black, sparsely to densely pubescent, not<br />
exceeding the calyx; disk loculicidal dehiscence<br />
present in old capsules. Seeds 0.5-0.63 x 0.35-0.5 mm.<br />
Distribution (Fig. 77B) and ecology. Primary and<br />
secondary rain forests, and respective remnants, in<br />
riverside forest, in alluvial, lateritic, sandy to clay<br />
soils, from low to high fertility, 150-800 m, of Ecua-
Systematic Treatment of the Genera Studied 163<br />
dorian and Peruvian Amazon. Flowering specimens (fl), Gutierrez 32 (MO); Prov. Pachitea, Dtto. Honoria, rd.<br />
were collected in January, February, May, June, Septo<br />
Turavista, 24 Jan 1963 (fl), Lao Magin 44 (F, K, NY),<br />
tember, and December. Specimens at intermediate 31 May 1963 (fr), 51 (F, NY, US). LORETO: Upper Rio<br />
stage between flowering and early fruiting have been Marafion, mouth of Rio Santiago, 160 m, 3 Dec 1924 (fl),<br />
Tessman 4668<br />
collected in April and<br />
(B, F, NY); Prov. Maynas, Pto. Allianza,<br />
August. Fruiting specimens<br />
72?55'W, 04?08'S, 160 m, 29 May 1981 (fr), Vdsquez &<br />
were collected in February, April, May (many), June, Criollo 1855 (F, MO, NY, TEX); Prov. Maynas, Yanomono,<br />
September, October, and December.<br />
Explorama Tourist Camp, between Indiana and mouth of Rio<br />
Napo, 130 m, 18 Feb 1981 (fr), Gentry et al. 31384 (F, G,<br />
Representative specimens examined. ECUADOR. MO, NY); Rio Nanay, 23 May 1929 (fl), Williams 369 (F,<br />
ESMERALDA: Canton Esmeralda, San Lorenzo-Lita, trail to K, US); Bosque Nacional Alexander Von Humboldt, km 91<br />
Punta, 00?49'N, 78?26'W, 200 m, Macias & Burbano 7788 on rd. Pucallpa-Lima, 300-500 m, 12 May 1975 (fr), Hart-<br />
(QCNE). NAPO: Estaci6n Experimental INIAP, San Carlos, shorn 1687 (F, MO[3]). MADRE DE Dios: Tambopata,<br />
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.<br />
al. 240 (AAU, F, MO, NY, QCNE, TEX, US); Reserva 45661 (MO, NY), 46255 (MO). UCAYALI: Prov. Coronel<br />
Biol6gica Jatun Sacha, 8 km downstream from Puerto Misa- Portillo, Pucallpa-Tingo Maria Hwy., 75?00'W, 08?41'S,<br />
hualli on Rio Napo, 01?04'S, 77?36'W, 450 m, Palacios et 250m, 11 Feb 1981 (fr), Gentryetal. 31219(F, MO); Prov.<br />
al. 365 (AAU, F, MO, NY, QCNE, TEX); Rio Aguarico, near Coronel Portello, Campo Verde, km 78 C.F. Basadre, 250the<br />
confluence of Rio Pavayacu, Sep 1981 (fr), Bravo & 270 m, 13 Mar 1989 (st), Chavez 265 (MO).<br />
G6mez 306 (QCA); Amazon primary forest, 77?28'W,<br />
01?20'S, 520 m, 19 May 1976 (fr), Oldeman & Arevalo 16 Local names and uses. Ecuador: intacchi<br />
(QCA); Parque Nacional Yasuni, road-oleoduct Maxus in (Quichua, Napo, Baker et al. 6000). Peru: itauba (Tello<br />
construction, km 46 to Capiron Well, primary forest, 862), itauba amarilla (Tello s.n.), itahuba (Pachitea,<br />
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<br />
(QCNE); 5 km SE of Las Sachas, 300 m, 13 Apr 1985 (fr), (Tessman 4668), huacapu masha (Hartshorn 1687),<br />
Baker et al. 6000 (MO, NY); Parque Nacional Yasuni, La<br />
quillo-bordon, purma-caspi (Quichua, = yellow-<br />
Joya de Los Sachas, rd. Maxus oleoduct, km 15, 00?25'S,<br />
wood),<br />
76?37'W, 250 m, 27 Feb 1993 (fl), Gudino &<br />
chollachaqui<br />
Grefa 2336<br />
caspi blanco (Ucayali, Chacvez<br />
(TEX); Canton Archidona, SE of Volcan Sumaco, rd. 265), sacha jagua (Leoncio Prado, Gutierrez 32),<br />
Hollin-Loreto, km 65, 00?44'S, 77028'W, 600 m, 26 Jun papelillo caspi (Huanuco, Lao Magin 44 and 51),<br />
1989 (fr), Alvarado 292 (AAU), Hurtado 2365; Cton. Tena,<br />
aromuhe (Maynas, Paitan 3b), pampa remo caspi<br />
8 km E of Puerto Misahualli, Reserva Biologica Jatun Sacha, (Maynas, Flores 61 [Quichua<br />
01?04'S, 77?36'W, 400 m, 20 Aug 1994 (fl-fr), Delprete &<br />
Verduga 6421 (AAU, F, G, K, MO, P, NY[2], QCA, QCNE,<br />
TEX, UPS[2], US), 6422 (COL, NY, QCA, TEX); Puerto<br />
Misahualli, Reserva Biologica Jatun Sacha, 9 Sep 1996 (fr),<br />
Delprete 6527 (NY, QCNE), 6528 (NY, QCNE). PASTAZA:<br />
Rio Curaray, 75040'W, 01?36'S, 200 m, 5 Jun 1980 (fl),<br />
Brandbyge & Asanza 31615 (AAU, MO, QCA, US);<br />
Lorocachi, 2 km S of village, left margin of Rio Curaray,<br />
75?58'W, 01?38'S, 200 m, 29 May 1980 (fl), Jaramillo et<br />
al. 31631 (AAU, QCA, US); rd. PETRO-CANADA, Via<br />
Auca, 115 km S of Coca, 4 km S of Rio Tigiifio, 01?15'S,<br />
76?55'W, 320 m, 22-28 Feb 1989 (fl), Zak 3979 (G, GB,<br />
MO, QCNE, TEX); Montalvo, 1 km NE of military camp,<br />
76058'W, 02005'S, 250 m, 17-19 May 1979 (fl), Lojtnant &<br />
Molau 13513 (AAU). SUCUMBIOS: Cant6n Lago Agrio, Parr.<br />
Tarapoa, 1 km from Mariann 3 oil well toward Reserva<br />
Faunistica Cuyabeno, 76022'W, 00?08'S, 240 m, 27 Apr 1990<br />
(fl-fr), Cer6n & Ayala 9656 (MO, TEX, QCNE).<br />
PERU. AMAZONAS: Rio Santiago Valley, 77?40'W,<br />
03?50'S, Quebrada Caterpiza, 3 Dec 1979 (fl), Tunqui 205<br />
(MO); Rio Cenepa, Quebrada Sasa, 30 May 1973 (fr),<br />
Ancuash 504 (F, MO, NY). HUANUco: Prov. Puerto Inca,<br />
Dtto. Yuyapichis, Unidad de Modelo y Producci6n Forestal<br />
DANTAS, 09?40'S, 75002'W, 270 m, 1-15 Dec 1990 (fr)<br />
Tello 862 (G); Prov. Huamalies, Cordillera Azul, Fundo<br />
Sinchono, km 209, 1600 m, 12 Oct 1944 (fr), Hodge 9 (F);<br />
Prov. Leoncio Prado, Hda. Laconyope, 870 m, 17 Jan 1962<br />
= paddlewood]),<br />
mukugd (Huambisa, Amazonas, Tunqui 945), pablo<br />
manchana, yemo prueba, and palo perro (the last three<br />
nemes reported by Duke & Vasquez, 1994).<br />
The label of Baker et al. 6000 reports: "Tree 50 m<br />
tall, I m dbh, with a crown roundish and flattened on<br />
top; its wood is very hard and the indigenous people<br />
use it to build their houses." Additional uses are reported<br />
as: "Its bark is used to make ropes, and its wood<br />
is of potential value for production of particle boards"<br />
(Lao Magin 44 and 51), and "used for construction of<br />
boats" (M.A. V. 42).<br />
This Amazonian species is recognizable by its<br />
small leaves, thin and readily woody leafy branchlets<br />
(Fig. 70), small capsules, and pubescent disks; leaf<br />
blades sparsely puberulent to glabrous below; leaf<br />
veins ascending-pubescent to canescent (rarely glabrous);<br />
and domatia often present as a tuft of pilose<br />
hairs converging toward the center of the axils. When<br />
the blades and the veins are glabrous the domatia are<br />
absent (Fig. 58G). Mature individuals of C. hookeri<br />
have a tree architecture that follows the Aubreville<br />
Model (cf. Halle et al., 1978).<br />
Chimarrhis hookeri is often confused with C.<br />
glabriflora (with which it is sympatric), the former
164 Flora Neotropica<br />
differing from the latter in having smaller, thinner<br />
leaves (Figs. 66A, 67B, 70), with 8-9 pairs of secondary<br />
veins (vs. 12-16 pairs of veins), slender woody<br />
(vs. semi-succulent and thick; see Fig. 70) branchlets,<br />
smaller, densely appressed-pubescent to canescent<br />
(vs. sparsely puberulent) stipules, and a pubescent (vs.<br />
glabrous) capsule disk.<br />
Chimarrhis williamsii was established by Standley<br />
(1930b) from material collected in the proximity of<br />
Rio Nanay (Prov. Loreto, Peru). He distinguished it<br />
from C. hookeri by erroneously stating that the latter<br />
has glabrous stipules. He also admitted that he did not<br />
see any material of the latter species. The disk and the<br />
stipules of C. williamsii (Ll. Williams 396 and 409)<br />
are sparsely pubescent, although the rest of the vegetative<br />
and reproductive characters closely match<br />
those of C. hookeri. The two taxa are here treated as<br />
synonymous.<br />
8. Chimarrhis microcarpa Standley, Bull. Torrey<br />
Bot. Club 53: 471. 1926, non Chimarrhis cymosa<br />
Jacquin subsp. microcarpa Urban, Symb. Ant. 1:<br />
411. 1899 (= C. cubensis Steyermark), non<br />
Chimarrhis cymosa Jacquin var. microcarpa (Urban)<br />
Standley, Field Mus. Nat. Hist., Bot. Ser. 11:<br />
192. 1936 (= C. cubensis Steyermark), Chimarrhis<br />
microcarpa var. microcarpa, Mem. New York<br />
Bot. Gard. 12(3): 183. 1965. Type. Trinidad.<br />
Maraval, 1904 (fr), Dannouse 6946 (holotype,<br />
NY). Figs. 71A-G, 73<br />
Chimarrhis longistipulata Bremekamp, Recueil Trav.<br />
Bot. Neerl. 31: 260. 1934. Type. Surinam. Kabalebo<br />
River, Avanero Falls, 3 Sep 1920 (fr), Pulle 379 (holotype,<br />
U, photo-U at NY, VEN; isotypes, K, US).<br />
Trees to 40 m tall, 100 cm or more dbh, canopy<br />
trees with lobed bole, small spreading crown, with<br />
buttresses to 3.35 m high; bark fissured, grayish<br />
brown; wood yellow, very hard. Leafy branchlets<br />
2.5-5(-6) mm thick, subterete to decussately compressed,<br />
green to brown, glabrous; older branches terete,<br />
grayish to pale brown; lenticels not found.<br />
Stipules free at base, contorted in bud, narrowly triangular,<br />
acuminate, sparsely appressed-puberulent<br />
(rarely glabrous) outside, glabrous with basal colleters<br />
inside, (20-)25-35 x (4-)5-7 mm, reddish brown,<br />
membranous, with many parallel veins departing from<br />
base, readily caducous, leaving a scar encircling the<br />
stem ca. 1 mm wide. Leaves (10-)16-22 x the deltoid acumen 0.5-1.8 cm long; dark green above,<br />
pale green below, thick-foliaceous; drying brown to<br />
olive-green, stiff-chartaceous; glabrous above, glabrous<br />
to sparsely appressed-puberulent below; primary<br />
and secondary veins glabrous to appressed-puberulent,<br />
secondary veins 8-11 each side; tertiary<br />
veins subparallel, laterally jointed in the center; petioles<br />
(13-)18-31 mm long, 1-1.5(-2) mm thick, adaxially<br />
flattened to concave, glabrous; tuft-domatia<br />
always present, tuft of hairs departing from the veins<br />
and converging toward the center of the axil (Fig.<br />
58E), only at the axils of primary with secondary<br />
veins, hairs 0.7-1 mm long, white to golden. Inflorescences<br />
2 per node, corymbose with opposite to<br />
subopposite decussate branches, cymose, very compacted<br />
at distal end of each branchlet, forming a continuous<br />
flowering plane, (8-)10-12(-15)<br />
(5-)6.5-<br />
9.5 cm, L/W = ca. 2:1, elliptic to obovate, acute-decurrent<br />
at base, acute to obtuse and mucronate at apex,<br />
x (4-)5-7<br />
(-9) cm, lateral branches 2(-3) pairs, basal portion of<br />
axis not branched (5-)6.5-10.5 cm long; L/A = ca.<br />
3.1; rachis terete to decussately compressed, rachis<br />
and branches short puberulent; distal bracts usually<br />
absent or narrowly triangular, to 4 x 3 mm; bracteoles<br />
subtending flowers absent. Flowers sessile to shortpedicellate;<br />
pedicels to 1 mm long, glabrous to sparsely<br />
puberulent; hypanthium obconical, 0.8-1 x 0.8-1.2<br />
mm, glabrous, reddish brown. Calyx extremely reduced,<br />
truncate to faintly undulate margin, 0.3-0.6 x<br />
1-1.4 mm, glabrous. Corolla funnelform, deeply<br />
lobed, 4-5 mm long, cream-white to greenish white<br />
(turning pale yellow at maturity); tube short cylindrical,<br />
0.9-1.5 x 0.6-1.1 mm, glabrous outside and inside;<br />
lobes 5, 1.8-3.2 x 0.4-0.6 mm, narrowly oblong,<br />
acute (rarely rounded) at apex, glabrous outside,<br />
sparsely white-villous at basal zone, with medial and<br />
distal zones glabrous inside. Stamens 5, exserted well<br />
above the corolla, equal, attached 1-1.7 mm from the<br />
base of the tube; filaments 3.5-5.5 mm long, terete,<br />
basally flattened, connate to throat, with a tuft of<br />
densely white-pilose hairs at base; anthers elliptic,<br />
0.6-0.7 x 0.3-0.4 mm, dorsifixed at medial zone, reversed<br />
at maturity, base rounded, apex shortly apiculate,<br />
truncate to rounded, dehiscing by longitudinal<br />
slit. Pollen exine foveolate-reticulate. Style exserted,<br />
3-5 mm long, as long as the corolla, glabrous, minutely<br />
costate to minutely winged; style branches<br />
extruding as two lips above corolla before anthesis,<br />
rounded to oblate, ca. 0.3 mm long, stigmatic surface<br />
microscopically (40x) papillose; ovary-galls not<br />
found. Capsules subglobose, obtuse at base, rounded<br />
at apex, 1.9-2.5 x 1.3-1.7 mm, dark brown without<br />
lenticels, glabrous throughout; disk black, exceeding<br />
the calyx; disk loculicidal dehiscence present in old<br />
capsules. Seeds 0.68-73 x 0.53-0.7 mm.
Systematic Treatment of the Genera Studied 165<br />
0<br />
FIG. 71. Chimarrhis microcarpa (B and G from Dannouse 6946, NY, holotype; C-F from Smith 3433, F). A. Architec-<br />
ture and dimensions of mature individual (from pers. obs.). B. Habit of inflorescences. C. Flower with exserted anthers. D.<br />
Detail of corolla, inside view. E. Stamen. F. Apical portion of style. G. Mature capsule.<br />
ic
166 Flora Neotropica<br />
Distribution (Fig. 73) and ecology. Primary and<br />
secondary rain forests, and respective remnants, mixed<br />
forests, and recent regrowth vegetation, on lateritic<br />
soils (often on red clay), usually in proximity of creeks<br />
and streams, on slopes at low elevation, 20-600 m,<br />
of eastern Venezuela, Trinidad, Guyana, French<br />
Guiana, Surinam, and northern Brazil. Flowering<br />
specimens were collected in January, February,<br />
March, April, and October. Fruiting specimens were<br />
collected in January, February, March, April, June,<br />
August, September, and December.<br />
Representative specimens examined. VENEZUELA. Local names and uses. Venezuela: carutillo (Delta<br />
ANZOATEGUI: Between Rio Le6n ofQuebrada Danta and Rio Amacuro, Marcano Berti 241), totumo amarillo (Mar-<br />
Zumbador, NE of Bergantin, 500-600 m, 24 Feb 1945 (fl), cano Berti 1426), amarilla (Anzoategui, Steyermark<br />
Steyermark 61136 (A, F[2], NY, US, VEN). BOLIVAR: Si-<br />
61136), agua fresca (Merida, Bernardi 5943). Guyana:<br />
erra de Lema, source of Rio Chicanan, 80 km SW of El white calabash (McDowell 4472). Steyermark (1974)<br />
Dorado, 60?05'N, 62?W, 500 m, 27 Aug 1961 (fr),<br />
Steyermark 89514 (NY, VEN); Reserva Forestal La<br />
reported that the hard wood of this species is used to<br />
Paragua, make<br />
margins of Rio Asa, Jun 1970 (fr), Blanco 822 (F, NY,<br />
wheel-spokes, beams, and posts.<br />
VEN).<br />
TERRITORIO FEDERAL DELTA AMACURO: Close to border with<br />
Estdo. Bolivar, E of Rio Grande, ENE of El Palmar, 21 Jun This species is unique in having narrow-funnel-<br />
1964 (fr), Marcano Berti 241 (F, K, MO[4], NY, VEN); form corollas 4-5 mm long, small, compactly<br />
Lower Orinoco, May 1896 (fl), Rusby & Squires 126 (A, F, branched inflorescences (L/A = 3:1), slender woody<br />
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<br />
Upata, 61?44'W, 08?14'N, 7 Apr 1967 (fl), Brujin 1646 (F, (Fig. 71B). Chimarrhis microcarpa is a rather com-<br />
M, MO, P, NY, US, VEN), 1647 (F, M, MO, P, NY, US, mon species in the lowland rain forests of northeast-<br />
VEN), 1651 (F, M, MO, P, NY, US, VEN). LARA: Dto. ern South America, reaching dimensions of large<br />
Iribarren, vic. of Laguna Negra, Loma de los Naranjos,<br />
Montafia de Macanillal and Fila de San Esteban, 10-19 km<br />
canopy-trees (40-50 m tall and to 1.5 m diam.; see<br />
S of Rio Claro, 1300-1500 m, 24-25 Mar 1975 Fig. 71A), and often described as trees with lobed<br />
(fl),<br />
Steyermark et al. 111625 (MO, NY, VEN). MONAGAS: boles, large buttresses, and small flattened crowns.<br />
Montafia de Aguacate, along Quebrada de Pajarral,<br />
The tree<br />
tributary<br />
architecture of this species is the Aubreville<br />
of Rio Caripe, NE of Alto de Aguacate, 600-900 m, 19 Apr<br />
Model (Terminalia branding; cf. Halle et al., 1978).<br />
1945 (fr), Steyermark 62185 (A, F, NY, VEN). SUCRE: Its bark is shallowly fissured and grayish brown; when<br />
Manacal, NW of Irapa, 800 m, 10 Dec 1966 (fl), Marcano freshly cut its sapwood is creamy-white to yellowish<br />
Berti et al. 1426 (VEN).<br />
white, and its heartwood pale-brown to dark-brown.<br />
TRINIDAD AND TOBAGO. TRINIDAD: Heights of Its leaves are thick-papery, dark-green and shiny<br />
Aripo, 10-26 Jan 1922 (fl), Broadway 10011 (NY, US); above and pale-green below, and its flowers are sweet-<br />
Blanchisense rd., 11 m post, 31 Dec 1926 (fr), Broadway<br />
6464<br />
fragrant.<br />
(K).<br />
Chimarrhis<br />
GUYANA. Kanuku Mtns., NW slopes, drainage of<br />
microcarpa differs from C. speciosa<br />
Moku-moku Creek, 150-400 m, Mar-Apr 1938 (fl), Smith<br />
in the larger funnelform corolla, slender and woody<br />
3433 (A, B, F, G, IAN, MO, NY, P, U, US); Rupununi, near leafy branchlets, and the much smaller, more compact<br />
Moco-Moco, 03?20'N, 59?35'W, 100 m, 29 Oct 1979, Maas inflorescences. The capsules in C. microcarpa and C.<br />
& Westra 3912 (U, VEN); Barima-Waini Region, Barima speciosa are similar in size and shape, and have gla-<br />
Head, along Barima River, 07?35'N, 60035'W, 91-122 m, brous disks; but in C. microcarpa the pedicels are<br />
15 Apr 1991 (fl), McDowell et al. 4411 (G, U, US); Barima- longer and more slender, while in C. speciosa the<br />
Waini Region, Matthew Ridge to Blue Mtn., 07?26'N, capsules are subsessile (vs. pedunculate).<br />
6011' W, 100-400 m, 28 Apr 1991 (fl), McDowell 4472<br />
Many herbarium specimens of C. microcarpa have<br />
(MO); Lower Essequibo River, Groete Creek, 19 Apr 1943<br />
(fl), Fanshawe F1249 (3985) (F, K[2], NY[2], U); Potaro-<br />
Siparuni region, 4-5 km N of Surama village, confluence of<br />
Burro-Burro and Surama rivers, 04?10'N, 59003'W, 75 m, 1<br />
May 1992 (fl), Hoffmann 1532 (US).<br />
FRENCH GUIANA. SAUL: Grand Boeuf Mort,<br />
03?37'N, 53?12'W, 250-300 m, 15 Jun 1988 (fl), Mori &<br />
Gracie 18943 (CAY); Petit BoefMort circuit, 21 Sep 1974<br />
(fr), Granville B-5167 (CAY, U, US); Monts La Fum6e,<br />
03?37'N, 53?12'W, 200-400 m, 2 Oct 1982 (fl), Mori et al.<br />
15109 (CAY, VEN), 15016 (P, US); Monts La Fumee, 200-<br />
400 m, 13 Oct 1982 (st), Boom & Mori 2037 (CAY), 2041<br />
(CAY), 2043 (CAY); layon toward Limonade bend, ca. 2<br />
km from village (crique Douille), 17 Jan 1976 (fr), Granville<br />
B-2671 (CAY, U); trail of Batardeau, 4 Mar 1977 (fr),<br />
Granville B-2799 (CAY, U); Mont Galbao, E sect., 600 m,<br />
03?35'N, 53020'W, 30 Jan 1986 (fr), Granville et al. 9041<br />
(B, CAY, US).<br />
BRAZIL. RORAIMA: Serra Tepequem, 1200 m, 17 Feb<br />
1967 (fr), Prance etal. 4452 (F, IAN, M, MG, NY, R, U, VEN).<br />
been misidentified as C. cymosa, but the latter has<br />
thick costate succulent branchlets, a much larger in-<br />
florescence, larger leaf blades (Fig. 65A-F), typical<br />
tuft-domatia (Fig. 58D), and is endemic to the Lesser<br />
Antilles (Fig. 63). As already noted by Steyermark<br />
(1965), C. longistipulata (from Surinam) is synony-<br />
mous with C. microcarpa.
Systematic Treatment of the Genera Studied 167<br />
9. Chimarrhis speciosa (Steyermark) Delprete, sp.<br />
et stat. nov. Chimarrhis microcarpa Standley var.<br />
speciosa Steyermark, Mem. New York Bot. Gard.<br />
12(3): 184. 1965. Type. Venezuela. Aragua:<br />
Rancho Grande, 18 Sep 1951, Garcia 147 (holo-<br />
type, VEN). Figs. 9H, 18A,B, 72A-G, 73<br />
Trees commonly 15-20 m tall, 30-40 cm dbh [exceptionally<br />
30 m tall, 80 cm dbh, Pittier 15275], with<br />
lobed bole, sometimes with small buttresses; bark<br />
grayish brown; wood yellow, very hard. Leafy<br />
branchlets (4-)5-8 mm thick, subterete to decussately<br />
compressed, greenish brown, often herbaceous,<br />
glabrous; older branches terete, grayish to pale brown;<br />
lenticels not found. Stipules free at base, contorted<br />
in bud, narrowly triangular, acuminate, sparsely appressed-puberulent<br />
(rarely glabrous) toward base and<br />
at margins outside, glabrous with basal colleters inside,<br />
25-30 x 5-7 mm, reddish brown, membranous,<br />
with many parallel veins departing from base, readily<br />
caducous, leaving a scar encircling the stem ca. 1 mm<br />
wide. Leaves (12-)18-27 x (5-)9-12 cm, L/W= 1.5:1<br />
to 2:1, widely elliptic to obovate, acute-decurrent at<br />
base, obtuse and mucronate at apex, the deltoid acumen<br />
0.5-1.8 cm long; dark green above, pale green<br />
below, semi-coriaceous; drying olive-green, stiffchartaceous;<br />
glabrous above and below; primary and<br />
secondary veins glabrous to sparsely appressed-puberulent,<br />
secondary veins (9-)11-12 each side; tertiary<br />
veins subparallel and laterally jointed in the center;<br />
petioles (10-)18-36 mm long, (1-)2-3 mm thick,<br />
adaxially flattened to concave, winged, glabrous; tuftdomatia<br />
always present, only at axil of primary with<br />
secondary veins, tuft of hairs departing from the veins<br />
and converging toward the center of the axil, hairs 0.4-<br />
0.6 mm long, golden-yellow (sometimes the hairs<br />
dense, straight, velutinous, white, starting before axil).<br />
Inflorescences 2 per node, corymbose with lax opposite<br />
to subopposite decussate branches, with small,<br />
very compacted cymules at distal end of each branchlet,<br />
not forming a continuous flowering plane but a rather<br />
sparse inflorescence, (10-)13-18(-22) x 7-13.5 cm,<br />
lateral branches (2-)3-4 pairs, basal portion of axis<br />
not branched (5-)7-13 cm long; L/A = ca. 2:1; rachis<br />
terete to decussately compressed, rachis and branches<br />
sparsely short-puberulent; distal bracts usually absent<br />
or deltoid to narrowly triangular, to 2 x 0.5 mm;<br />
bracteoles subtending flowers absent. Flower sessile<br />
to short-pedicellate; pedicels to 0.7 mm long, glabrous<br />
to sparsely puberulent; hypanthium obconical, 0.8-<br />
1.4 x 0.9-1.3 mm, glabrous, reddish brown. Calyx<br />
extremely reduced, truncate to obviously lobed, 0.3-<br />
0.6 x 1-2 mm, glabrous; lobes absent to rounded, to<br />
0.3 mm long. Corolla cyathiform, deeply lobed, 1.6-<br />
2.4 mm long, cream-white to greenish white (turning<br />
pale yellow at maturity); tube short cylindrical, 0.8-<br />
1 x 1.2-1.5 mm, glabrous outside and inside; lobes 5,<br />
0.8-1.3 x 1-1.6 mm, ovate to oblong-ovate, rounded<br />
to obtuse at apex, glabrous outside, with a dense tuft<br />
of white-villous hairs at basal zone, with medial and<br />
distal zones glabrous inside. Stamens 5, exserted well<br />
above the corolla, equal, attached ca. 1 mm from the<br />
base of the tube; filaments 2.4-3.7 mm long, terete,<br />
basally flattened, connate to throat, with a tuft of dense<br />
white-pilose hairs at base; anthers elliptic, 0.5-0.7 x<br />
0.4-0.5 mm, dorsifixed at medial zone, base rounded,<br />
apex rounded to obtuse-mucronate. Pollen exine fo-<br />
veolate-reticulate. Style exserted, 2-2.8 mm long, as<br />
long as the corolla, glabrous, minutely costate to mi-<br />
nutely winged; style branches extruding as two lips<br />
above the corolla before anthesis, rounded to oblate,<br />
0.2-0.3 mm long, stigmatic surface microscopically<br />
(40x) papillose; ovary-galls not found. Capsules<br />
subglobose, obtuse at base, rounded to hemi-ovoid at<br />
apex, 1.8-2.3 x 1.5-2 mm, dark brown without len-<br />
ticels, glabrous throughout; disk black, exceeding the<br />
calyx; disk loculicidal dehiscence present in old cap-<br />
sules. Seeds 0.7-0.95 x 0.5-0.75 mm.<br />
Distribution (Fig. 73) and ecology. Restricted to<br />
cloud forests on steep slopes, usually in rich mixed<br />
evergreen forest, 1000-1500 m (500 m in Guatopo<br />
National Park), of the coastal Cordillera of northeast-<br />
ern Venezuela (states of Carabobo, Aragua, Lara, and<br />
Miranda). Flowering specimens were collected only<br />
in January, while specimens at late blooming-early<br />
fruiting stage have been collected in March, Septem-<br />
ber, and October. Fruiting specimens were collected<br />
March, May, July, October, and November.<br />
Specimens examined. VENEZUELA. ARAGUA: Parque<br />
Nacional Henry Pittier, Quebrada Palo Vaco, trail up Periquito<br />
Parque Nacional Henry Pittier, 1300-1400 m, 25 Oct<br />
1961 (fr), Steyermark 89885 (NY, VEN[2]); Parque Nacional<br />
Henry Pittier (Rancho Grande), 16 Jul 1963 (fr), Uferide 13<br />
(VEN); rd. Rancho Grande-Gran Regresiva, 18 May 1984<br />
(fr), Field 424 (K[2]); Rancho Grande, S slope, 1000 m, 8<br />
Jan 1937 (fl), Pittier 13854 (F, NY, US, VEN); Rancho<br />
Grande, N slope, 1100 m, s.d. (fl), Pittier 14138 (F, NY, US,<br />
VEN); vic. of Alto de Rancho Grande, 29 Oct 1946 (fl-fr),<br />
Pittier 15275 (US, VEN); without locality, 30 Dec 1946 (fl),<br />
Lasser 2264 (VEN). CARABOBO: Between La Toma and<br />
"Capuchinos," S of Borburata, above Rio San Gian, 750-<br />
1000 m, 28 Mar 1966 (fr), Steyermark & Steyermark 95211<br />
(F, NY, U, US). FALC6N: Dto. Acosta, Mun. Jacura, Cerro<br />
La Mina, 350-650 m, 17 Feb 1961 (fl), Ruiz Teran 488<br />
(VEN[2]). MIRANDA: Parque Nacional de Guatopo, 48 km<br />
NNW of Altegracia, 500 m, 11 Sep 1960 (fl-fr), Steyermark
168 Flora Neotropica<br />
0~~~~<br />
~~~~~~~~~~<br />
' It<br />
FI.72 hiarhssecoa AF rm iter185,NY fo ah 33,NY.A.Hbt finlrecnc n<br />
mature!leaf B. Flower wih exserted anthers. C. ariation in calyx lobes.D. Detail of corolla, iside view. E. Stamen. F<br />
A<br />
o i.<br />
,,.~\:: ?<br />
:..:'<br />
'~..__.....~~<br />
~~' '? ~ ~ ???<br />
FIG. 72 Chimarhis speiosa (AF from ittier 3854, Y; G frm dahn 339, NY. A. Hait of iflorescnce an<br />
Apical portion style. of<br />
G. Mature capsule.~;T;'
Systematic Treatment of the Genera Studied 169<br />
*6'<br />
FIG. 73. Distribution of Chimarrhis microcarpa (circles)<br />
and C. speciosa (stars).<br />
87104 (NY, US, VEN). WrrmoIT LOCAurT: Aguacatal, 1000<br />
m, Jul 1919 (fr), Jahn 1339 (NY).<br />
Local name. Venezuela: santonino (Jahn 1339).<br />
This species is unique in having the smallest corolla<br />
(1.6-2.4 mm long; see Fig. 72D) in the genus<br />
(followed by C. parviflora of Central America), shortcyathiform<br />
corolla tubes, large laxly branched inflorescences<br />
(Fig. 72A), with irregularly separated distal<br />
units of compacted cymules, and semi-succulent<br />
leafy branchlets.<br />
Chimarrhis speciosa is closest to C. microcarpa<br />
in having capsules similar in size and shape (subsessile<br />
in the former), glabrous disks, and leafdomatia as tuft<br />
of pilose hairs at the vein axils; the former differing<br />
from the latter in being much larger (rain forest<br />
canopy) trees with bigger buttresses, longer corollas,<br />
smaller inflorescences, thinner woody leafy branchlets,<br />
and longer pedicels. The two species also occupy<br />
two different altitudinal ranges: C. microcarpa occurs<br />
in lowland (50-500 m) rain forests (on lateritic soils)<br />
of northeastern South America, and C. speciosa is<br />
restricted to steep-sloped cloud forests (1000-1400 m)<br />
of the Venezuelan coastal Cordillera.<br />
As already noted by Steyermark (1965, 1974),<br />
there are several collections that are transitional between<br />
C. microcarpa and C. speciosa, which were<br />
collected in the state of Miranda (where the two species<br />
come into contact) at intermediate elevations (50-<br />
500 m) in Guatopo National Park and Cerros del<br />
Bachiller. These specimens have transitional morphological<br />
characters: inflorescences not laxly branched<br />
and of intermediate length, leafy branchlets of intermediate<br />
thickness, capsules short-pedicellate (neither<br />
subsessile nor long-pedicellate). None of these tran-<br />
sitional specimens were collected in flower, so the<br />
dimensions and shapes of their corollas are unknown.<br />
Steyermark 87104 from Guatopo, which was reported<br />
as transitional by Steyermark (1965, 1974), is a good<br />
representative of C. speciosa. The transitional speci-<br />
mens between Chimarrhis microcarpa and C.<br />
speciosa are as follows: VENEZUELA. MIRANDA:<br />
Parque Nacional Guatopo, rd. Santa Teresa-<br />
Altagracia de Orituco, Rio Santa Cruz, 520 m, 23 Nov<br />
1961 (fr), Steyermark 89961 (NY, US[2], VEN);<br />
Bosque del Guatopo, 450-600 m, 29 Nov 1956 (fr),<br />
Bernardi 5943 (G[2], NY, VEN); Cerros del<br />
Bachiller, S of Santa Cruz, 10009'N, 65?48'W, 20-65<br />
m, 16 Mar 1978 (fr), Steyermark & Davidse 116278<br />
(MO, NY), 20-26 Mar 1978 (fr), 116885 (MO, VEN).<br />
CHIMARRHIS JACQUIN SUBGEN.<br />
PSEUDOCHIMARRHIS<br />
Chimarrhis Jacquin subgen. Pseudochimarrhis<br />
(Ducke) Delprete, subgen. et stat. nov.<br />
Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio<br />
de Janeiro 3: 255. 1922 (and Arch. Jard. Bot.<br />
Rio de Janeiro 4: 177, pl. 23. 1925). Type spe-<br />
cies. Pseudochimmarhis turbinata (A. P. de Can-<br />
dolle) Ducke (= Chimarrhis turbinata A. P. de<br />
Candolle).<br />
10. Chimarrhis turbinata A. P. de Candolle, Prodr.<br />
4: 404. 1830. Pseudochimarrhis turbinata (A. P.<br />
de Candolle) Ducke, Arch. Jard. Bot. Rio de<br />
Janeiro 3: 255. 1922. Chimarrhis duckei Rizzini,<br />
nom. superfl., Rev. Bras. Biol. 7: 277. 1947. Type.<br />
French Guyana. Ile de Cayenne, date unknown,<br />
Patris s.n. (lectotype, G-DC, here selected).<br />
Figs. 3Q, 10C, 18D, 58L, 74A-F, 75A-E,<br />
76A,B, 77A<br />
Elaeagia brasiliensis Standley in Gleason & Smith, Bull.<br />
Torrey Bot Club 60: 395. 1933. Type. Brazil. Para:<br />
Tapajos River region, Boa Vista, Fordlandia, 6 Sep<br />
1931 (fr), Krukoff 1018 (holotype, NY; isotypes,<br />
K, U).<br />
Pseudochimarrhis difformis (Benoist) Benoist, Arch.<br />
Bot. 5 (Mem. 1): 264. 1933. Bathysa difformis<br />
Benoist, Bull. Mus. Hist. Nat. Paris 26: 185. 1920.<br />
Type. French Guiana. Vic. of St-Jean-du-Maroni, 8<br />
May 1914 (fl-fr), Benoist 1190 (holotype, CAY;<br />
isotype, P).
170 Flora Neotropica<br />
g1<br />
FIG74Cimrrhsurbnaa(-EroIin a544,N; from Mo&Cardoso1712,GH).<br />
FIG. 74. Chimarrhis turbinata (A-E from Irwin et al. 55443, NY; F from Mori & Cardoso 17128, GH). A. Habit of<br />
inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style (note short antrorse<br />
pubescence). F. Mature capsule.
Systematic Treatment of the Genera Studied 171<br />
Trees to 40 m tall, to 170 cm dbh, bole irregular,<br />
much-branched at crown, fluted trunk, with buttresses<br />
to 3 m tall and extending several meters from trunk;<br />
bark deeply sulcate, lamellate, grayish white; wood<br />
yellow, very hard. Leafy branchlets (4-)7-11 mm<br />
thick, semi-succulent to woody, quadrangular to<br />
decussately compressed, terminal 2-3 nodes sparsely<br />
puberulent to densely pubescent; older branches<br />
glabrate, exfoliating in narrow strips, showing the redbrown<br />
bark just below; lenticels not found. Stipules<br />
free to connate at base, more or less contorted in bud,<br />
often adnate to petioles, deltoid to narrowly triangular,<br />
strongly acuminate, golden-pubescent outside,<br />
often with ciliate margins, glabrous with sparse small<br />
colleters at base inside, 9-20 x 6-8 mm, subcaducous,<br />
reddish green, leaving a scar encircling the stem, adnate<br />
to the petiole scar 1-3 mm wide. Leaves 9-19<br />
(-33) x 6-10.5(-16) cm, L/W = 1.5:1 to 2:1, oblong<br />
to elliptic to obovate, acute (rarely decurrent) to obtuse<br />
at base, obtuse to shallowly acute at apex, acumen<br />
usually absent; dark green, often shiny above,<br />
pale green-yellowish below, thick-chartaceous to<br />
semi-coriaceous; drying olive-green above, rustbrown<br />
below, thick-chartaceous to leathery; glabrous<br />
above, glabrous to puberulent (rarely pubescent) below;<br />
primary and secondary veins glabrous to puberulent<br />
(rarely pubescent), prominent below; secondary<br />
veins 8-12(-16) each side; tertiary veins alternatively<br />
subparallel and reticulate, quaternary veins finely reticulate,<br />
very distinctive from other species; petioles<br />
4-15(-22) mm long, 1-2.5 mm thick, terete to adaxially<br />
flattened or concave, (glabrous-) puberulent to pubescent;<br />
domatia absent or present, usually a tuft of<br />
sparse yellow hairs, puberulent to pubescent (Fig.<br />
57L). Inflorescences 2 per node, corymbose, laxly<br />
branched, with opposite to subopposite decussate<br />
branches, cymules lax to fairly compacted at distal end<br />
of each branchlet, (8.5-)11-13.5(-32) x cm long. Corolla narrowly funnelform, deeply lobed,<br />
the lobes reflexed at anthesis, 3.5-6 mm long, white<br />
to yellowish white; tube long-cylindrical, 1.5-3 x 0.7-<br />
1 mm, glabrous outside and inside; lobes (4-)5, 2.5-<br />
3.5 x ca. 0.5 mm, oblong with acute apex and round<br />
tip, glabrous outside, with a basal area pilose to tomentose<br />
(ca. 1 mm wide), with distal zone glabrous<br />
inside. Stamens 5, exserted well above the corolla,<br />
equal, attached 1-2.5 mm from the base of the tube;<br />
filaments 3.5-5.5 mm long, terete, minutely costate,<br />
basally flattened, connate to throat, sparsely white<br />
long-pilose at base, the hairs decreasing in length toward<br />
medial zone; anthers elliptic, 0.8-1.5 x 0.3-0.5<br />
mm, dorsifixed near medial zone, base rounded, apex<br />
rounded or minutely mucronate. Pollen exine foveolate-reticulate.<br />
Style exserted, 4-6 mm long, as long<br />
as the corolla, terete, densely antrorse-strigose; style<br />
branches extruding as two lips above the corolla before<br />
anthesis, oblate to oblong, 0.3-0.5 mm long, stigmatic<br />
surface microscopically (40x) papillose; ovaries<br />
often infested by insect, causing production of<br />
pyriform fruit-galls. Capsules turbinate, acute at base,<br />
rounded at apex, 3.5-6 x 2.5-3.3 mm, rust-brown,<br />
without lenticels, sparsely to densely puberulent below<br />
the disk; disk densely white-pilose, not exceeding<br />
the calyx; disk loculicidal dehiscence present in<br />
old capsules. Seeds 0.75-1.95 x 0.75-1.25 mm.<br />
Distribution (Fig. 77A) and ecology. Primary and<br />
disturbed non-inundated rain forests (terra firme), and<br />
riverine forests, in lateritic soils, clay to sandy-clay<br />
soils, lowland to mild slopes, 50-400 m, of Guyana,<br />
Surinam, French Guiana, and Brazil. Flowering specimens<br />
were collected in March, June, July, August,<br />
September, and October. Fruiting specimens were<br />
collected in all the months from June through Janu-<br />
5-10(-18)<br />
cm, lateral branches 1-2(-4) pairs, basal portion of<br />
axis not branched 5-8(-15) cm long; L/A = 2:1 to<br />
2.8:1; rachis basally terete, distally decussately compressed,<br />
rachis and branches sparsely to densely yellow-puberulent;<br />
distal bracts from reduced deltoid<br />
scales 0.2-0.3 mm long to linear bracts to 7 x 1-2 mm,<br />
sometimes leaf-like, to 7 x 2.7 cm, glabrous to puberulent,<br />
the margin sometimes ciliate; bracteoles<br />
subtending flowers absent. Flowers sessile to shortary.<br />
Boom and Campos (1991) reported that C.<br />
turbinata "is ecologically the most important <strong>Rubiaceae</strong><br />
in the BDFF [Biological Dynamics of Forest<br />
Fragments; located north of Manaus] reserves" and<br />
that its "distribution... suggests a tendency of clumping.<br />
Geographic distribution is mostly eastern Amazonian;<br />
this species' occurrence in this study are the<br />
westernmost records we have seen. Ecologically, it<br />
is restricted to terra firme, at elevations of up to 350<br />
m. Flowering: Jun-Oct. Fruiting: Jun-Jan."<br />
pedicellate; pedicels to 1.5 mm long, densely puberulent;<br />
hypanthium obovoid-turbinate, 1.5-2.5 x 1-1.6<br />
mm, densely puberulent. Calyx extremely reduced,<br />
truncate to short-lobed, 0.4-0.7 x 1-1.6 mm, glabrous<br />
to sparsely puberulent; lobes shallowly triangular, to<br />
0.2 mm long; rarely in some flowers one of the calyx<br />
lobes enlarged into a pale-green calycophyll, to 1.5<br />
Specimens examined. GUYANA. Gunns Strip,<br />
Essequibo River, 01?39'N, 58?38'W, 240-260 m, 30 Sep<br />
1989 (fr), Jansen-Jacob et al. 1899 (B, U).<br />
SURINAM. Forest reserve, section 0, tree n. 577, Forestry<br />
Bureau 1151 (U), 3 Jun 1916 (fr) 2040(U), 7 Sep 1916<br />
2415 (A, BR, IAN, U), 31 Oct 1916 (fr) 2464 (G, MO, U),
172 Flora Neotropica<br />
FIG. 75. Leaf and inflorescence variation in Chimarrhis turbinata (A from Boom et al. 8756, NY; B and D from Schultz<br />
7289, G; C from unknown collector 2907 [Surinam], MO). A, B. Inflorescence range. C-E. Leaf range. All drawn to the<br />
indicated scale bar.
Systematic Treatment of the Genera Studied 173<br />
11 Jan 1917 (fr) 2585 (K, NY, U, US), 5 Jun 1917 (fl) 2907<br />
(G, MO, U), 10 Oct 1920 (fl) 4952 (NY, U, US); vie. of Camp<br />
8, 17 Aug 1955 (fl), Schulz 7289 (G, NY, U); Komaramaraballi,<br />
1 Nov 1944 (st), Stahel 261 (A[2], B, MO, NY, U),<br />
Jun 1945 261a (U); Lucie River, below confluence of Oost<br />
River, 03020-32'N, 56?26-49'W, 225 m, 7 Sep 1963 (fl),<br />
Irwin et al. 55443 (F, GH, MO, NY, US); vic. of Nassau, 18<br />
Mar 1949 (st), Lanjouw & Lindeman 2810 (U).<br />
FRENCH GUIANA. CAMOPI: Trois Sauts, Mitake, 18<br />
Sep 1977 (fr), Grenand 1500 (CAY). CAYENNE: Approague<br />
River, confluence with Arataye River, Parare waterfalls, 25<br />
Nov 1985 (st), Barrier 5060 (CAY); Orapu, rd. to Fourgessi6,<br />
km 8,200 m, 15 Oct 1966 (fl), Oldeman B-674 (CAY,<br />
P); Sinnamary River, Petit Saut, 05?03'N, 53?03'W, 26 Jul<br />
1988 (fr), Sabatier & Prevost 2116 (CAY, US); Sinnamary<br />
River, above Petit Saut, between Petit Crique and Plomb and<br />
Crique Tigre, 87 m, 05?00'N, 53001'W, 6 Sep 1993 (st),Mori<br />
et al. 23645 (NY); Sinnamary-Counamama River Basin,<br />
Piste de Saint-Elie, 05020'N, 53?00'W, 13 Aug 1991 (st),<br />
Sabatier & Prevost 3731 (CAY, US). MARIPA-SOULA: Maroni<br />
River Basin, Grand Inini River, 03?40'N, 53?50'W, 19 Jul<br />
1990 (st), Sabatier & Prevost 3356 (CAY). SAOL: Bellevue,<br />
250 m from bridge, left shore of Crique Bellevue, 1.2 km<br />
from Me Madere Victor, 6 Dec 1956 (fr), Bena 1262 (CAY);<br />
Gallion, km 30, Cascade, 13 Sep 1956 (fl), Bena 1187<br />
(CAY); rd. to Belizon, between Eaux Claires and Layon<br />
Biche, 03037'N, 53?12'W, 200-400 m, 4 Aug 1993 (fl), Mori<br />
et al. 23098 (CAY); Montagne du Chevaux, near Route de<br />
l'Est, 04?45'N, 52?28'W, 50 m, 27 Jun 1988 (fr), Gentry &<br />
Morawetz 63183 (CAY, MO); Limonade, ORSTOM, 15 Oct<br />
1977 (fr), Granville 5377 (CAY, U, US); rd. Charvein-<br />
Acarouany, km 1, 11 Mar 1954 (fl-fr),BAFOG 166M(CAY, U).<br />
BRAZIL. AMAPA: Rio Araguari, 01009'-26'N, 51?52'-<br />
Isabel, Jun 1908 (fl), Museu Goeldipersonnel 9425 (F, G),<br />
9713 (F, G, NY); Belem, Bosque Municipal, 17 Jun 1943<br />
(fl), Ducke 1235 (MG19119) (IAN, MG); Belem, Bosque<br />
Municipal, 16 Jul 1947 (fl), N. T. Silva 51 (B); Belem,<br />
Utinga-Providencia, 20 Aug 1914 (fr), Ducke s.n.<br />
(MG15757) (MG); Belem, 31 Aug 1922 (fr) and 23 Jun 1923<br />
(fl), Ducke s.n. (RBI 7356) (RB, U, US); Belem, Lago Catu,<br />
10 Sep 1914 (fr), Ducke s.n. (MG15480) (MG); Belem,<br />
Reserva do Mocambo, 16 Aug 1982 (fr), Ramos & Rosario<br />
15 (MG); Itacaiunas, affluent of Rio Tocantins, Serra<br />
Buritirama, 05?31'S, 50013'W, Sep 1970 (st), Pires & Belem<br />
13006 (IAN); Belterra, 26 Jun 1947, Black F8 47-955 (US);<br />
Melgaco, Reserva Nacional de Caxiufla, site of the Ferreira<br />
Penna Scientific Station, 2-15 Feb 1991 (st), Chagas et al.<br />
2217 (MG); Mun. Oriximina, Serra da Preciosa, 21 km N of<br />
Rio Cumina-Mirim, 130 m, Martinelli 6835 (NY); Carajas,<br />
Serra Norte, Maraba, 7 Aug 1982 (fr), Maciel et al. 776(IAN,<br />
MG, NY); Belem, Mata da Utinga, 15 Dec 1945 (st), Fr6es<br />
20777 (IAN, NY); rd. Belem-Brasilia, km 2, 28 Sep 1959<br />
(fr), Kuhlmann & Jimbo 310 (IAN, TEX), 329 (IAN, TEX);<br />
Planalto de Santarem, Currupiru, loc. Gato, 22 Aug 1954 (fr),<br />
Fr6es 31049 (UB); Santar6m, km 70 on rd. to Cachoeira do<br />
Palhao, of Rio Curua Una, 5 Oct 1966 (fr), Cavalcante &<br />
M. Silva 1598 (IAN, MG); Santar6m, km 35 on Palhao rd.,<br />
Igarape Curupira Camp, 28 Aug 1969 (fr), M. G. Silva &<br />
Rosdrio Souza 2420 (MG, US), 2 Sep 1969 (fr), 2475 (CAS,<br />
MG, US); Mun. Mez, confluence of Rio Pinaticaua and Rio<br />
Jaracu, 19 Sep 1955 (fr), Fr6es 32097 (UB); Mun. Curralinho,<br />
between Rio Mapua and Rio Piria, 18 Jul 1950 (fl),<br />
Black et al. F8 50-9831 (IAN); Moju, rd. to Acara, km 60,<br />
15 Aug 1975 (fr), N. T. Silva 3924 (IAN); Fazenda Uriboca,<br />
Pirelli, Jun-Aug 1958 (fr), Pires 6826 (UB), 6922 (UB),<br />
6969 (UB), 6978 (UB), 7107 (UB).<br />
58'W, 11 Sep 1961 (fl), Pires et al. 50853 (G, GH, IAN, M,<br />
MG, U, US); Serra do Navio, 26 Sep 1961 (fl), Pires et al.<br />
51224 (IAN, NY, U, US); rd. Calcoene-Oiapoque, 60 km<br />
SSE of Oiapoque, 03? 18'N, 51039'W, 2 Dec 1984 (fr), Mori<br />
Local names and uses. French Guiana: bois<br />
pagaie (Bena 1262, Granville B5377), bois chapelle<br />
(Bena 1262),citronelle (BAFOG 166-M), samaati-<br />
& Cardoso 17128 (F, GH, MG, MO, NY, US); Rio Oiapoque,<br />
3 km E of mouth of Rio Matura, 02?34'N, 52?31'W, 22<br />
Sep 1960 (fl), Irwin et al. 48440 (GH, IAN, MG, NY, U,<br />
US); Rio Jari, Monte Dourado, 9 Sep 1968 (fl), N. T. Silva<br />
905 (CAS, NY), 17 Sep 1968 (fl), 999(NY, US), 17 Oct 1968<br />
(fr), 1246 (NY); Rio Jari, Planalto A, km 11, 2 Aug 1969<br />
(fl), M. Silva 2568 (IAN, NY, TEX, UB); Porto Platon, Perimetral<br />
Norte, Cupixi, 14 Sep 1976 (fl), M. Silva 2794<br />
(INPA); Rio Jari, Monte Dourado, 4 Dec 1967 (fr), Oliveira<br />
3836 (NY), 29 Jan 1968 (fr), 4036 (NY); Camaipi,<br />
EMBRAPA Reserve, 0010'N, 51037'W, 3 Sep 1983 (fr),<br />
palioudou (BAFOG 166-M), citronelle rouge (commercial<br />
name, Bena 1262, BAFOG 1262). Surinam:<br />
samaati-palioudou (Paramaka, BAFOG Service<br />
Forestiere 7608), walalu (Way pi, Grenand 1500),<br />
sinjahoedoe (Sur., Lanjouw & Lindeman 2810), koemaramaraballi<br />
(Arow., Lanjouw & Lindeman 2810),<br />
kaire-e (Car., Lanjouw & Lindeman 2810), sienjahoedoe<br />
(Schultz 8581). Brazil: pau de remo (Para,<br />
Irwin et al. 48440, Oliveira 3854, Pires 48902)), ramal<br />
do pau rosa (Para, Silva & Rosario 3732), carapa-<br />
Mori et al. 15786 (MG, NY[2]). AMAZONAS: Rd. Manaus-<br />
Caracari, km 146,25 Sep 1973 (fr), Bisby et al. P18095 (US);<br />
Mun. Manaus, rd. ZF-3, km 24, Reserva N0 1202, 17 Jun<br />
1980 (fr), Lima & Zimmerman 519 (NY, U); rd. Manaus-<br />
Itacoatiara, km 165, 16 Oct 1965 (st), Rodrigues 8098<br />
(INPA), km 140, 4 Oct 1965 (st), 8099 (INPA), km 125, 4<br />
Feb 1970 (st), Rodrigues 8692 (INPA); rd. between Rio<br />
Araqa and Lago Castanho, toward Careiro, 13 Jul 1972 (fl),<br />
M. Silva 5073 (US). MARANHAO: Rio Alto Turiacu, Nova<br />
Esperanca, 02?55'S, 45?45'S, 29 Nov 1978 (fr), Jangoux &<br />
Bahia 86 (MG, NY, UB); PARA: Rd. Belem-Braganca, Santa<br />
nauba (Para, Black F8-4 7-955), carapanauba cinzeiro<br />
(Para, Black F8-47-955), paraka'y (MaranhAo, Balee<br />
& Ribeiro 431), paraku-'iran-'y (Maranhao, Balee &<br />
Ribeiro 2826).<br />
This species offers pale-yellow hard wood of first<br />
quality for construction of houses. In French Guiana,<br />
the Saramacas use the wood to make paddles, toys,<br />
combs, wooden spoons and benches, and as construction<br />
wood for various purposes; the bark is also used<br />
by the same tribe to make ropes and floral artifacts
174<br />
A<br />
?<br />
.?<br />
.<br />
.<br />
iii.=izi'i'<br />
B ? .. .<br />
!f '<br />
t.....<br />
*D?? L ,,: ? . i<br />
A''!<br />
.....:; i?" ''?..<br />
B ~~~X.~~S~B~ i~ ~ .;t~,<br />
FIG. ?6. Chiman'6i~<br />
'<br />
Flora Neotropica<br />
turbina~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~a.<br />
A. Inflorescence. B. Trunk with bnsnl Boom.)~~~~~~~~~~~~~~~~sf~i<br />
buttresses. (Photog~~~~~~~~~~~~~~~~~~~~<br />
by Brinn
Systematic Treatment of the Genera Studied 175<br />
for decoration of their houses. Various indigenous<br />
tribes in Brazil and in the Guianas use the plank wood<br />
from the buttresses (without cutting down the tree) to<br />
make paddles, from which the common name is derived<br />
["Pau de remo" (Portuguese), "Bois Pagaie"<br />
(French) = fore it belonged to the Cinchoneae. Bremekamp<br />
(1934) considered this separation unacceptable and<br />
returned them to Chimarrhis. Pseudochimarrhis is<br />
here treated as a subgenus of Chimarrhis, characterized<br />
by persistent, deltoid stipules.<br />
paddle wood].<br />
Benoist (1920) described Bathysa difformis (using<br />
his collection from French Guiana), which he later<br />
This species is a frequent canopy tree, and is rec- (Benoist, 1933) transferred to Pseudochimarrhis.<br />
ognizable by its large, turbinate capsules (hence the Although this name is based on material with immaname),<br />
pubescent disk not exceeding the calyx, and ture capsules, it is clear that these specimens belong<br />
ascending-puberulent slender style. The leaf blades to C. turbinata.<br />
and the inflorescences are very variable in size and The trunk of C. turbinata (Fig. 76B) is usually irshape<br />
(Fig. 75A-E) even within the same individual. regular, alveolate and fluted, with bark deeply sulcate,<br />
In some inflorescences of C. turbinata (Mori & rust-brown and fibrous, sapwood pale-yellow, and<br />
Cardoso 17128; Pires et al. 51224) the distal bracts heartwood dark brown. Color photographs of this<br />
subtending secondary branches are foliose, delicate, species can be seen in Parrotta et al., 1995 (306-307).<br />
and pale-green, which might be superficially confused<br />
with calycophylls, but these are growing at the inflo- Dubious collection: FRENCH GUIANA.<br />
rescence nodes, or more rarely at the tips of inflores- CAMOPI: Trois Sauts, Aft iti, 17 Apr 1976 (fr),<br />
cence branches (but not calyx lobes, and with pinnate Grenand 1238 (CAY[3]). The young branches are<br />
venation). Also, as observed by Ducke (1922), very sterile and the leaves very much resemble those of<br />
rarely one of the calyx lobes expands into small, pale C.turbinata; the infructescences are detached and<br />
green calycophyll (with palmate venation). probably collected from the forest floor. The fruits are<br />
Leafvestiture in this species is rather variable, but much smaller than those of C. turbinata, and similar<br />
in most cases the blades are glabrous. In a few col- to those of C. microcarpa. Since both species occur<br />
lections (Boom et al. 8756, Mori et al. 16248, Balee in this area, it is uncertain whether these specimens<br />
& Ribeiro 2826, and Pires 48902) the blades are<br />
represent a mixed collection or a chance hybrid bedensely<br />
golden soft-hirtellous (velvety to the touch) tween these two species.<br />
beneath, exceptionally large and often asymmetrical<br />
(see below). Domatia in C. turbinata are absent (on<br />
the pubescent blades a few short hairs are present at Deviating collections. A few collections (see bethe<br />
axils).<br />
low) have distinct morphological traits, as compared<br />
Chimarrhis turbinata is similar to C. barbata in with typical C. turbinata, having stipules free at base,<br />
having large capsules and nonglabrous styles; C. bar- not contorted in bud; obovate capsules (2.5-)3.5-8<br />
bata differs from the former in having smaller flowers,<br />
smaller anthers, ascending-pubescent thicker and<br />
shorter styles, and capsules with minute-puberulent<br />
disks exceeding the calices.<br />
Chimarrhis turbinata was first described by A. P.<br />
de Candolle (1830) based on material collected by<br />
Patris in Cayenna (French Guiana). One specimen<br />
preserved at Geneva (G, Candolle Herbarium) has<br />
been found and selected to be the lectotype. Elaeagia<br />
brasiliensis, as already pointed out by Steyermark<br />
(1965), is synonymous with C. turbinata. Rizzini<br />
(1947), declaring C. turbinata of dubious identity,<br />
proposed C. duckei (a superfluous name), which is<br />
here treated as synonymous with C. turbinata.<br />
Ducke (1922) separated the genus Pseudochimarrhis<br />
from Chimarrhis so as to include C. turbinata and<br />
C. barbata [as P. barbata], stating that the general<br />
morphology of the flowers and the vertically inserted<br />
seeds resembled those of Calycophyllum, and there-<br />
x<br />
2-5 mm, golden-puberulent; uncommonly large<br />
(23.5-34 x 10-16 cm, L/A = 2:1) often asymmetrical<br />
leaf blades, densely golden-hirsute; and uncommonly<br />
large inflorescences (ca. 33 cm long). Flowers, pollen,<br />
and seeds of these specimens are still unknown,<br />
and future collections might show that these samples<br />
deserve nomenclatural recognition.<br />
Specimens examined. BRAZIL. AMAPA: Camaipi,<br />
EMBRAPA reserve and vic., 00?10'N, 51?37'W, 17 Sep<br />
1983 (st), Mori et al. 16248 (NY). AMAZONAS: Distrito<br />
Agropecuario, Reserva 1501 of the WWF/INPA Project, 50-<br />
125 m, 6 Dec 1988 (fr), Boom et al. 8756 (INPA, K, NY,<br />
US); Distrito Agropecuirio da SUFRAMA, Mun. Manaus,<br />
rd. BR-174, 02?25'S, 59?51'W, 13 May 1985 (fr), INPA/<br />
WWF 1109.25 (NY). MARANHAO: Mun. Carutapera, Gurupiuna,<br />
Ka'apor Indian Reserve, affluent of Rio Gurupi,<br />
02?43'S, 46026'W, 4 Nov 1986 (st), Balee & Ribeiro 2826<br />
(NY). PARA: Belem, grounds of Instituto Agronomico do<br />
Norte, 1 Nov 1960 (st), Pires 48902 (NY, SP).
176 Flora Neotropica<br />
11. Chimarrhis barbata (Ducke) Bremekamp,<br />
Recueil Trav. Bot. Neerl. 31: 260. 1934.<br />
Pseudochimarrhis barbata Ducke, Arch. Jard. Bot.<br />
Rio de Janeiro 4: 177, pl. 23. 1925. Type. Brazil.<br />
Para: Tapajoz River, between Boim and Serra<br />
Humayta, 8 Apr 1924 (fl-fr), Kuhlmann s.n. (RB<br />
17384) (holotype, R; isotypes, B*, U, US; photo-<br />
B at MO, F, NY). Figs. 77B, 78A-F<br />
Trees to 30 m tall, bole irregular; bark deeply fissured;<br />
wood yellow, very hard. Leafy branchlets 2-<br />
4 mm thick, thin-woody, glabrous to sparsely puberulent;<br />
older branches glabrate, exfoliating in narrow<br />
fibrous strips, dark brown; lenticels punctiform to lin-<br />
ear, to 2 mm long. Stipules connate at base, adnate to<br />
the petioles, not contorted in bud, deltoid to narrowly<br />
triangular, mucronate, puberulent outside, often with<br />
ciliolate margins, glabrous and with sparse colleters<br />
at base inside, 4.5-15 x 4.5-7 mm, subcaducous, leaving<br />
a scar encircling the stem. Leaves 6-16 x 4-7 cm,<br />
L/W = 1:1 to 2:1, elliptic, obovate, to widely ovate,<br />
acute to obtuse at base, acute to obtuse at apex, sometimes<br />
short mucronate, semi-coriaceous; glabrous<br />
above and below, drying olive-green; primary and<br />
secondary veins glabrous, prominent below, secondary<br />
veins 6-10 each side; tertiary veins subparallel and<br />
reticulate; petioles 6-20 mm long, 1-3 mm thick,<br />
adaxially flattened or concave, glabrous to minutely<br />
puberulent; domatia absent or few sparse hairs present.<br />
Inflorescences 2-4 per node, corymbose, short pedunculate,<br />
5-22 x 4.5-12 cm, lateral branches one<br />
pair; rachis basally terete, decussately compressed<br />
distally; L/A = 2.1:1 to 2.5:1; rachis and branches<br />
sparsely puberulent; distal bracts extremely reduced,<br />
1-5 x ca. 1 mm, linear to deltoid, rarely foliose, to<br />
1.5 cm long, narrowly elliptic; bracteoles subtending<br />
flowers deltoid scales ca. 0.5 mm long. Flowers<br />
sessile to short-pedicellate; pedicels to 0.5 mm long,<br />
sparsely short-puberulent; hypanthium obconical,<br />
1.5-2.5 x 1-1.5 mm, golden short-puberulent. Calyx<br />
extremely reduced, truncate to short-lobed, 0.5-1 x<br />
1.5-2.5 mm, glabrous to sparsely puberulent; lobes<br />
absent to shallowly triangular, to 0.5 mm long; rarely<br />
one or two flowers per inflorescence with one calyx<br />
lobe expanding into a small greenish white<br />
calycophyll. Corolla funnelform, deeply lobed, the<br />
lobes reflexed at anthesis, 3-4.5 mm long, white to<br />
greenish white; tube short cylindrical, 1-1.8 x 1-2 mm,<br />
glabrous outside and inside; lobes 5, 2-3.5 x 0.7-1.5<br />
mm, oblong with round apex, glabrous outside, with<br />
a basal area white pilose and distal zone glabrous inside.<br />
Stamens 5, exserted well above the corolla,<br />
equal, attached 1-1.5 mm from the base of the tube;<br />
filaments 2.5-3 mm long, terete, basally flattened,<br />
densely white pilose at base; anthers elliptic, 0.8-1 x<br />
0.3-0.5 mm, medially dorsifixed, base rounded, apex<br />
rounded or minutely mucronate. Pollen exine foveolate-reticulate.<br />
Style exserted, 4-5 mm long, terete,<br />
densely strigose, appressed-retrorse; style branches<br />
protruding as two lips above corolla before anthesis,<br />
oblate to oblong, reflexed, 0.6-1 mm long, stigmatic<br />
surface minutely papillose. Capsules pedicellate,<br />
pedicels to 2 mm long, narrowly turbinate, acute at<br />
base, rounded at apex, 5-9 x 2.5-5 mm, rust-brown<br />
to black-brown, without lenticels, sparsely goldenpuberulent<br />
below the disk; disk short white-puberulent,<br />
obviously exceeding the calyx; disk loculicidal<br />
dehiscence not seen. Seeds 1-1.5 mm long. 0.8-1.25<br />
mm wide.<br />
Distribution (Fig. 77B) and ecology. Primary,<br />
secondary, and fragmented non-flooded Amazon for-<br />
est (terra firme), in the Brazilian states of Para (Rio<br />
Tapajoz, Boim, and Itaituba) and Amazonas (Manaus,<br />
Tef6, and Villa Braga). Flowering specimens were<br />
collected in October, November, and December. Fruit-<br />
ing specimens were collected in April and August.<br />
Boom and Campos (1991) reported that the distri-<br />
bution of C. barbata "within the BDFF reserves is<br />
sporadic. Geographic distribution is strictly central<br />
Amazonian. The species is rare and/or rarely col-<br />
lected. Ecologically, is known only from terra firme<br />
forest." Prance et al. (1976) cited C. barbata in the<br />
<strong>Rubiaceae</strong> encountered in a hectare plot 30 km north<br />
of Manaus.<br />
Specimens examined. BRAZIL. AMAZONAS: Villa<br />
Braga, 1924 (fr), Ducke s.n. (RB17358) (B*, NY, U, frag-<br />
F); Manaus, Mindu, 22 Oct 1929 (fl), Ducke s.n. (RB22847)<br />
(K, U, US); Manaus, Villa Municipal, 11 Nov 1931 (fl),<br />
Ducke 38 (Yale 20719) (A, F, US); Manaus, Villa Munici-<br />
pal, 26 Dec 1941 (fl), Ducke 860 (F[2], IAN, MG, NY, R[2]);<br />
Rio Negro, SAo Gabriel, 16 Feb 1936 (young fr), Ducke s.n.<br />
(RB35055) (K, RB); upper Rio Solimfes, Mun. Tef6, Villa<br />
Nogueira, Xelele, 02?18'S, 64?48'W, 10 Nov 1986 (fl), Daly<br />
et al. 4312 (NY); rd. Manaus-Itacoatiara, km 26, Reserva<br />
Florestal Ducke, 02?53'S, 59?58'W, Reserva Florestal<br />
Ducke, tree N. 3220-15, 26 Sep 1995 (fl), Sothers et al. 568<br />
(INPA 185102) (INPA, NY); Reserva Florestal Ducke, tree<br />
N. 3220-15, 9 Feb 1996 (fr), Campos et al. 468 (INPA<br />
185019) (INPA, NY); Reserva Florestal Ducke, tree N. 3357-<br />
15, 12 Oct 1995 (fl), Sothers et al. 625 (INPA 185105)<br />
(INPA, NY); Reserva Florestal Ducke, tree N. 3357-15, 17<br />
Jan 1996 (fr), Sothers et al. 750 (INPA 185110) (INPA, NY);<br />
Reserva Florestal Ducke, tree N. 4384-15, without date,<br />
Assuncao 369 (INPA); Itaituba, Parque Nacional do Tapaj6s<br />
(IBDF), km 85 on rd. to Itaituba, 14 Nov 1978 (fl), M.G.<br />
Silva & Rosdrio 3732 (MG). PARA: Santar6m, Reserva For-
Systematic Treatment of the Genera Studied 177<br />
so 70 s So / 0<br />
16 l<br />
0t<br />
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.<br />
FIG. 77. Distribution of Chimarrhis. A. C glabrflora (circles), C turbinate (diamonds), and C brevipes (stars). B. C<br />
hooleri (circles), C. barbata (solid slurs), C. dulckeana (open sic-s), and C. gentryana (Uriangles).<br />
estal Curuana (SUDAM), right margin of Rio Curuana,<br />
Barreirinha, 19 Aug 1988 (fr), Rosdrio et al. 852 (MG), 858<br />
(IAN, MG).<br />
Local name Brazil: pau amarelo (Prance et al., 1976).<br />
Chimarrhis barbata is distinguished by its large capsules<br />
with puberulent to minutely puberulent disks, which<br />
exceed the calices (Fig. 78F). Chimarrhis barbata is a<br />
rare species of large canopy-trees endemic to the Brazilian<br />
Amazon, with deeply fissured, fibrous, reddish<br />
brown bark, extremely hard pale-yellow wood, semileathery<br />
leaves, and cream-white to greenish white,<br />
very fragrant flowers visited by bees. Because of its<br />
scanty collections, this is probably the least known<br />
species of Chimarrhis from the Amazon basin.<br />
Chimarrhis barbata is similar to C. turbinata in<br />
having large capsules, the latter differing from the<br />
former in having capsules with pubescent disk not<br />
exceeding the calyx, longer corollas, larger anthers,<br />
longer and thinner style with shorter style branches.<br />
At least one specimen of C. barbata [Ducke s.n.<br />
(RB53055)] has flowers with obvious calycophylls,<br />
about 4.5 x 2 cm, and greenish white when fresh (ac-<br />
cording to the label-data).<br />
Chimarrhis barbata is easily distinguishable from<br />
C. duckeana by its smaller leaves that turn pale ol-<br />
ive-green (to light-brown) when dry, capsules with<br />
ovoid disks much exceeding the calices, longer and<br />
slenderer antrorse-pubescent style, slender leafy<br />
branchlets exfoliating in small pieces, and smaller<br />
flowers.
178 Flora Neotropica<br />
Ic<br />
FI. Chimrribabr(ADfoDue224,U;EfoDayeal43,N;FfoKhman18,U,<br />
8<br />
isoyp). . abi oinforscnce. . Fowr wthexsrtd nthrs C.Deailofcorll, nsie iew D Stme. E Syl<br />
(note long antrorse-pubescence).<br />
F. Mature capsule.??<br />
FIG. 78. Chimarrhis barbata (A-D from Ducke 22847, US; E from Daly et al. 431, NY; F from Kuhlmann 17384, US,<br />
isotype). A. Habit of inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style<br />
(note long antrorse-pubescence). F. Mature capsule.
Systematic Treatment of the Genera Studied 179<br />
12. Chimarrhis duckeana Delprete, sp. nov., non<br />
oblong with round apex, glabrous outside, with basal<br />
Chimarrhis duckei Rizzini, nom. superfl., (based area densely white-pilose and distal zone glabrous<br />
on de Candolle's description of C. turbinata), inside. Stamens 5, exserted well above the corolla,<br />
Revista Brasil. Biol. 7: 277. 1947 . Type. Brazil.<br />
equal, attached ca. 1 mm from base of tube; filaments<br />
Amazonas: Manaus, Villa Municipal, non-flooded 4-5 mm long, terete, basally flattened, densely pilose<br />
secondary forest, 2 Jan 1930 (fl), Ducke s.n. (RB at base; anthers elliptic, 0.8-1 x 0.2-0.4 mm, medi-<br />
22846) (holotype, RB; isotypes, K, P, RB, U). ally dorsifixed, base rounded, apex rounded. Pollen<br />
Figs. 10B, O1F, 18E, 77B, 79A-F<br />
exine foveolate-reticulate. Style exserted, 2.5-3 mm<br />
long, minutely costate, densely retrorse white puberu-<br />
Arbores Chimarrhi barbatae similes, sed ab illa foliis lent; style branches protruding as two lips above comajoribus<br />
ambitu ellipticis ovatisve basi cordatis cordatisve, rolla before anthesis, ovate, reflexed, 0.3-0.5 mm<br />
brevius petiolatis, lamina in statu sicco nigrescenti (nec long, stigmatic surface minutely papillate. Capsules<br />
olivacea), stylo breviori crassiori minutim costato, secus<br />
pedicellate, pedicels to 2 mm long, turbinate to widely<br />
costas tantum puberulo (nec gracili tereti ubique retrorsus<br />
turbinate, acute at base,<br />
strigoso), necnon styli ramulis brevioribus diversa.<br />
oblong-rounded at apex, 6-<br />
12 x 4-6 mm, dark-brown, without lenticels, microscopically<br />
puberulent throughout; disk obviously exceeding<br />
the calyx; disk loculicidal dehiscence not<br />
seen. Seeds 1.15-2.38 x 0.83-1.5 mm.<br />
Trees 20-25 m tall, 30-55 cm dbh, mid-canopy<br />
trees; bark fissured, brown. Leafy branchlets<br />
decussately compressed, 4-6 mm thick, (4-)5-8 mm<br />
wide, semi-succulent, glabrous; older branchlets gla-<br />
brous, smooth (not exfoliating in narrow strips), gray-<br />
ish brown; lenticels linear, to 3 mm long. Stipules free<br />
or connate at base, not contorted in bud, narrowly tri-<br />
angular to deltoid, acuminate, glabrous outside (with-<br />
out ciliolate margins), glabrous and with sparse<br />
colleters inside, 7-19 x 7-10 mm, dark green,<br />
subcaducous to persistent. Leaves (8-) 18-21(-32) x<br />
(7-)10-18 cm, L/W = 1.1:1 to 1.2:1, widely elliptic<br />
to widely ovate, obtuse to cordate at base, rounded to<br />
obtuse at apex, drying dark-brown to black; primary<br />
and secondary veins glabrous, prominent below; sec-<br />
ondary veins 8-10 each side, widely spaced (2-3 cm<br />
intervein); tertiary veins subparallel, faintly evident<br />
above; petioles 25-43 mm long, 2-5 mm thick, ad-<br />
axially concave, glabrous; domatia absent. Inflores-<br />
cences 2-4 per node, corymbose, short to long pedun-<br />
culate, with opposite decussate branches terminating<br />
in dense cymules, (6.5-)12.5-22 x (3-)6-12 cm, lat-<br />
eral branches 1-4 pairs, basal portion of the axis not<br />
branched 5-14 cm long; L/A = 2:1 to 3:1; rachis ba-<br />
sally terete, decussately compressed distally, rachis<br />
and branches glabrous to minutely puberulent; distal<br />
bracts extremely reduced, 0.5-4 x 0.5-1 mm, deltoid<br />
to lanceolate (no foliose bracts observed); bracteoles<br />
subtending flowers deltoid scales ca. 0.5 mm long.<br />
Flowers pedicellate; pedicels 1-2 mm long, puberu-<br />
lent; hypanthium obconical, ca. 2 x 1.5-2 mm, pu-<br />
berulent. Calyx extremely reduced, short-lobed, 0.5-<br />
1.5 x 1.5-2.5 mm, glabrous to sparsely puberulent;<br />
lobes deltoid, ca. 0.5 mm long; flowers with<br />
calycophylls not found. Corolla funnelform, deeply<br />
lobed, the lobes reflexed at anthesis, 3-4.5 mm long,<br />
white; tube short-cylindrical, 1-1.5 x 1-1.5 mm, gla-<br />
brous outside and inside; lobes 5, ca. 2.5 x 1-1.5 mm,<br />
Distribution (Fig. 77B) and ecology. Primary,<br />
secondary, and fragmented forests in non-flooded<br />
Amazon (terra firme), in clay or sandy lateritic soil.<br />
Several specimens were collected in the surroundings<br />
of Manaus, but one specimen was collected on the Rio<br />
Negro. Flowering specimens were collected in January<br />
and February. Fruiting specimens were collected<br />
in March, July, August, September, and October.<br />
Specimens examined. BRAZIL. AMAZONAS: Rd.<br />
Manaus-Itacoatiara, km 26, Reserva Florestal Ducke,<br />
02?53'S, 59?58'W, parcel 22, tree N. 2938,22 Mar 1966 (fr),<br />
Rodrigues & Coelho 7600 (INPA 16903) (US); Reserva<br />
Florestal Ducke, tree N. 3553-15, 12 Dec 1995 (fl), Ribeiro<br />
& Assuncdo 1776 (INPA 185097) (INPA, NY); Reserva<br />
Florestal Ducke, 10 Aug 1976 (st), Oliveira s.n. (INPA<br />
60564) (INPA); Reserva Florestal Ducke, tree N. 1613<br />
(inventario florestal), 4 Sep 1968 (fr), W. Rodrigues 8573<br />
(INPA 21522) (INPA); Reserva Florestal Ducke, 16 Jul 1976<br />
(st), Adair s.n. (INPA 72795) (INPA); Manaus, estrada da<br />
Raiz, between Cachoerinha and Mind6, 30 Jul 1929 (fr),<br />
Ducke s.n. (RB 22845) (RB) [mixed collection]; Manaus,<br />
Campus Universitario do INPA, estrada do Japiim, 7 Dec<br />
1977 (st), Falcao & Coelho 209 (INPA); Manaus, Campus<br />
Universitario do INPA, estrada do Aleixo, 21 Feb 1977 (fl),<br />
Leite 2 (INPA); Manaus, margem do igarape do BuiAo, 10<br />
Jan 1955 (fl), J. Chagas 486 (MG25131) (IAN, INPA, MG);<br />
Dto. Agropecuario da SUFRAMA, rd. 174, km 64, Fazenda<br />
Esteio, Reserva N. 1109, tree N. 302, 02?25'S, 59049'W, 20<br />
Aug 1981 (fr), Nascimento et al. s.n. (INPA/WWF 1109.302)<br />
(NY); Manaus, Ponta Negra, Hotel Tropical, tree N. 17, 19<br />
Jul 1977 (fr), Rodriguez 9954 (INPA), 9976 (INPA); Rio<br />
Negro, Paduary, Mata do Tucano, 14 Oct 1947 (fr), Fr6es<br />
s.n. (RB 22596) (NY[2], SP, U, US).<br />
Local names and uses. Amarelinha<br />
(Chagas 486),<br />
pau amarelo (Adair s.n., INPA 72795). Label data
180 Flora Neotropica<br />
IN<br />
A~<br />
Pe<br />
~~~~~~~~~~~'5*<br />
r~~~~~~~~~~~~~ ?.<br />
?~~~~~~~~<br />
..: ~-;<br />
i "i' '"' ~...<br />
-.~~~~~~ o<br />
.<br />
i :'~~~~~~'<br />
~~~~~~~~~~~~~~.~:~<br />
.~~~r<br />
??<br />
!~~~~~.<br />
.....<br />
'.?'? ? B,<br />
FIG 7. hiarhi dckan (-Efrm uce 286,P,istye;F ro Nscmetoetal IPA! 09$0, Y) A<br />
pubescence inlniuiaros.FMauecpl.<br />
~ ~ ~<br />
FIG. 79. Chimarrhis duckeana (A-E from Ducke 22846, P, isotype; F from Nascimento et aL INPA/1 109.302, NY). A.<br />
Habit of inflorescences. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Stamen. E. Style (note antrorse<br />
pubescence in longitudinal rows). F. Mature capsule.<br />
o<br />
A<br />
i .o
Systematic Treatment of the Genera Studied 181<br />
from Chagas 486 reports that the wood of this spe- petioles, deltoid, glabrous outside and inside, densely<br />
cies is used in sawmills.<br />
covered with colleters throughout inside, 6-9 x 7-9<br />
mm, dark green, subcaducous, leaving a scar encir-<br />
Chimarrhis duckeana is distinguishable from C. cling the stem 1-2 mm wide. Leaves 17-23 x 11.5barbata<br />
by its larger ovate to cordate leaves with 22 cm, L/W = 1:1 to 1.5:1, widely pyriform, rounded<br />
shorter petioles; blades that turn black-brown when to subcordate at base, rounded to obtuse at apex, acudry<br />
(vs. olive-green to light-brown in C. barbata); men absent; dark green above and below, semi-coriashorter<br />
and stouter minutely costate style, puberulent ceous; drying dark olive-green above, reddish brown<br />
only on the costae (vs. slender terete [non costate], below, semi-leathery to leathery; glabrous above and<br />
and appressed-retrorse throughout in C. barbata); and below; primary and secondary veins glabrous, promishorter<br />
style branches. Nascimento et al. s.n. (INPA/ nent below, secondary veins 5-6 each side; tertiary<br />
WWF 1109.302) was reported to have been collected veins subparallel throughout; petioles (4-)7-11 mm<br />
from a tree 25 m tall and 55 cm dbh. The morpho- long, 3-4 mm thick, terete to adaxially flattened or<br />
logical variation in size and shape of the leaves and concave, glabrous; domatia absent. Inflorescences 2-<br />
architecture of the inflorescences of this species (as 4 per node, corymbose, densely branched distally,<br />
already reported in C. turbinata) makes it difficult to with opposite to subopposite decussate branches teridentify<br />
sterile specimens.<br />
minating in cymules, forming a divided flowering<br />
Fr6es 22596 has atypically long-pedunculate in- plane, (6.5-)9-15<br />
florescences and ovate leaves, but these two characters<br />
alone are, in my opinion, not sufficient for the<br />
establishment of a new taxon; future collections with<br />
flowers in anthesis may prove that this collection deserves<br />
at least varietal recognition.<br />
Ducke s.n. (RB 22845) sheet A (RB) is a mixed<br />
collection of C. barbata and C. duckeana. The specimen<br />
mounted on the upper portion of the sheet is C.<br />
duckeana, with blackish brown shiny leaves, stout<br />
inflorescences with mature capsules, and three calycophylls.<br />
The specimen in the lower portion of the sheet<br />
is C. barbata, with light-brown opaque leaves, smaller<br />
slenderer inflorescences in anthesis, and no calycophylls<br />
present. The presence of calycophylls is not<br />
uncommon in this species, which were also observed<br />
in Chagas 486 (IAN, INPA, MG).<br />
Adolpho Ducke (1876-1959) is the botanist that<br />
most contributed toward the understanding of the systematics,<br />
taxonomy, and phenology of the Amazonian<br />
species of Chimarrhis (see Delprete, 1996b: 37). It is<br />
a great honor to dedicate the epithet of this species to<br />
such a famous Amazonian botanist.<br />
13. Chimarrhis brevipes Steyermark, Mem. New<br />
York Bot. Gard. 12: 179. 1965. Type. Venezuela.<br />
Amazonas: Cerro Sipapo (Paraque), forest near<br />
Base Camp, 125 m, 25 Dec 1948 (fr), Maguire &<br />
Politi 27971 (holotype, NY; isotypes, U, US,<br />
VEN). Figs. 10A, 581, 77A, 80A,B<br />
Small trees. Leafy branchlets glabrous; older<br />
branches smooth, not exfoliating in narrow strips,<br />
grayish pale brown; lenticels not found. Stipules connate<br />
at base, not contorted in bud, often adnate to<br />
x 4.5-7.5 cm, lateral branches one<br />
pair, basal portion of axis not branched (3.5-)6-9.5<br />
cm long; L/A = 2.5 to 3:1, rachis basally subterete,<br />
distally decussately compressed, rachis and branches<br />
sparsely puberulent; distal bracts extremely reduced,<br />
deltoid to lanceolate, to 2.5 x 1 mm; bracteoles subtending<br />
flowers absent or reduced to microscopic<br />
scales. Flowers not seen, sessile to short-pedicellate,<br />
puberulent (observed from young fruits). Pollen not<br />
seen. Calyx extremely reduced, truncate to minutely<br />
lobed, 0.5-1 mm long, minutely puberulent; lobes<br />
barely distinguishable, to 0.5 mm long; rarely one<br />
calyx lobe expanded into a calycophyll, usually<br />
present on aborted flowers. Calycophylls ca. 5.5 x<br />
ca. 3.5 cm, blades obovate, with peltate venation,<br />
whitish; stalks ca. 1.8 cm long. Capsules obovoid,<br />
acute at base, rounded at apex, 4.5-6 x 2.5-3.5 mm,<br />
pale brown, without lenticels, minutely white puberulent<br />
below the disk; disk white-pubescent, not exceeding<br />
the calyx; disk loculicidal dehiscence present in<br />
old capsules. Seeds 0.85-1.05 x 0.63-1 mm.<br />
Distribution (Fig. 77A) and ecology. This species<br />
is known only from a few collections from lowelevation<br />
forests, on lateritic soil, 125-190 m, in the<br />
Venezuelan Amazon. Fruiting specimens were collected<br />
in February, April, and December.<br />
Specimen examined. VENEZUELA. TERRITORO FED-<br />
ERAL AMAZONAS: Dpto. Atabapo, plains of Rio Padamo,<br />
03?20'N, 65?14'W, 190 m, Feb 1990 (fr), Marin 740 (MO,<br />
NY); San Carlos de Rio Negro, 30 Apr 1982 (fr), Dezzeo 73<br />
(VEN).<br />
Dubious specimen. BRAZIL. RONDONIA:<br />
Projecto RADAM, Ponto 26-SC-20-XD, 30 Aug 1975<br />
(fr), Cordeiro 676 (VEN). This specimen has leath-
182 Flora Neotropica<br />
- -- --'--<br />
?<br />
| 1l.. . .S..* 1 | ? -<br />
. .<br />
--..gw -.1.... Ul<br />
?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...<br />
.. ? ..<br />
?I ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
V~~~~~~~~~~~?<br />
LA;<br />
tv1~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
? .i ?~~?<br />
~m~s ~ ~ ~ ??<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
. .?...P; ?<br />
?~~i .? ir<br />
ii i~~~~~~~~~~~~~~~~~~~~~~~~~<br />
..i~i' .. .. :.<br />
B . Mature capsule.<br />
".:~ ~~~~~~~~~~~~?.?<br />
.. ~ ~~<br />
.l ": i~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
? '~~~~~~~~~~~~~~~~~~~~~~~~~~~~?<br />
?<br />
.... ... ?= .<br />
''~~~~~~~~~~~~~~~~~~~~~.<br />
? ,~ .: .. .,~<br />
FIG. FIG. 80. Chimarrhris<br />
Chimarrhis brevipes brevipes (Maguire (Maguire &r & Politi 27971, 27971, US, isotype). A. Habit of infiorescences<br />
inflorescences and mature<br />
leaf.<br />
B. Mature capsule.<br />
>~<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.:....?<br />
?i~~~~~~~~~~~~~~~~~~~~~~~~~<br />
. "''<br />
?? ~ :? ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
?' .. ''<br />
r~~~~~~~~<br />
7Y. l??E'i~~~~~~~~~~~~~~~~~~~~:<br />
g rJ~~~~~~~~~~~I 'j:34~~~~~~. j...<br />
?... '.<br />
r ?: ~~~?~:~~ii~ ~~~ ? '~ i. E lr ?? . . . . . . . . ....... . ...... ?<br />
'<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~I.<br />
.:<br />
'......... .<br />
?
Systematic Treatment of the Genera Studied 183<br />
ery, pandurate leaves, and subsessile fruits, as in C. brevipes,<br />
but the fruits of this specimen are abnormally large<br />
and the collection locality is outside the expected range.<br />
brown; lenticels not found. Stipules connate at base,<br />
not contorted in bud, often adnate to petioles, deltoid,<br />
not acuminate, glabrous outside, microscopically<br />
(40x) ciliolate, glabrous with a basal triangular area<br />
Chimarrhis brevipes resembles C. gentryana in of colleters inside, 8.5-9 x 6.5-10 mm, reddish green,<br />
having leathery, pandurate leaves (usually much larger<br />
in C. brevipes), but the latter differs from the previous<br />
in having much smaller and narrowly obconical<br />
subcaducous, leaving a scar encircling the stem and<br />
connected to the petiole scars, 1-2 mm wide. Leaves<br />
(8-)13-24<br />
capsules.<br />
It is one of the three species of Chimarrhis (with<br />
C. microcarpa and C. speciosa) present in Venezuela<br />
(Chimarrhis venezuelensis is a Dioicodendron dioicum,<br />
q.v.). One isotype specimen (Maguire & Politi<br />
27971, NY) has an evident calycophyll on an aborted<br />
flower, not mentioned by Steyermark (1965, 1974).<br />
This calyx-lobe expansion is a calycophyll proper<br />
because it has peltate venation. In the other collections<br />
(Marin 740, MO; Dezzeo 73, VEN), no calycophylls<br />
are present; therefore, this feature is sporadic<br />
and of no taxonomic significance in C. brevipes. The<br />
occurrence ofcalycophylls in Chimarrhis is very rare,<br />
being sporadically present in this species and C. barbata,<br />
and commonly present only in C. gentryana.<br />
Chimarrhis brevipes is similar to C. turbinata in<br />
having puberulent capsules with small pubescent disks<br />
not exceeding the calices, the former differing from<br />
the latter in having leathery broadly pandurate leaves,<br />
and smaller obovoid capsules. From C. barbata this<br />
species differs in having smaller obovoid capsules<br />
with pubescent disk not exceeding the calices (vs. turbinate<br />
capsules with puberulent disk exceeding the<br />
calyx), and pandurate leaves.<br />
Chimarrhis brevipes differs from C. microcarpa<br />
and C. speciosa by: capsular disks white-pubescent<br />
(vs. glabrous in the latter two), domatia absent (vs.<br />
tufted domatia), bigger, obovoid capsules (vs. subglobose),<br />
and leaves usually pandurate (vs. elliptic to<br />
obovate).<br />
14. Chimarrhis gentryana Delprete, Brittonia 48:<br />
37. 196. Type. Peru. Loreto: Maynas, Mishana,<br />
Rio Nanay, halfway between Iquitos and Santa<br />
Maria de Nanay, ca. 150 m, 73?30'W, 03?50'S,<br />
upland forest mostly on white sand, 1 Mar 1980<br />
(fl), Gentry & Aronson 25307 (holotype, MO;<br />
isotype, NY). Figs. 10E, 58J, 77B, 81A-F<br />
Trees usually 15-20 m tall, exceptionally to 30 m<br />
tall; bark deeply fissured, fibrous; wood extremely<br />
hard. Leafy branchlets 4-8 mm thick, semi-succulent,<br />
terete to subterete, glabrous; older branches<br />
smooth (not exfoliating in narrow strips), grayish pale<br />
x (4-)8-13 cm, L/W = 1.5:1 to 1.7:1,<br />
pandurate, obtuse to rounded at base, concave at basal<br />
area, obtuse at apex, acumen absent; dark green above,<br />
pale green below, semi-coriaceous; drying reddish<br />
brown, semi-leathery to leathery; glabrous above and<br />
below; primary and secondary veins glabrous, prominent<br />
below, secondary veins 7-10 each side; tertiary<br />
veins subparallel throughout; petioles 6-12 mm long,<br />
2-3 mm thick, terete to adaxially flattened or narrowly<br />
concave, glabrous; domatia absent. Inflorescences 2<br />
per node, corymbose, densely branched distally, with<br />
opposite to subopposite decussate branches terminating<br />
in cymules, forming a continuous flowering plane,<br />
9.5-15 x 7.5-11 cm, lateral branches 1-2 pairs, basal<br />
portion of axis not branched 4.5-7.5 cm long; L/A =<br />
1.5 to 2:1; rachis basally subterete, distally decussately<br />
compressed, rachis and branches white or golden<br />
minute-puberulent; distal bracts absent or extremely<br />
reduced, deltoid to linear, to 4 x 1 mm; bracteoles subtending<br />
flowers absent or reduced to microscopic<br />
scales. Flowers sessile to short-pedicellate; pedicels<br />
to 1.5 mm long, puberulent; hypanthium narrowly<br />
obconical, 1-1.5 x ca. 1 mm, puberulent. Calyx extremely<br />
reduced, truncate to short-lobed, 0.4-0.8 x<br />
1.3-2 mm, golden puberulent, pale green; lobes barely<br />
distinguishable, to 0.5 mm long; in some flowers one<br />
calyx lobe expanded into a calycophyll. Calycophylls<br />
5-7 x 1.6-3 cm, blades ovate to elliptic, with peltate<br />
venation, pale green to white; stalks 1.5-2 cm long.<br />
Corolla funnelform, deeply lobed, the lobes reflexed<br />
at anthesis, 3.5-4.5 mm long, white to greenish white;<br />
tube short cylindrical, 1.5-2.5 x 0.5-0.7 mm, glabrous<br />
outside and inside; lobes 5, 2-2.5 x 0.6-0.8 mm, narrowly<br />
oblong with acute apex, glabrous outside, with<br />
a basal area white-pilose and with distal zone glabrous<br />
inside. Stamens 5, exserted well above the corolla,<br />
equal, attached 1-2 mm from the base of the tube; filaments<br />
3.5-5 mm long, terete, basally flattened, connate<br />
to throat, sparsely white pilose at base; anthers<br />
elliptic, 0.6-0.7 x 0.3-0.5 mm, medially dorsifixed,<br />
base rounded, apex acute to short mucronate. Pollen<br />
not seen. Style exserted, 5-6 mm long, terete, densely<br />
strigose, appressed-retrorse; style branches extruding<br />
as two lips above corolla before anthesis, oblate, 0.3-<br />
0.6 mm long, stigmatic surface microscopically (40x)<br />
papillose; ovary-galls present. Capsules narrowly<br />
obconical to turbinate, acute at base, rounded at apex,
184 Flora Neotropica<br />
A<br />
FIG. 81. Chimarrhis gentryana (A-E from Huashikat 2136, MO; F from Vdsquez & Arevalo 8969, TEX). A. Habit of<br />
inflorescence with mature leaf. B. Flower with exserted anthers. C. Detail of corolla, inside view. D. Style (note antrorse<br />
pubescence). E. Stamen. F. Mature capsule. (Reproduced with permission from Brittonia 48, fig. 1. 1996.)<br />
' o i'<br />
"~~~~~~,<br />
FIG81Charhsgnraa(-frmHahkt23,M;Ffo sqe&Aral899TX)A.Hbto<br />
pu~scn?) E Samn.F Mtue 8,f.1.19.<br />
apul. Rerduedwih erisio ro Bitona
Systematic Treatment of the Genera Studied 185<br />
3.5-5 x 1.5-3.5 mm, rust-brown, without lenticels,<br />
densely golden puberulent below the disk; disk white-<br />
pubescent, not exceeding the calyx; disk loculicidal<br />
dehiscence present in old capsules. Seeds 0.95-1.13<br />
x 0.58-0.88 mm.<br />
Distribution (Fig. 77B) and ecology. Primary and<br />
secondary high-canopy non-flooded rain forests (terra<br />
firme) with high canopy, in white sand and clay soil,<br />
of low elevations, 115-390 m, of Amazonian Ecua-<br />
dor, Peru, and Brazil. Flowering specimens were col-<br />
lected in February and March. Fruiting specimens<br />
were collected in March, April, July, and August.<br />
Specimens examined. ECUADOR PASTAZA: UNOCAL<br />
petroleum exploration well site, Mazaramu, 01?27'S,<br />
76?54'W, 390 m, 28 Apr 1990 (fr), Beck et al. 1055 (paratype,<br />
NY, QCNE); Kapawi, Rio Pastaza, 76?48'W, 02?31'S,<br />
235 m, 14-20 Jul 1988 (fr), Lewis et al. 13543 (QCNE).<br />
PERU. AMAZONAS: Valley of Rio Sagrado, 65 km N of<br />
Pinglo, Quebrada Caterpiza, 200 m, 19 Feb 1980 (fl),<br />
Huashikat 2136 (MO). LORETO: Prov. Maynas, Mishana, Rio<br />
Nanay, between Iquitos and Santa Maria de Nanay, 73?30'W,<br />
03?50'S, 150 m, 25 Jul 1984 (fr), Vdsquez et al. 5405 (MO,<br />
NY); Prov. Loreto, Nauta, road Nauta-Iquitos, 73?35'W,<br />
04?29'S, 200 m, 28 Mar 1987 (fr), Vdsquez & Arevalo 8969<br />
(F, TEX).<br />
BRAZIL. AMAZONAS: Mun. Sio Paulo de Olivenca,<br />
basin of Bel6m creek, Oct-Dec 1936 (fr), Krukoff8782 (MO,<br />
NY, U).<br />
This species is easily recognizable by its numerous<br />
cream-white calycophylls (Fig. 81A), rust-brown<br />
puberulent capsules, and puberulent disk which does<br />
not exceed the calyx. The calycophylls usually darken<br />
and break off at capsule maturity. The leaf blades of<br />
C. gentryana are commonly pandurate, semi-leathery,<br />
and olive-green when fresh, turning reddish brown<br />
when dry.<br />
Chimarrhis gentryana is most similar to C. brevipes<br />
(of the Venezuelan Amazon), both possessing<br />
pandurate leaves and calycophylls; the former differs<br />
from the latter in having narrowly obconical capsules<br />
(vs. broadly obovoid), which are rust-brown (vs.<br />
beige-brown) and densely golden-puberulent (vs.<br />
sparsely white-puberulent).<br />
Chimarrhis gentryana is often confused with Calycophyllum<br />
obovatum (Ducke) Ducke, because of its<br />
white calycophylls, minute white flowers, and obovate<br />
to pandurate leaves (Fig. 81A-E). The genus<br />
Calycophyllum differs from Chimarrhis in having<br />
terminal inflorescences (vs. axillary and subterminal),<br />
seeds with strongly bipolar wings (vs. minute seeds<br />
with small concentric wing), and smooth thin bark<br />
exfoliating into long vertical strips (vs. soft-fibrose<br />
bark falling off in small irregular pieces).<br />
Chimarrhis gentryana is commonly encountered<br />
as medium-sized trees (15-20 m tall; but two emergent<br />
individuals were reported to be 25 and 30 m tall),<br />
growing below the high canopy Amazon forest (30-<br />
35 m high), with small buttresses, rust-brown fibrous<br />
bark, leathery leaves, cream-white fragrant flowers,<br />
and cream-white calycophylls.<br />
Excluded and Dubious Species<br />
Chimarrhis bathysoides Steyermark, Mem. New York<br />
Bot. Gard. 12(3): 181. 1965. Type. Venezuela.<br />
Amazonas: Cerro Sipapo (Paraque), forest along<br />
trail near Base Camp, 25 Jan 1949 (fl), Maguire<br />
& Politi 28626 (holotype, NY; isotypes, B, GH,<br />
US[2], VEN[2]) = Bathysa bathysoides (Steyermark)<br />
Delprete (see Delprete, 1996b).<br />
Chimarrhis clausicorollata J. H. Kirkbride, Bio-<br />
Llania, Edicion Esp. 6: 394. 1997. Type. Guyana.<br />
Distr. Brokopondo, Kentoegoni Island, S of<br />
Gansee, 21 Jul 1966 (fl), van Donselaar 1471<br />
(holotype, U; photo-U at NY). = Simira tinctoria<br />
Aublet.<br />
This taxon has been recently described by Kirk-<br />
bride (1997a), based on a single specimen with young<br />
flowers. The terminal inflorescence and the reddish<br />
tinge in the leaves and the wood of this specimen prove<br />
that this species belongs to the genus Simira (Chimarrhis<br />
has lateral inflorescences and yellowish wood).<br />
Additional collections from virtually the same site<br />
(van Donselaar 1033-U, 1297-U, and 1324-U) have<br />
convinced me that the C. clausicorollata is synonymous<br />
with Simira tinctoria Aubl. More specifically,<br />
the specimen van Donselaar 1297 (U) has flowers that<br />
match Kirkbride's description of C. clausicorollata.<br />
Also, label data of all the van Donselaar's collections<br />
report that the vernacular name for this species is<br />
"tjintji-oedoe" (Sranan), which means tinged wood<br />
(Lindeman, pers. comm.).<br />
Chimarrhis decurrens Steyermark, Ceiba 3: 18. 1952.<br />
Type. Panama. Cocle: El Valle de Anton, 600 m,<br />
28 Jun 1946 (fl), Allen 3566 (holotype, F; isotypes,<br />
F[2], MO) = Allenanthus erythrocarpus Standley.<br />
Chimarrhis dioica K. Schumann & K. Krause, Bot.<br />
Jahrb. Syst. 40: 312. 1908. Type. Ecuador. Loja:<br />
Rio El Vargro, shrubs growing on oriental Andean<br />
slopes, 3400-3600 m, Oct-Nov 1906 (fl), Lehmann
186 Flora Neotropica<br />
7718 (B*, photo-B at G, GH, MO; lectotype, K) =<br />
Dioicodendron dioicum (K. Schumann & K.<br />
Krause) Steyermark.<br />
Chimarrhisferruginea (Standley) Standley, N. Amer.<br />
Fl. 32: 6. 1918. <strong>Rustia</strong>ferruginea Standley, Contr.<br />
U.S. Natl. Herb. 18: 142. 1916. Type. Panama.<br />
Col6n: Along Rio Fato, in forest or thickets, 10-<br />
100 m, Aug 1911 (fl), Pittier 4201 (holotype, US).<br />
(not Chimarrhis, not <strong>Rustia</strong>) = Bathysa (?) sp.<br />
Dwyer (1980) did not cite this species in either Chi-<br />
marrhis or <strong>Rustia</strong>. This species, in my opinion, does<br />
not belong to either of the two genera under consid-<br />
eration, and the absence of fruiting material (and the<br />
depauperate inflorescence) does not allow its place-<br />
ment with certainty. Future flowering or fruiting col-<br />
lections will hopefully clarify its generic position.<br />
Chimarrhis goudotii Baillon, Adansonia 12: 307.<br />
1879. Sickingia goudotii (Baillon) Standley, Publ.<br />
Field Columbian Mus., Bot. Ser. 7: 27. 1930. Type.<br />
Colombia. Quinindi6, 1844 (fl), Goudot s.n. (lec-<br />
totype, P, here selected; B*, photo-B at F, GH,<br />
NY) = Simira goudotii (Baillon) Steyermark.<br />
Baillon (1879) described a number of species of<br />
Simira under Chimarrhis, which were treated by him<br />
as closely related genera. Chimarrhis is easily recognizable<br />
by its axillary inflorescences and peltate seeds<br />
with concentric wing, while Simira has terminal inflorescences<br />
and horizontal seeds with a lateral orbicular<br />
wing. He treated Sickingia [= Simira] as a section<br />
of Chimarrhis. Chimarrhis goudotii was described by<br />
Baillon (1879) as "C. (Sickingia) goudotii" using<br />
specimens collected by Goudot in Colombia. One<br />
specimen preserved at Paris (P) has two labels:<br />
"Sickingia - Chimarrhis goudotii Baill [Baillon handwriting].<br />
- Quindiu [Goudot handwriting]- NC"e Grenade<br />
- M. Justin Goudot 1844," and the other (in<br />
Baillon handwriting) bears a description of the species,<br />
which declares it related to Simira longifolia, and<br />
a sketch of the flower. I select the specimen preserved<br />
at P as the lectotype of C. goudotii (= Simira goudotii).<br />
An additional specimen of Goudot's collection was<br />
preserved at Berlin (now destroyed, photos at F, GH,<br />
NY) was labeled as "Chimarrhis (Sickingia) goudotii<br />
Baill. - Quindiu et Tolima - leg Goudot" (Baillon's<br />
handwriting). This species was correctly transferred<br />
by Steyermark (1972) to Simira.<br />
Chimarrhis<br />
longifolia (Willdenow) Baillon, Adanso-<br />
nia 12: 306. 1879. Sickingia longifolia Willdenow,<br />
Gesellsch. Naturf. Freunde Schr. 3: 445. 1801.<br />
Type. Venezuela. "Prov. Caracas," Bredemeyer<br />
s.n. (no types found) = Simira longifolia<br />
(Willdenow) Bremekamp.<br />
Baillon (1879) transferred Sickingia longiflora to<br />
Chimarrhis. Bremekamp (1954a) concluded that<br />
Sickingia and Simira are congeneric and transferred<br />
C. longifolia to Simira. Inspired by Bremekamp's<br />
conclusions, Steyermark (1972) transferred all of the<br />
species of Sickingia known to him to Simira.<br />
Chimarrhis paraensis Baillon, Adansonia 12: 308.<br />
1879. Sickingiaparaensis (Baillon) K. Schumann<br />
in Martius, Fl. Bras. 6(6): 232. 1889. Type. Bra-<br />
zil. Para: Saint-Hilaire s.n. (ex herb. Lisboa), ([P<br />
not found]; B*, photo-B at F, GH, NY; fragment<br />
of isolectotype, F [ex herbario parisiensis]) =<br />
Simiraparaensis (Baillon) Steyermark.<br />
The specimen used by Baillon in P to describe C.<br />
paraensis was not found, and the specimen once pre-<br />
served at B is now destroyed. The photos of the Ber-<br />
lin type allow enough definition to conclude that also<br />
this specimen was seen by Baillon (his handwriting<br />
is on the label). A fragment (one leaf) with the stamp<br />
"Ex Herbario Musei parisiensis" and preserved at F,<br />
is labelled "Sickingia paraensis - Para - herb.<br />
lusitanicum [Lisboa, Portugal]"; this is probably a<br />
portion of the Paris type.<br />
Chimarrhisperijaensis Steyermark, Acta Bot. Venez.<br />
8: 248. 1973. Type. Venezuela. Zulia: Sierra Perija,<br />
forest near Rio Omira-Kuna, SW de Pishikako,<br />
1440-1460 m, 28 Mar 1972 (fl), Steyermark &<br />
Dunsterville 105658 (holotype, VEN) = Bathysa<br />
perijaensis (Steyermark) Delprete (see Delprete,<br />
1996b).<br />
Chimarrhis pisoniaeformis Baillon, Adansonia 12:<br />
307. 1879. Sickingia pisoniiformis (Baillon) K.<br />
Schumann in Martius, Fl. Bras. 6(6): 225. 1889.<br />
Type. Brazil. Para(?): Saint-Hilaire s.n. [ex herb.<br />
Lisboa] (lectotype, P, here selected; photo-P at A,<br />
F, MO, NY). Ixora podocarpa J. D. Hooker in<br />
Bentham & Hooker, Gen. PI. 2: 113. 1873. Type.<br />
Brazil. Rio Negro, above Barraroa, Spruce 1994<br />
(lectotype, K, here selected; photo-K at NY). =<br />
Simira pisoniiformis (Baillon) Steyermark.<br />
The selected lectotype of C. pisoniaeformis pre-<br />
served at P has not been directly seen, but its photo<br />
(at A, F, MO) is adequate to identify it. This speci-
Systematic Treatment of the Genera Studied 187<br />
men has two labels (Baillon handwriting): one with a<br />
sketch of a flower, and the other with the listing of its<br />
morphological characters. Steyermark (1974) noted<br />
that Ixora podocarpa (as "nomen subnudum") is synonymous<br />
with Sickingia pisoniiformis. I did not see<br />
the type specimen of Ixora podocarpa preserved at<br />
K, but its photo (NY) is adequate to recognize the<br />
characteristic shape of its capsules, and its label<br />
(Spruce handwriting) reports the collection locality as<br />
given in the original description; consequently the K<br />
specimen is selected as lectotype of I. podocarpa.<br />
Chimarrhis pittieri Standley, Publ. Field Columbian<br />
Mus., Bot. Ser. 8: 53. 1930. Type. Venezuela.<br />
Distrito Federal: Caracas, 1926 (fr), Pittier 11954<br />
(holotype, US, US-frag at F, US-photo at F, G;<br />
isotypes, G, NY, VEN) = Bathysa pittieri<br />
(Standley) Steyermark. (see Delprete, 1996b).<br />
Chimarrhis sessile Rusby, in herb., ined. Basis: Bo-<br />
livia, Yungas, 1890, Bang 418 (G, GH, NY).<br />
Bang 418 was annotated and distributed by Rusby<br />
as "C. sessile," a name that remained unpublished.<br />
Bang 418 is the type of Malanea grandis Rusby<br />
(1893), which was later transferred by Rusby (1895)<br />
to Elaeagia. About its identity, Steyermark (1965)<br />
stated "Standley [1931b] placed this species under<br />
Elaeagia in his <strong>Rubiaceae</strong> of Bolivia, but it is obvi-<br />
ously not this genus. The 5-6-lobed corolla, 5-6 sta-<br />
mens, conspicuously dentate 5-6-lobed calyx, and<br />
other characters would exclude it from Elaeagia, and<br />
the multi-ovulate cells of the ovary would exclude it<br />
from Malanea, with which it has also been filed in<br />
some herbaria." This species also does not belong to<br />
Chimarrhis because of its terminal inflorescences. To<br />
place this taxon in a genus, fruiting specimens are<br />
necessary. At the present time I can only suggest<br />
placement in either Bathysa or Warszewiczia.<br />
Chimarrhis venezuelensis Standley & Steyermark,<br />
Fieldiana, Bot. 28: 162. 1930. Type. Venezuela.<br />
Lara: Slopes of mtn. below Santo Domingo and<br />
Los Quebraditos, S of Las Sabanetas, above<br />
Humocaro Bajo, 2430-2475 m, 8 Feb 1944 (fl),<br />
Steyermark 55408 (holotype, F; isotypes, GH, NY,<br />
US, VEN). = Dioicodendron dioicum (K.<br />
Schumann & K. Krause) Steyermark (see Dioicodendron,<br />
below).<br />
DIOICODENDRON<br />
7. Dioicodendron Steyermark, Bol. Soc. Venez.<br />
Cienc. Nat. 25: 24. 1963. Schumann & Krause,<br />
Engl. Bot. Jahrb. 40: 312. 1908; Standley, Publ.<br />
Field Columbian Mus., Bot. Ser. 7: 203. 1931;<br />
Standley & Steyermark, Fieldiana, Bot. 28: 570-<br />
571, fig. 121. 1953; Steyermark, Bol. Soc. Venez.<br />
Cienc. Nat. 23-28. 1963; Steyermark in Lasser &<br />
Steyermark, Fl. Venez. 9(1): 253-256, fig. 40.<br />
1974; Delprete in Harling & Andersson, Fl. Ecuador<br />
(in press). Type species. Dioicodendron<br />
dioicum (K. Schumann & K. Krause) Steyermark.<br />
Shrubs to trees, much-branched, without buttresses;<br />
bark light brown. Stipules interpetiolar, free<br />
at base, not adnate to the petioles, persistent. Leaves<br />
short-petiolate, elliptic to ovate, short-acuminate at<br />
apex; petioles not thickened at base; domatia absent.<br />
Inflorescences terminal, frondose, pyramidal, laxly<br />
paniculate, with opposite to subopposite lateral<br />
branches terminating in corymbose cymules. Flowers<br />
dioecious, 4-merous, short-pedicellate, fragrant;<br />
hypanthium of male flowers shallowly obconical; hypanthium<br />
of female flowers oblong-elliptic. Calyx<br />
extremely reduced, with 4 narrowly triangular to ovate<br />
lobes, persistent. Corolla funnelform, deeply lobed,<br />
with reflexed lobes, white to greenish white; tube<br />
extremely reduced, entirely glabrous; lobes 4; aestivation<br />
valvate with contact zone. Male flowers whitish<br />
green, fragrant, with 4(-5) stamens alternate to<br />
the petals, exserted well above the corolla; filaments<br />
attached on the upper part of the tube, slender, basally<br />
flattened, entirely glabrous; anthers oblong-elliptic,<br />
dorsifixed, dehiscing by lateral slits. Pollen tricolporate,<br />
exine thinly reticulate. Female flowers<br />
whitish green, fragrant, with exserted style; style<br />
branches protruding above the corolla only after anthesis,<br />
arcuate, with the tips remaining in contact during<br />
maturity and pollination, ovate. Ovary 2-celled,<br />
turbinate to obovate, placentation peltate on the septum;<br />
ovules many in each locule, horizontally inserted.<br />
Capsules woody, obovoid to turbinate; dehiscing<br />
septicidally, disk-loculicidal dehiscence present in old<br />
capsules. Seeds obovoid to suborbicular, 3-5-angular,<br />
with a microscopic concentric wing.<br />
Dioicodendron<br />
is a unique genus of trees in the Rubi-<br />
aceae, having dioecious tetramerous sweet-fragrant<br />
flowers, whitish green to bluish green corollas, foliose-<br />
ovate spreading stipules, cinnamoneous (light-brown)<br />
bark, and capsules containing minute ovoid seeds.
188 Flora Neotropica<br />
Dioicodendron is similar to (and often confused Semiscandent shrubs 2-4 m tall to much<br />
with) Chimarrhis in having minute fragrant flowers<br />
and peltate placentation; the previous differing from<br />
the latter in having terminal, paniculate, frondose inflorescences<br />
(Fig. 82A; vs. lateral, subterminal), flowers<br />
4-merous and dioecious (vs. 5-merous and monoecious),<br />
ovate-foliose persistent stipules (Fig. 82B,C;<br />
vs. narrowly lanceolate and readily caducous), and<br />
minute, ovoid, non-winged seeds (vs. minute, flattened,<br />
and with concentric wing). Dioicodendron has<br />
also been confused with Chimarrhis, because the latter<br />
has been erroneously reported to be dioecious or<br />
heterostylous (Wright 1262 in label data; Grisebach,<br />
1862; Urban, 1899).<br />
The flowers of Dioicodendron (Fig. 83A-F) have<br />
valvate-reduplicate aestivation. Its female flowers<br />
(Fig. 83A-D) are short-pedicellate, with spreading<br />
corolla lobes, narrowly obovoid hypanthium, and the<br />
stigmas convergent at maturity. Its male flowers (Fig.<br />
83E,F) are, in contrast, long-pedicellate, with erect<br />
corolla lobes, shallowly obconical hypanthia, filaments<br />
curved toward the center, and anthers often in<br />
contact with each other.<br />
Dioicodendron is here treated as a monotypic genus,<br />
occurring at medium-high elevations of the South<br />
American Andes.<br />
1. Dioicodendron dioicum (K. Schumann & K. Krause)<br />
Steyermark, Bol. Soc. Venez. Ci. Nat. 25: 24. 1963.<br />
Chimarrhis dioica K. Schumann & K. Krause, Bot.<br />
Jahrb. Syst. 40: 312. 1908. Type. Ecuador. Loja:<br />
Rio El Vagro, shrubs growing in E Andean slopes,<br />
3400-3600 m, Oct-Nov 1906 (fl), Lehmann 7718<br />
(lectotype, K, selected by Delprete, 1999b; photo-K<br />
at VEN; isolectotype, B*, photo-B at G, GH, MO,<br />
VEN).<br />
Figs. 3R, 4L,M, 11A, 18G,H, 82A-E, 83A-F, 84<br />
Dioicodendron cuatrecasasii Steyermark, Bol. Soc.<br />
Venez. Ci. Nat. 25: 25. 1963. Type. Colombia. Valle:<br />
Cordillera Oriental, W slopes, riverbank of Rio<br />
Digua, left side, Piedra de Moler, 900-1180 m, 19-<br />
28 Aug 1943 (fr), Cuatrecasas 15192 (holotype,<br />
VEN; isotypes, F, US).<br />
Chimarrhis venezuelensis Standley & Steyermark,<br />
Fieldiana, Bot. 28: 570, fig. 121. 1953. Type. Venezuela.<br />
Lara: Slopes of mtn. between Santo Domingo<br />
and Los Quebraditos, S of Las Sabanetas, above<br />
Humocaro Bajo, 2430-2475 m, 8 Feb 1944 (male<br />
fl), Steyermark 55408 (holotype, F; isotypes, A, NY,<br />
US, VEN).<br />
branched trees 10-12 m tall, 10-20 cm dbh (rarely<br />
single-stemmed trees 20 m tall); trees with slender<br />
trunk, much branched at crown; bark smooth, thin,<br />
grayish brown (cinnamoneous). Leafy branchlets<br />
subtetragonal, 3-5 mm thick, pale brown, densely<br />
puberulent to pubescent (to rarely hirtellous); older<br />
branches glabrate, grayish to pale brown. Stipules free<br />
at base, foliose to subfoliose, widely to narrowly ovate<br />
(rarely deltoid), rounded at apex, sparsely to densely<br />
puberulent outside, sparsely pubescent with few basal<br />
narrow colleters inside, 3-8 x 4-7 mm, light green,<br />
persistent, splitting and breaking when old. Leaves<br />
(4-)7-16(-20) x (1.5-)3-8.5 cm, L/W= 1.6:1 to 2.5:1,<br />
lanceolate, oblanceolate to narrowly ovate, attenuate<br />
to rounded at base, acute at apex, gradually tapering<br />
into an acumen 0.5-2 cm long; dark green above, pale<br />
green below, foliaceous, drying brownish green<br />
above, yellowish green below; glabrous to sparsely<br />
puberulent above; glabrous, puberulent, or pubescent<br />
(rarely hirsutulous) below; the blade plane or often<br />
rugose (depressed secondary venation); primary, secondary,<br />
and tertiary veins sparsely to densely puberulent<br />
(to appressed-pubescent), secondary veins 5-9<br />
each side, ascending-arcuate, anastomosing 2-5 mm<br />
from the margin; tertiary veins subparallel-reticulate;<br />
petioles 3-20 mm long, 1-2(-3) mm thick, flattened<br />
to concave adaxially, glabrescent, puberulent to shortpubescent;<br />
Inflorescences terminal, frondose, laxly<br />
to densely paniculate, with opposite to subopposite<br />
lateral branches subtended by leaf-like bracts, terminating<br />
in dense corymbose cymules, pyramidal to<br />
umbelliform; 16-29 cm long, basal branches (10-)12-<br />
23 cm long, lateral branches 3-7 pairs; rachis<br />
subtetragonal, rachis and branches sparsely puberulent<br />
or erect to appressed-pubescent; flowers on distal<br />
branches in dense cymules; foliose bracts subtending<br />
the secondary, tertiary and quaternary branches<br />
leaf-like, decreasing in size toward the distal portion<br />
of the inflorescence, 8-13 x 3-7 cm at basal branches,<br />
gradually reduced to 1 x 0.5 cm at most distal<br />
branches; bracteoles subtending flowers 0.7-3 x 0.2-<br />
0.4 mm, linear, sparsely puberulent. Flowers shortpedicellate<br />
(female flowers) to pedicellate (male flowers).<br />
Calyx extremely reduced, 0.5-1 x 0.7-1 mm,<br />
pale green to bluish green; lobes 4, narrowly triangular<br />
to widely ovate, 0.7-1.2 mm long. Corolla funnelform,<br />
deeply lobed with reflexed lobes, 1.5-3.5<br />
mm long, entirely glabrous, white to greenish white;<br />
tube extremely reduced, 0.3-0.5 x 0.7-1.3 mm; lobes<br />
4, 1.5-2.3 x 1-1.4 mm, oblong-ovate and acute at<br />
apex. Male flowers with pedicels 1.5-2 mm long,
Systematic Treatment of the Genera Studied 189<br />
trt<br />
ALi<br />
N~, . . i ..<br />
FIG. 82. Dioicodendron dioicum. A. Habit of frondose inflorescence (Lozano & Diaz 3 734, COL). B, C. Stipules (Ham-<br />
mered et al 5233, NY). B. Lateral view. C. Frontal view. D, E. Mature capsule (Little 8505, US). D. Lateral view. E. Top<br />
view, with circular disk and central hole.
190 Flora Neotropica<br />
AI[<br />
i ' & ,~~~~~~~~~~~~~~~~~~~~~~~~~<br />
]~~~~~7<br />
FIG. 83. Dioicodendron dioicum. A-D. Female flower (Barclay et al. 3424, US). A. Flower in anthesis. B, C. Female<br />
flower bud. B. Side view. C. Top view. D. Close up of receptive stigma. E, F. Male flower (Cuatrecasas 8665, US). E.<br />
Male flower in anthesis. F. Close up of anther (note reticulate thecae).
Systematic Treatment of the Genera Studied 191<br />
00 '70<br />
'0<br />
cleared slopes, (450-)900-3500 m, of Colombia,<br />
Venezuela, Ecuador, and Peru. Flowering specimens<br />
were collected in January, February, March, April,<br />
May, July, August, October, and November, but<br />
mainly in October. Fruiting specimens were collected<br />
in February, April, May, July, August, October, No-<br />
vember, and December.<br />
Specimens examined. COLOMBIA. BOYACA: Cerro<br />
Berlin, between Arcabuco and La Palma, 1780-1800 m, 28<br />
Mar 1973 (male fl), Cuatrecasas et al. 28672 (COL, US[2]);<br />
Cordillera Oriental, Mun. Arcabuco, 2850 m, 20 Oct 1965<br />
(male fl), Huertas & Camargo 6282 (COL); Mun. Arcabuco,<br />
La Cumbre, 2400 m, 9 Oct 1980 (male fl), Lozano & Diaz<br />
3734 (COL); Cordillera Oriental, NE of Arcabuco, 2650 m,<br />
12 Oct 1966 (male fl), Garcia-Barriga 18763 (AMES,<br />
COL); Arcabuco, 2750 m, Nov 1965 (male fl), Uribe-Uribe<br />
5468 (COL, NY). CAQUETA: Cordillera Oriental, Ruidosa,<br />
5 km NW of camp Ruidosa, 2800 m, 11 Jul 1944 (fr), Little<br />
8205 (COL, NY[2], US, VEN). CAUCA: Cerro San Pedro,<br />
2700 m, 10 Sep 1944 (fr), Hutter 1215 (US). CHoc6: Mun.<br />
San Jose de Palmar, 11 Nov 1985 (fr), Lozano et al. 4885<br />
(COL); Mun. N6vita, Curund6, right bank of Rio Ingari,<br />
400-550 m, 12 Jan 1983 (male fl), Diaz-Pedrahita 3446<br />
(COL); Cieneguita, s.d., Triana 1869 (P, US). CUNDINA-<br />
FIG. 84. Distribution ofDioicodendron dioicum (circles) MARCA: Rd. Cundinamarca-Fusagasuga, km 38-39,2300 m,<br />
and Dolichodelphys chlorocrater (stars).<br />
17 May 1972 (female fl), Barclay et al. 3424 (COL, US);<br />
rd. Bogota-Fusagasuga, below San Miguel, Quebrada Malagana,<br />
28 km SW of Bogota, 2600 m, 30 Apr 1944 (female<br />
sparsely puberulent; hypanthium shallowly obconical, fl), Fosberg 21857 (NY, VEN). HUILA: Cordillera Oriental,<br />
0.5-1 x 0.7-1 mm, glabrous to sparsely puberulent;<br />
Vertiente Occidental, between Gabinete and Andalucia,<br />
stamens 4(-5) exserted well above the 2200-2300<br />
corolla, m, 25 Mar 1940<br />
equal,<br />
(male fl), Cuatrecasas 8665<br />
attached at the superior zone of the tube, alternate to (COL, US); Mun. La Plata, Vereda Arrabal, Turbera de<br />
Paramo, 2380 m, 1 Oct 1984<br />
the lobes; filaments 2.5-3 mm<br />
(female fl), Lozano et al. 4395<br />
long, terete, glabrous,<br />
(COL). META: Ridge between Rio Tigre and Rio Papamene,<br />
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<br />
medially dorsifixed, reversed at maturity, microscopi- 1942 (fr), Fosberg 19527 (US). PUTUMAYO: Punto Buenos<br />
cally (40x) reticulate (Fig. 78F), base rounded; male Aires, Cerro Portachuelo, 2800 m, 25 Jul 1964 (male fl),<br />
flowers sometimes with a non-functional style (with- Soejarto 1086 (ECON, US). QuINDfo: Quindio, 24 Nov 1853<br />
out functional stigmas). Female flowers with pedicels (fl), Holton 410 (K, NY). VALLE: Mun. Argelia, vereda Las<br />
0.5-1 mm long, sparsely puberulent; hypanthium Brisas, 2050-2200 m, 21 Jan 1983 (male fl), Diaz-Pedrahita<br />
oblong-elliptic, 2-2.5 x 1.3-1.6 mm, glabrous to 3817 (COL); Cordillera Occidental, N of Las Brisas,<br />
sparsely puberulent; style exserted, (2.5-)3-3.5 mm<br />
Gibraltar, 2100-2200 m, 25 Oct 1946 (male fl), Cuatrecasas<br />
22516<br />
long, glabrous, terete; style branches arcuate, with the<br />
(US); Cordillera Occidental, mt. above Alto de Mira<br />
(between Tabor and Carrizales), 2100-2350 m, 23 Oct 1946<br />
tips connected at maturity, thick, widely ovate, 0.7-1<br />
(fr), Cuatrecasas 22458 ([2]US); Cordillera Occidental,<br />
mm long, stigmatic surface microscopically (40x) Serrania de Los Paraguas, Mun. El Cairo, Cerro del Ingl6s,<br />
papillose. Capsules obovoid to turbinate, acute at 2260 m, 5 Jan 1987 (male fl), Silverstone-Sopkin 3001 (US).<br />
base, apex rounded, 3-4.5 x 2-3.5 mm, dark brown WITHOUT LOCALITY: 1760-1808, Mutis 2194 (US).<br />
without lenticels, glabrous throughout; disk donut- VENEZUELA. LARA: Las Sabanetas, SW slopes of<br />
shaped, with a hole at the center (caused by the cadu- Aposentos, above Humocaro Bajo, 2500-2530 m, 3 Feb<br />
cous style); loculicidal dehiscence present in old 1944<br />
cap-<br />
(fr), Steyermark 55220 (US); SW-facing slopes at Los<br />
sules. Seeds obovoid to orbicular,<br />
Aposentos, above Las Sabanetas, 2500-2530 m, 3 Feb 1944<br />
3-5-angular,<br />
(fr), Steyermark 55223 (VEN). TRUJILLO: Dtto. Bocon6, 20<br />
0.5-0.9 mm long, testa reticulate.<br />
km E of Bocon6, Guaracamal, 09?14'N, 70011'W, 1900-<br />
2300 m, 7 Feb 1987 (female fl), Aymard et al. 5233 (NY,<br />
Distribution (Fig. 84) and ecology. Rare shrubs VEN).<br />
and trees of understory in seasonal dry forest and its ECUADOR. CARCHI: Cant6n Tulcan, above Maldoremnants,<br />
rarely in seasonal cloud forest, also on nado, Chilma, 78002'W, 005 1'N, 2000 m, 20 May 1991 (fr),
192 Flora Neotropica<br />
Palacios & Rubio 7290 (MO, QCNE, TEX); Espejo, El<br />
MOLOPANTHERA<br />
Guantal, slopes of Cerro Golondrinas, 00?51'N, 78?07'W,<br />
8.<br />
2450 m, 21 Aug 1994 (fr), Palacios & Clark 12633<br />
Molopanthera Turczaninow, Bull. Soc. Imp.<br />
(QCNE).<br />
LOJA: Parque Nacional Podocarpus, above Nudo de<br />
Naturalistes Moscou 21: 580. 1848. Forsteronia<br />
Cajanuma,<br />
trail to Mirador, 3000-3500 m, 79?10'W, 04?05'S, 10 Casaretto ex K. Schumann, in synon., in Martius,<br />
Apr 1989 (fr), Madsen 85961 (QCNE); rd. Loja-Malacotas, Fl. Bras. 6(6): 200. 1889; Turczaninow, Flora 31:<br />
Nudo de Cajanuma, 10 km S of Loja, 2670 m, 4 Oct 1944 711-712. 1848; Miquel, Linnaea 22: 804-805.<br />
(female fl), Wiggins 10880 (DS, NY, TEX, US, VEN); be- 1849; Walpers, Ann. bot. syst. 2: 799. 1852; J. D.<br />
tween Tambo Cachiyacu (La Entrada) and Nudo de Hooker in Bentham & Hooker, Gen. P. 2: 41-42.<br />
Sabanillas, 2500-3500 m, 7 Oct 1943 (female fl), Steyermark 1873; Baillon, Hist. P1. 7: 346-347, 491. 1880;<br />
54450 (NY, US, VEN). MORONA-SANTIAGO: Rd. Gualaceo- Schumann in Martius, Fl. Bras. 6(6): 198-200, fig<br />
El Lim6n, 2500 m, 15 Aug 1989 (fl), Van der Werff &<br />
109. 1889; Schumann in<br />
Gudinfo 11098<br />
Engler & Prantl, Nat.<br />
(QCNE).<br />
Pflanzenfam.<br />
PERU. AMAZONAS: Prov. Chachapoyas, 1-5 km W of<br />
4(4): 54.1891. Type species. Molop-<br />
Molinopampa, Jalca, 2400-2450 m, 18 Jul 1962<br />
anthera<br />
(fr),<br />
paniculata Turczaninow.<br />
Wurdack 1383 (K, NY, UC, US, VEN); Shanyan, Nov (fl),<br />
Jelski 376 (paratype of C. dioica [frag-W at F]).<br />
Trees; bark longitudinally fissured, brown; sapwood<br />
pale-yellow, very hard; heartwood light-brown.<br />
Local names and uses. Venezuela: quina blanca<br />
Stipules interpetiolar, free at base, adnate to petioles,<br />
(Steyermark 55220). The bitter bark is used as an alpersistent.<br />
Leaves elliptic to narrowly elliptic, acumiternative<br />
source of quinine to cure malaria (Standley nate at apex; not pellucid punctate; petioles slightly<br />
& Steyermark, 1953).<br />
thickened at base; domatia absent. Inflorescences<br />
terminal, frondose, pyramidal, laxly paniculate, sec-<br />
This species was first described by Schumann and<br />
ondary branches subtended by leaf-like bracts, tertiary<br />
Krause (1908a) as Chimarrhis dioica, based on speci- branchlets thyrsoid. Flowers zygomorphic (flower<br />
mens collected in Ecuador and Peru; of its two syn- buds and stamens curved upward), protandrous,<br />
types, I studied Lehmann 7718 preserved at Kew, sessile to short-pedicellate, fragrant; hypanthium obowhich<br />
was selected as lectotype by Delprete (1999b). void. Calyx extremely reduced, with minute deltoid<br />
A duplicate specimen was once preserved at Berlin<br />
lobes, persistent. Corolla rotate, curved upward,<br />
(probably the original type, but now destroyed), of<br />
deeply lobed, with reflexed unequal lobes, glabrous<br />
which I studied the photos (G, GH, MO).<br />
throughout, white to cream-white; tube short-cylin-<br />
Standley and Steyermark (1953) described C.<br />
drical; lobes (4-)5; aestivation imbricate. Stamens 5,<br />
venezuelensis, distinguishing it from C. dioica, the alternate to petals, unequal (the lower ones longer than<br />
previous having longer spreading hairs, larger cap- the upper ones), exserted; filaments attached near the<br />
sules, and shorter filaments.<br />
base of the tube, slender, basally flattened, glabrous;<br />
Steyermark (1963) founded the genus Dioico- anthers narrowly elliptic, dorsifixed near base, dehiscdendron,<br />
the name meaning dioecious tree, to which<br />
ing by lateral slits. Pollen tricolporate, exine densely<br />
he transferred C. dioica, and described D. cuatrereticulate.<br />
Style exserted, elongating after anther decasasii.<br />
Chimarrhis venezuelensis was reported by hiscence; style branches narrowly oblong, reflexed at<br />
Steyermark (1963) to be synonymous with D. diomaturity.<br />
Ovary 2-celled, obovoid, bilobed, with a<br />
icum. The characters (vestiture, size of capsules, and<br />
globose placenta supported by a transverse stalk atsize<br />
and shape of calyx lobes) used by Steyermark tached to the septum; ovules many per locule. Cap-<br />
(1963) to separate D. cuatrecasasii from D. dioicum<br />
sules thin-woody, strongly bilobed, the two sides subare<br />
solely a geographical variation of secondary morglobose;<br />
dehiscing loculicidally, disk septicidal<br />
phological features, and the two taxa are synonymous. dehiscence absent. Seeds with irregular outline, with<br />
The collections Holton 410 (GH, NY) annotated a concentric deeply fringed wing.<br />
by Standley as "Molopanthera colombiana Standl"<br />
(unpublished name), are D. dioicum.<br />
Molopanthera is a rare genus of trees endemic to<br />
Dioicodendron is a rare genus most commonly the forests of eastern Brazil, easily recognizable by<br />
found in the Colombian cordilleras at 450-3500 m. its minute curved flower buds (Figs. 4T, 85B), small<br />
Human pressure on these environments will likely bilobed-globose capsules, capitate placenta supported<br />
cause the species' demise.<br />
by a short stalk, seeds peltate to the placenta, umbilically<br />
attached, and concentric seed-wing with<br />
deeply fringed margins (Fig. 11B).
Systematic Treatment of the Genera Studied 193<br />
Genera with zygomorphic flowers are rare in the<br />
<strong>Rubiaceae</strong>. In Molopanthera, zygomorphy is more<br />
noticeable in its minute flower buds, which are curved<br />
upward (Fig. 85B), than in its open flowers (Fig. 85C).<br />
Nevertheless, when magnified, it is possible to appreciate<br />
the unequal length of the filaments and corolla<br />
lobes (longer in the lower portion of the corolla). The<br />
filaments remain bent upward in two couples with the<br />
two adjacent anthers connected to each other by their<br />
pointed extensions at both ends, forming two cupshaped<br />
units (Fig. 85C), the fifth filament remaining<br />
free. This particular morphology is correlated with its<br />
peculiar pollination syndrome. The anthers in couples<br />
release pollen in a spherical mass, which is thrown<br />
onto the pollinator (i.e., small bees) by the single anther<br />
(pers. obs.)<br />
Turczaninow (1848) in describing M. paniculata<br />
as having leaves glabrous throughout. The generic<br />
name is derived from the Greek gioXoWxV (molops =<br />
Trees 5-10 m tall, rarely to 30 m tall, to 80 cm<br />
dbh; bark shallowly longitudinally fissured. Leafy<br />
branchlets thin-woody, glabrous or densely ascending<br />
pubescent; lenticels sparse on older branches.<br />
Stipules free at base, broadly deltoid, long-acuminate,<br />
glabrous or densely pubescent outside, long-sericeous<br />
with few basal colleters at base inside, 3-4 x 4-5 mm,<br />
persistent, leaving a scar encircling the stem. Leaves<br />
5-16 x 2.5-5 cm, elliptic to narrowly elliptic, acute<br />
at base and at apex, sometimes short-acuminate, the<br />
acumen 0.5-1.2 cm long; stiff-foliaceous; glabrous or<br />
sparsely pubescent above, glabrous or densely golden<br />
pubescent below; primary and secondary veins glabrous<br />
or densely pubescent, prominent below, secondary<br />
veins 5-8 each side; tertiary veins subparallel-reticulate;<br />
petioles 0.7-1.5 mm long, 1-1.5 mm thick,<br />
adaxially concave, glabrous to densely pubescent;<br />
domatia absent. Inflorescences frondose, pyramidal,<br />
laxly paniculate, with opposite decussate branches,<br />
bruise or weal) and av9rTo (anteros = anther), mean- each lateral branch subtended by leaf-like bracts, tering<br />
bruised anthers, probably in allusion to the fact minating in tertiary thyrsoid branchlets, 13-18 cm<br />
that they are often fused (in couples) during anthesis. long; basal branches 9-13 cm long, lateral branches<br />
Molopanthera has been treated as incertae sedis 5-9 pairs; rachis basally terete, decussately com-<br />
(Turczaninow, 1848), tentatively placed in the Cinpressed<br />
distally, without lenticels, rachis and branches<br />
choneae (Hooker, 1873; Baillon, 1880; Schumann<br />
densely short-pubescent; flowers in tertiary thyrsoid<br />
1889, 1891), transferred to the Condamineeae (sensu branchlets single or in few-flowered units. Flowers<br />
Hooker) by Andersson and Pearsson (1991), and in- sessile to short-pedicellate; pedicels to 0.5 mm long,<br />
cluded in the "genera associated with Portlandia" by short-pubescent; hypanthium obovoid, faintly bilobed,<br />
Robbrecht (1993). Molopanthera is here treated as a 0.5-0.9 x 0.6-0.7 mm, densely pubescent; flower buds<br />
monotypic genus with the two varieties recognized by curved upward, zygomorphic. Calyx minutely<br />
Schumann (1889).<br />
cupular, short-lobed, 0.4-0.7 x 0.9-1.2 mm, pubescent;<br />
lobes deltoid, 0.3-0.5 mm long. Corolla rotate,<br />
deeply lobed, the lobes reflexed at anthesis, 3-4 mm<br />
long, glabrous throughout, white to cream-white; tube<br />
1. Molopanthera paniculata Turczaninow, Bull. Soc.<br />
short cylindrical, 0.3-0.5 x 0.5-0.6 mm; lobes (4-)5,<br />
Nat. Imperialistes Moscou 21: 581. 1848.<br />
2.5-3.5 x 0.4-1<br />
Molopmm,<br />
oblong-linear with round apex.<br />
anthera panaiculata K. Schumann, orth. var., in Stamens (4-)5, exserted well above the corolla, un-<br />
Martius, Fl. Bras. 6(6): 200, tab 109. 1889. equal, attached 0.1-0.2 mm from the base of the tube;<br />
Forsteronia panniculata Casaretto ex K. Schu- filaments 0.7-1.5 mm long, the lower ones longer than<br />
mann, in syn., in Martius, Fl. Bras. 6(6): 200. 1889. the upper ones (because of flowers buds curved up-<br />
Type. Brazil. Bahia: Serra de Jacobina, 1837, ward), slender-terete, basally flattened, glabrous<br />
Blanchet 2557 (lectotype, K, here selected; photo- throughout; anthers narrowly elliptic, 1-1.5 x 0.3-0.4<br />
K at NY; isolectotypes, F, G, MG, NY). Possible mm, dorsifixed near the base, with pointed dark exisolectotypes<br />
might be found at KW and LE. tensions at both ends, which are often fused in adja-<br />
Figs. 3T, 4N, 1 lB, 85A-D, 86 cent anthers. Pollen exine minutely foveolate-reticulate.<br />
Style exserted, elongating only after anthers<br />
Coffea floribunda Miquel, nom. superfl., Linnaea 22:<br />
dehiscence, 1-1.5 mm long, terete, glabrous; style<br />
804. 1849. Specimens cited: Brazil. Bahia: Serra de<br />
branches linear-oblong, reflexed, 0.5-1 mm long, stig-<br />
Jacobina, 1837, Blanchet 3282 (F, G-n.v., KW?,<br />
matic surface minutely papillose. Capsules short-<br />
LE?, MG, RB, U).<br />
Non Coffeafloribunda Martius, Flora 24, suppl. 2: 85. pedicellate, pedicels 0.5-1 mm long; strongly bilobed<br />
1841. Type. Brazil. Cuiaba: Morro do Rio da Casca, the two sides globose, rounded at base and at apex,<br />
collector unknown (BR?). [= Ixora densiflora 2-3.5 x 3.5-5 mm, golden to greenish yellow (to pale-<br />
(Martius) Mull. Arg.].<br />
green at maturity), without lenticel, glabrous to
194 Flora Neotropica<br />
: ~ "- I ...- '- _. _ A . -<br />
ii<br />
FIG. 85. Molopanthera paniculata var. paniculata. (A-C from Williams & Assis 6861, GH; D from Orlandi 2237, F).<br />
A. Habit of inflorescences. B. Curved flower bud. C. Flower with exserted and connected anthers. D. Mature capsule.
Systematic Treatment of the Genera Studied<br />
sparsely short-pubescent below the disk; disk gla- 1972 (fl), Spada 31 (RB[2]); Reserva Forestal Linhares, near<br />
brous, not exceeding the calyx; disk septicidal dehis- rd. 143, talhAo 602, 29 May 1972 (fl), Lino 75 (RB). BAHIA:<br />
cence absent. Seeds 0.8-3 x 1-2.2 mm, with concen-<br />
Mun. Sta. Cruz de Cabralia, Pau-Brasil Ecological Station,<br />
ca. 16 km W of Porto<br />
tric<br />
Seguro, 27 Jul 1978 (fl), Mori et al.<br />
wing deeply fringed.<br />
10328 (K, NY); Bahia, Encruzilhada, rd. to Itamb6, Mata<br />
Cip6, 27 May 1968 (fl), Belem 3677 (CEPEC, NY);<br />
Turczaninow (1848) described Molopanthera Jacobina, 1 1?16'S, 40?27'W, 25 Aug 1980 (fr), Orlandi 223<br />
paniculata citing two syntypes, Blanchet 2557 and (CEPEC, F, HRB, MG, RB); Mun. Jacobina, Serra do Brite,<br />
Blanchet 3282. Curiously, one year later, Miquel 11009'S, 40?01'W, 500-600 m, 21 Aug 1984 (fl), Bautista<br />
(1849) described Coffeafloribunda based on Blanchet & Orlandi 985 (CEPEC, HRB, MG); Dto. Itaitu, Mun.<br />
3282, the syntype of M. paniculata, present at U, Jacobina, km 8 on rd. Jacobina-Capim Grosso, 27 Oct 1995<br />
which is here selected as the holotype of (fr), Jardim et al. 711 (CEPEC); Mun. Anderai, km 10 on<br />
C.floribunda.<br />
rd.<br />
Because of the difficulties in receiving material on<br />
Andarai-LengSis (BA 142), Fda. Lagoa Encantada, 22<br />
May 1989 (fl), Mattos Silva et al. 2842 (CEPEC); Eunapolis,<br />
loan from KW and LE, Blanchet 255 7 preserved at K<br />
Itabela, Itamaraju, Prado, 8 Jul 1970 (fl), Santos 920<br />
is here selected as the lectotype of M. paniculata. (CEPEC); Itamaraju, Fazenda Boa Vista, 27 Jul 1971 (fl),<br />
The specific epithet is sometimes misspelled as Monteiro 23498 (RB); Barra da Estiva, 13?38'S, 41?07'W,<br />
panniculata (Schumann 1889, 1891), but it was origi- 21 Jun 1978 (fl), Vaillant 21 (CEPEC, HRB, MG, RB).<br />
nally published as paniculata and should be main- MINAS GERAIS: Mun. Serro, 3 km from Serro, 12 May 1945<br />
tained as such.<br />
(fl), Williams & Assis 6861 (F, GH, K, NY, RB); Itaobim, 4<br />
Apr 1979 (fl), Rizzini & Mattos 1126 (RB); Mun. Aracuai,<br />
16?53'S, 41056'W, 6 Jul 1981 (fl-fr), Salim J. Filho 148<br />
(HRB, MG); Mun. Jequitinhonha, 16?33'S, 41?06'W, 8 Jul<br />
1981<br />
Key to the varieties of (fl), Salim J. Filho 159 (HRB, MG). Rio DE JANEIRO:<br />
Molopanthera paniculata<br />
Rio de Janeiro, Lagoinha, Silvestre, 16 Jun 1959 (fl-fr),<br />
1. Leaves glabrous beneath, glabrous-shiny above; Duarte 4842 (RB110068) (F[2], RB).<br />
stipules glabrous outside; lower nodes of the<br />
inflorescences glabrous ................. 1a. var. paniculata Local name. Brazil: saco de mono (Bahia, Mori<br />
1. Leaves densely golden-pubescent beneath, coarse- et al. 10328).<br />
pubescent above; stipules pubescent outside;<br />
lower nodes of the inflorescences short-<br />
This variety is easily distinguishable from var.<br />
pubescent ......................................... Ib. var. burchellii<br />
burchellii) in having by the characters given in the<br />
key; it occurs mainly in the provinces of Bahia and<br />
northernmost Minas Gerais. Individuals of this varila.<br />
Molopanthera paniculata Turczaninow var. ety have been reported to reach 30 m in height and 80<br />
paniculata. Figs. 3T, 4N, 85A-D, 86 cm dbh (Mori et al. 10328).<br />
Distribution (Fig. 86) and ecology. Primary and<br />
disturbed seasonal forests, remnant in pastures, of<br />
Brazil, mainly in the provinces of Bahia, Espirito<br />
Santo, and northernmost Minas Gerais; commonly in<br />
clay-siliceous soils. Flowering specimens were col-<br />
lected in May, July, and August. Fruiting specimens<br />
were collected in August and October.<br />
Pollination ecology. The flowers of this variety<br />
(and probably the other variety) were reported to be<br />
cream-white, fragrant, and visited (and most likely<br />
pollinated) by small bees (label data of Bautista &<br />
Orlandi 985).<br />
Specimens examined. BRAZIL. ESPIRITO SANTO:<br />
Reserva Forestal Linhares, near rd. 161, talhao 604, 16 Jul<br />
1973 (fl), Spada 293 (RB); Reserva Forestal Linhares, near<br />
rd. 143, talhao 403, 10 Jul 1973 (fl), Spada 289 (RB);<br />
Reserva Forestal Linhares, near quadra 1, talhao 605, 2 Jul<br />
195<br />
lb. Molopanthera paniculata Turczaninow var.<br />
burchellii (J. D. Hooker) K. Schumann in Martius,<br />
Fl. Bras. 6(6): 200. 1889. Molopanthera burchellii<br />
J. D. Hooker in Bentham, G. & J. D. Hooker, Gen.<br />
P1. 2: 42. 1873. Type. Brazil. Minas Gerais: Lagoa<br />
Santa, 1863-65 (fr), Warming s.n. (B*; lectotype,<br />
F, here selected; isolectotypes, C-n.v., GH).<br />
Figs. 1 B, 85, 86<br />
Distribution (Fig. 86) and ecology. Primary and<br />
disturbed forests of southeastern Brazil, mainly in the<br />
provinces of Minas Gerais and Rio de Janeiro. Flow-<br />
ering specimens were collected in February, May, and<br />
June. Fruiting specimens were collected in May, July,<br />
and August.<br />
Specimens examined. BRAZIL. MINAS GERAIS:<br />
Caratinga Biological Station, 19?40'S, 41?50'W, 24 Aug
196 Flora Neotropica<br />
FIG. 86. Distribution of Afolopanthera paniculata var.<br />
paniculata (circles) and M. paniculata var. burchelli (tri-<br />
angles).<br />
1983 (fr), Strier 1007 (NY); Mun. Tombos, Fazenda da<br />
Cachoeira, 29 Jul 1935 (fr), Barreto 1789 (JBBH16686) (A,<br />
F); Mun. Santa Luzia, Lagoa Santa, 3 May 1934 (fl-fr),<br />
Barreto 8305 (JBBH26371) (F); Mun. Santa Luzia, Lagoa<br />
Santa, 8 May 1937 (fl), Magalhaes s.n. (IAN); Mun. Belo<br />
Horizonte, Bento Pires (Eng. Nogueira), Feb 1945 (fl), Williams<br />
5066 (F, GH); Grno Mogol, valley of Riberio creek,<br />
ca. 900 m, 24 May 1987 (fl), Pirani & Mello Silva SPF47733<br />
(NY). Rio DE JANEIRO: Morro de S. Joao, Botafogo, Jul 1914<br />
(fl), Kulhmann 1221 (RB); Horto Florestal, 9 Jun 1927 (fl),<br />
Personnel ofthe Herbario Floresta s.n. (RB); Depto. Veiase,<br />
between Arapinga and Caman, 21 Apr 1935 (fl), Kuhlmann<br />
82 (RB149083) (RB).<br />
Local name. Brazil: pesquim (Minas Gerais,<br />
Barreto 1789).<br />
This variety is distinct from the typical variety in<br />
having blades densely golden-pubescent below and<br />
coarse sparsely pubescent above, stipules externally<br />
pubescent, and lower nodes of the central rachis of<br />
inflorescences short-pubescent, correlated with its<br />
geographical distribution. The minute bent corollas<br />
and small bilobed capsules are morphologically consistent<br />
throughout the range of the species. Future<br />
collections throughout the range of this species may<br />
prove that the two varieties here recognized are not<br />
distinguishable.<br />
Schumann annotated Warming s.n. (the lectotype<br />
here selected) as "Molopanthera panniculata Turcz.<br />
var. scabrida K. Schum. et Burchell II - det.<br />
Schumann."<br />
DOLICHODELPHYS<br />
9. Dolichodelphys K. Schumann & K. Krause in E.<br />
Ule, Verh. Bot. Vereins. Prov. Brandenburg 50:<br />
102, fig. 1 (103). 1908; Standley, Publ. Field<br />
Columbian Mus., Bot. Ser. 8: 159. 1930;<br />
Steyermark in Lasser & Steyermark, Fl. Venez.<br />
9(1): 262-265, fig. 42. 1974; Delprete in Harling<br />
& Andersson, Fl. Ecuador (in press). Type species.<br />
Dolichodelphys chlorocrater K. Schumann<br />
& K. Krause.<br />
Shrubs to treelets, much branched shrubs to<br />
single-stemmed trees; bark grayish. Stipules<br />
interpetiolar, free at base, often connate to the petiole,<br />
deltoid to narrowly triangular, persistent. Leaves<br />
petiolate, lanceolate to obovate, acuminate to obtuse,<br />
mucronate or acuminate at apex; petioles thickened<br />
at base; domatia absent. Inflorescences terminal, solitary,<br />
pyramidal, openly paniculate; lateral branches<br />
opposite to subopposite, decussate. Flowers protandrous,<br />
extremely variable in size within the same inflorescence,<br />
first flowers to 3 cm long, gradually<br />
smaller toward the end of flowering season. Calyx<br />
cupular, lobes ovate to lanceolate, persistent. Corolla<br />
campanulate (in earlier, big flowers) to urceolate (in<br />
laster, smaller flowers), membranous, cream-white to<br />
yellowish green; tube cylindrical to narrowly funnelform;<br />
lobes 5(-6); aestivation narrowly imbricate.<br />
Stamens 5(-6), included, equal or subequal; anthers<br />
cylindrical, narrowly oblong, green, dorsifixed near<br />
the base, base sagittate, dehiscing by longitudinal slits.<br />
Pollen tricolporate, exine openly reticulate. Style included;<br />
style branches membranous, reflexed. Ovary<br />
2-celled, placentation axile, turbinate to narrowly<br />
obconical, glabrous, with many ovules in each locule<br />
horizontally inserted. Capsules woody, long, narrowly<br />
cylindrical, pluricostate; dehiscing septicidally,<br />
the septa detaching and expanding. Seeds ovoid,<br />
rhomboid to trapezoid, not winged, exotesta reticulate.
Systematic Treatment of the Genera Studied 197<br />
Dolichodelphys is a rare monotypic genus endemic<br />
to northwestern South America. Its name is derived<br />
from the Greek Dolicho- (= long) and delphys (=<br />
ovary), referring to the slender elongated capsules<br />
typical of this genus. Dolichodelphys is readily distinguishable<br />
by its inflorescences with the first internal<br />
flowers campanulate and 1-3 cm long, and gradually<br />
smaller on outer branchlets of inflorescence<br />
(down to 3-5 mm long), its yellowish green to creamwhite<br />
thinly membranous flowers, and its long-slender<br />
multicostate capsules (see Fig. 87A-G). This<br />
threatened species is mainly encountered in the Cordilleras<br />
of Colombia, and few rare collections are<br />
available from other northwestern South American<br />
countries.<br />
Schumann and Krause (1908c) described this genus<br />
from plants with only immature capsules and<br />
flower buds. They assigned it to the "Cinchonoideae-<br />
Gardenieae group" and related to Retiniphyllum,<br />
Synisoon [= Retiniphyllum], and Tocoyena, because<br />
of its two-parted ovaries and linear calyx lobes.<br />
Steyermark (1974) did not cite this genus in Flora de<br />
Venezuela, but treated the specimens of<br />
Dolichodelphys as <strong>Rustia</strong> longifolia. Robbrecht (1988,<br />
1993) maintained Dolichodelphys in the Gardenieae,<br />
but in his survey of the tribe (Robbrecht & Puff, 1986)<br />
lar, acuminate, glabrous outside and inside, 13-30 x<br />
8-15 mm, older stipules sometimes splitting into two<br />
parts, giving the impression of being bilobed. Leaves<br />
31-62 x 10-30 cm, L/W 2:1 to 3:1, lanceolate to oblanceolate<br />
to obovate, acute at base, acuminate to obtuse<br />
and mucronate at apex, sometimes tapering to an<br />
acuminate apex up to 3 cm long; dark green above,<br />
pale green below, semi-coriaceous; drying brown to<br />
olive-green, semi-coriaceous to stiff-chartaceous; glabrous<br />
above and below; primary and secondary veins<br />
glabrous (hirtellous in Fernandez & Oyos 182), secondary<br />
veins 17-25 each side, tertiary veins starting<br />
subparallel and reticulate in the middle; petioles 10-<br />
15 mm long, 3-5 mm thick, terete to costate, sometimes<br />
adaxially flattened. Inflorescences openly paniculate<br />
with opposite decussate branches with terminal<br />
cymules of 10-25 flowers; inflorescence (15-)25-65<br />
cm long, basal branches 15-28 cm long, apical<br />
branches 4-7 cm long, lateral branches 3-4 pairs,<br />
basal portion of axis not branched (4-)15-25 cm long;<br />
rachis and branches laterally compressed, 4-5 costate,<br />
glabrous (rarely short hirtellous); bracts subtending<br />
basal branches foliose, to 7 x 3 cm, upper bracts 10-<br />
20 x 3-6 mm, lanceolate to linear; bracteoles subtending<br />
flowers 1-2 mm x 1-2 mm, ovate to linear. Flowers<br />
pedicellate, pedicels 1-11 mm long, glabrous (but<br />
it was not mentioned. Its terminal, paniculate inflo- hirtellous in Fernandez & Oyos 182); hypanthium<br />
rescences, seed coat morphology, bilocular ovary, and obconical, 1-5 x 1-2 mm, glabrous. Calyx cupular,<br />
T-shaped placenta resemble those of Raritebe 0.5-2 x 2-6 mm, glabrous; lobes ovate to narrowly<br />
(Isertieae s.l.). Nevertheless, Dolichodelphys is ten- lanceolate (margins entire to fringed), 1-6 x 1-4 mm,<br />
tatively included in the <strong>Rondeletieae</strong> s.l. (Delprete, ciliolate. Corolla size and shape extremely variable,<br />
1996d), although future investigations may result in first largest flowers opening, 1.5-3 cm long, campanuits<br />
placement elsewhere.<br />
late; later flowers gradually smaller, (1-)6-8 mm long,<br />
tubular to urceolate, the smallest ones lacking sexual<br />
structures; tube cylindrical to narrowly funnelform,<br />
1-20 mm x 1-6 mm, glabrous outside and inside,<br />
1. Dolichodelphys chlorocrater K. Schumann & K. whitish cream to yellowish green to pale-green; lobes<br />
Krause, Verh. Bot. Vereins. Prov. Brandenburg 5(-6), 1-4 x 1-4 mm, triangular to ovate, sometimes<br />
50: 102. 1908. Type. Peru. Loreto: Cerro de<br />
mucronate, glabrous outside and inside, margins en-<br />
Escaler, 1200 m, Jan 1903 (fl), Ule 6774 (B*, tire to fringed, cream-white to yellow (rarely greenphoto-B<br />
at F; lectotype, K, selected by Delprete, ish). Stamens 5(-6), included, attached V2 to '/io from<br />
1999b; isolectotypes, frag-F, G[2], frag-IAN). the base of the tube; filaments 1-7 mm long, flattened,<br />
Figs. 3S, 12A,B, 19A,B, 84, 87A-G<br />
glabrous; anthers cylindrical, narrowly oblong, green,<br />
<strong>Rustia</strong> longifolia Standley, Publ. Field Columbian Mus., 4-7 x 1-2 mm, base sagittate, dehiscing by longitu-<br />
Bot. Ser. 8: 159. 1930. Type. Colombia. Caqueta: dinal slits. Pollen exine reticulate, with denser reticu-<br />
Between Santa Marta and Marsella, Aug 1926 (fr), lation at the<br />
Woronow & Jusepczuk 6426<br />
poles. Style included, 5-23 mm long,<br />
(holotype, LE-n.v.;<br />
glabrous; style branches ovate to obovate, membraisotypes,<br />
F, LE-n.v.).<br />
nous, reversed, entire to fringed, ca. 3 mm long, gla-<br />
Shrubs to treelets to 8(-10) m tall, to 10 cm dbh,<br />
much branched, rarely a single-stemmed treelet; bark<br />
grayish-whitish. Leafy branchlets glabrous to golden<br />
pubescent, slender, tetragonous to costate, grayish;<br />
lenticels absent. Stipules deltoid to narrowly triangubrous,<br />
stigmatic surface smooth. Capsules narrowly<br />
cylindrical, pluri-costate, apex truncate, 14-23 x 3.5-<br />
5 mm, dark brown, glabrous (hirtellous in Hernandez<br />
& Hoyos 182) throughout, without lenticels; disk truncate,<br />
not exceeding the calyx. Seeds ovoid, rhomboid to<br />
trapezoid, 0.1-0.3 long, not winged, exotesta reticulate.
198 Flora Neotropica<br />
-. \ I<br />
I '<br />
G.~~~~~ Stle H. Fruit.' ~ ~ ~<br />
?~~~~~~~~~~~~~~~~~~~~1 ?<br />
" ~ ~ ~ ~<br />
~~~~'rl ?' 'IE<br />
?~~~~~~~~~~~~~ ~ ?. ..<br />
\~~~~~~~~~~~~~~~~~~~~~~~~~?<br />
?I ..~" ?<br />
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. i<br />
~~~~. . ?<br />
~~~~~~~~=?<br />
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'i~ ' ? .. I.<br />
, ~'~~~~~~~~~~~~~~~~~~~<br />
~~~~~~~?.??<br />
~~~~~? i<br />
?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?<br />
~~~~~~~~~~~. , 'E.<br />
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G~~. S H.Fut<br />
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Systematic Treatment of the Genera Studied 199<br />
Distribution (Fig. 84) and ecology. Primary and<br />
secondary rain forests and cloud forests, to second-<br />
ary moist forests, of medium elevations, 400-1400 m,<br />
on the Andes of Venezuela, Colombia, Ecuador, and<br />
Peru. Flowering specimens were collected in January,<br />
March, June, August, October, November, and De-<br />
cember. Specimens in intermediate late flowering-<br />
early fruiting stage have been collected in July, and<br />
September. Fruiting specimens were collected in Feb-<br />
ruary, March, and August.<br />
Reproductive biology. The inflorescences of this<br />
species are laxly paniculate, and its flowering pattern<br />
is unique. The first flowers to open are large and campanulate<br />
(1.5-3 cm long). As the blooming continues<br />
on outer lateral branchlets, the flowers decrease in size<br />
(down to 3-5 mm), gradually changing shape from<br />
campanulate to short-tubular to urceolate (Fig. 87B-<br />
E). Also, as the flowers become smaller, they gradually<br />
lose their reproductive parts, first only with functional<br />
anthers (and non-functional ovary), down to the<br />
minute flowers, and the most reduced without any<br />
sexual organs and entirely sterile.<br />
Probably the flowering strategy of this species<br />
relies on the memory of its pollinators (Larry Gilbert,<br />
pers. comm.): the first few large perfect flowers have<br />
the main function to attract pollinators, the following<br />
flowers gradually diminish in corolla size (but repro-<br />
ductive organs are still functional), economizing en-<br />
ergy devoted to attraction. The pollinators, remem-<br />
bering the locality of the individuals, continue to visit<br />
the gradually less visible, but still functional flowers.<br />
The pollinators of this species have never been ob-<br />
served, but considering the size and shape of the flow-<br />
ers, they are probably butterflies.<br />
Specimens examined. COLOMBIA. ANTIOQUIA: Valley<br />
of Rio Anori, Planta Providencia, 26 km S of Saragoza,<br />
07?13'N, 75?03'W, 400-700 m, 10 Feb 1975 (fr), Denslow<br />
2680 (COL, US); Planta Providencia, 30 Sep 1984 (fl-fr),<br />
Zarucchi 3338 (MO, NY); Planta Providencia, 27 Jun 1976<br />
(fl), Shepherd 459 (COL, MO[2]); Hwy. Medellin-Bogota,<br />
Rio Samani-Rio Claro, San Luis, 600 m, 7 Jul 1981 (fl-fr),<br />
Herndndez & Hoyos 182 (COL); Mun. San Luis, quebrada<br />
La Cristalina, 06?N, 74?45'W, 770 m, 25 Aug 1987 (fr),<br />
Ramirez & Cdrdenas 179 (MO), 1017 (COL, MO), 1501<br />
(COL), 1620 (COL), 1879 (MO). CAQUETA: Florencia, Cerro<br />
de la Sardina, 500 m, 30 Mar 1940 (fl), Cuatrecasas 8926<br />
(COL, US). META: Parque Nacional La Macarena, macizo<br />
Renjifo, 1300-1900 m, 6-20 Jan 1951 (fl), Idrobo & Schultes<br />
1011 (COL, US); Parque Nacional La Macarena, cafto 20,<br />
850 m, 2 Mar 1962 (fr), Idrobo 4874 (COL[2]); Parque<br />
Nacional La Macarena, Central mtns., N ridge, 1400 m, 27<br />
Dec 1949 (fl), Philipson & Idrobo 1955 (COL[2], US[2]);<br />
Parque Nacional La Macarena, Rio Guapaya, 450 m, 26 Nov<br />
1949 (fl), Philipson et al. 1576 (COL, US). PUTUMAYO:<br />
Umbria, 00?54N, 76?10'W, 325 m, Oct 1930 (fl), Klug 1804<br />
(A, K, MO, NY, US); Rio Rumiyaco, 800 m, 9 Aug 1964<br />
(fl), Soejarto et al. 1287 (COL, GH, US[2]).<br />
VENEZUELA. CAABOBo: On steep rocky slopes above<br />
Rio San Gean, above La Toma, E of Los Tanques, S of<br />
Borburata, 1000 m, 6 Dec 1967 (fr), Steyermark 100495<br />
(G[2], VEN). WrIHour LOCALrTY: "Venezuela," Linden 1604<br />
(BR, photo-BR at VEN).<br />
ECUADOR. MORONA-SANTIAGO: Along new rd.<br />
M6ndez-Morona, km 25-30, forest remnant, 800 m, 21 Aug<br />
1989 (fl), van der Werff& Gudinio 11326 (QCNE).<br />
PERU. LORETO: Prov. Alto Amazonas, N slopes of Cerro<br />
Campanquiz, Pongo de Manseriche, right bank of Rio<br />
Marafion, 300-550 m, 19-21 Oct 1962 (fl), Wurdack 2348<br />
(US). SAN MARTIN: Prov. Lamas, Rio Curiyacu, affluent of<br />
Rio Cumbasa, 8 km above San Antonio, 450 m, 6 Nov 1937<br />
(fl), Belshaw 3593 (BH, GH, K, LL, NY, UC, US).<br />
Standley (1930b) placed <strong>Rustia</strong> longifolia Standl.<br />
close to R. splendens (= R. thibaudioides). Steyermark<br />
(1974) maintained Standley's (1930b) point of view<br />
(both authors without seeing mature flowering material),<br />
keying it and describing it under <strong>Rustia</strong>. Simpson<br />
(1976) and Andersson (1992) erroneously reported R.<br />
longifolia to be synonymous with <strong>Tresanthera</strong><br />
condamineoides. The type of R. longifolia (Woronow<br />
& Jusepczuk 6426, F) has leaves and a portion of the<br />
inflorescence that exactly matches those of D.<br />
chlorocrater, with which is here treated as synonymous.<br />
Ferndndez & Oyos 182 (Prov. Antioquia, Colombia)<br />
differs from the rest of the collections of this species<br />
in having young vegetative parts, rachis, and capsules<br />
golden-hirtellous. I do not consider this character<br />
alone sufficient so as to assign any formal taxonomic<br />
recognition to this collection.<br />
PARACHIMARRHIS<br />
10. Parachimarrhis Ducke, Arch. Jard. Bot. Rio de<br />
Janeiro 3: 253. 1922. Type species. P.breviloba<br />
Ducke.<br />
Trees; bole irregular, without buttresses; bark<br />
smooth; wood brownish yellow, hard. Stipules<br />
interpetiolar, above leaf attachment, free at base, ad-<br />
nate to the petioles, caducous, leaving a scar encir-<br />
cling the stem above the petiole attachments. Leaves<br />
elliptic to ovate to obovate, usually acuminate at apex;<br />
microscopically glandular-punctate on the abaxial<br />
side; petioles slightly thickened at base; domatia ab-<br />
sent (exceptionally a few short hairs). Inflorescences<br />
terminal corymbiform panicles, with opposite to
200 Flora Neotropica<br />
subopposite lateral branches terminating in cymules.<br />
Flowers protandrous, sessile to short-pedicellate, very<br />
fragrant; hypanthium obconical. Calyx reduced, with<br />
acute lobes, persistent; in some of the flowers one<br />
calyx lobe expanded into a small, white calycophyll.<br />
Corolla short-tubular, shortly lobed, with reflexed<br />
lobes, white; tube cylindrical, with a ring of pubescent<br />
hairs inside on the tube; lobes 5, aestivation narrowly<br />
imbricate, superficially resembling valvate aestivation.<br />
Stamens 5, alternate to the petals, exserted;<br />
filaments attached on the upper part of the tube, slender,<br />
basally flattened, with a tuft of pilose hairs at base;<br />
anthers elliptic, dorsifixed, dehiscing by lateral slits.<br />
Pollen tricolporate, exine sparsely, irregularly reticulate.<br />
Style exserted; style branches rounded to ovate,<br />
reflexed at maturity. Ovary 2-celled, placentation<br />
horizontal on the septum, obconical, with many ovules<br />
in each locule; immature fruits green and semi-fleshy<br />
when fresh. Capsules woody, bi-globular; dehiscing<br />
loculicidally, disk septicidal dehiscence absent. Seeds<br />
hemi-elliptic, dorsiventrally flattened, laterally attached<br />
in two rows each locule, margin entire, with<br />
lateral wing.<br />
Ducke (1922) treated Parachimarrhis as closely<br />
related to Chimarrhis (hence the name), stating that<br />
it differed by having deeply laciniate calyx, a shortly<br />
lobed corolla, and capsules with few seeds per locule.<br />
In Chimarrhis the inflorescences are axillary (vs.<br />
terminal in Parachimarrhis), the flowers are<br />
protogynous (vs. protandrous), the capsules are mostly<br />
obconical to turbinate (vs. biglobular), with peltate<br />
placentation (vs. horizontal) and septicidal (vs. loculicidal)<br />
dehiscence, minute seeds with deeply fringed,<br />
concentric wing irregular in outline (vs. small and with<br />
entire, lateral wing hemielliptic in outline), and testa<br />
reticulate (vs. minutely foveolate; cf. Fig. 12E).<br />
Because of the terminal inflorescences, protandrous<br />
flowers with exserted anthers, hemi-elliptic<br />
seeds (Figs. 12G, 88A-G), seed inserted horizontal<br />
on the central placenta, testa sculpturing (Fig. 12G),<br />
and the biglobular capsules (Fig. 87F), Parachimarrhis<br />
is related to Simira (Fig. 12E,F). From Simira it<br />
differs in having much smaller capsules and seeds, and<br />
yellowish wood (most species of Simira have wood<br />
with a reddish tinge when exposed to air).<br />
1. Parachimarrhis breviloba Ducke, Arch. Jard. Bot.<br />
Rio de Janeiro 3: 254. 1922. Type. Brazil. Para:<br />
Rio Tapajoz, close to Maranhaosinho waterfalls,<br />
Maria Luisa, 6 Dec 1919 (fl), Ducke s.n. (RB<br />
15687) (lectotype, RB, here selected; isotypes, B*,<br />
F, US; photo-B at F, MO, NY).<br />
Figs. 12G, 88A-G, 89<br />
Trees to 20 m tall, to 40-50 cm dbh, without buttresses;<br />
bark smooth, pale brown; wood brownish<br />
yellow, hard. Leafy branchlets 2.5-5 mm thick, terete<br />
to compressed, glabrous to puberulent,<br />
cinnamoneous; older branches glabrate, terete, brown;<br />
lenticels sparse, whitish, linear. Stipules free at base,<br />
contorted in bud, narrowly triangular, acuminate, appressed<br />
puberulent to gray-sericeous outside, glabrous<br />
with basal colleters inside, 15-27 x 2-4 mm, readily<br />
caducous, deep-green, leaving a white scar encircling<br />
the stem above petiole attachments, 0.5-1 mm wide.<br />
Leaves (10-)13-30 x 4-10 cm, L/W = 2.5:1 to 3:1,<br />
elliptic to oblong-obovate, acute-decurrent at base,<br />
acute and acuminate at apex, the narrowly triangular<br />
acumen 1.5-2.5 cm long; dark green above, pale green<br />
below, foliaceous to thinly coriaceous; drying olivegreen<br />
to pale-brown, stiff-chartaceous to coriaceous;<br />
glabrous above and below; primary, secondary and<br />
tertiary veins glabrous to very sparsely puberulent<br />
below; secondary veins 10-18 each side; tertiary veins<br />
subparallel; petioles 10-26 mm long, 1-2 mm thick,<br />
adaxially concave, glabrous; domatia absent or a few<br />
puberulent hair. Inflorescences corymbiform<br />
panicles, with opposite decussate branches, short-pedunculate,<br />
5.5-11 cm long; basal branches 2-8 cm<br />
long, lateral branches 2-5 pairs; rachis basally terete<br />
to subterete, decussately compressed distally, without<br />
lenticels, rachis and branches densely puberulent;<br />
flowers in terminal few-flowered cymules. Flowers<br />
sessile to short-pedicellate; pedicels to 1.5 mm long,<br />
puberulent; hypanthium obovoid, faintly bilobed, 0.5-<br />
0.9 x 0.6-0.7 mm, puberulent. Calyx minutely<br />
cupular, short-lobed, 1-1.5 x 1.5-2 mm, glabrous;<br />
lobes narrowly triangular, 0.6-1.7 x 0.3-0.6 mm; several<br />
flowers of the same inflorescences with one calyx<br />
lobe expanded into small calycophyll.<br />
Calycophylls 7-13 x 3-5 mm, blades elliptic to ovate,<br />
with the adaxial side covered with a thick mat of glandular<br />
hairs (0.2-0.4 mm long). Corolla tubular-campanulate,<br />
shortly lobed, the lobes reflexed at anthesis,<br />
3-4 mm long, glabrous throughout, white to<br />
cream-white; tube short cylindrical, 2-3 x 0.5-0.6<br />
mm; lobes 5, 0.5-0.7 x 0.4-0.6 mm, oblong-linear,<br />
round at apex. Stamens 5, exserted well above the<br />
corolla, equal in length, attached 1-1.5 mm from the<br />
base of the tube; filaments 4-5 mm long, slender-terete,<br />
basally flattened and short-pubescent; anthers<br />
narrowly elliptic, 1-1.5 x 0.3-0.4 mm, dorsifixed near<br />
the middle. Pollen exine irregularly reticulate. Style<br />
exserted, elongating only after anthers dehiscence, 3-
Systematic Treatment of the Genera Studied 201<br />
,'- ;<br />
,<br />
. . .o,'...<br />
~"<br />
FIG. 88. Parachimarrhis breviloba. (A-E from Fundaci6n Biol6gica Puerto Rastrojo 5052, NY, and Ducke (RB15567),<br />
RB, lectotype; F, G from McDaniel & Rimachi 20503, NY). A. Habit of inflorescence, with two mature leaves. B. Flower<br />
with exserted anthers. C. Detail of expanded calyx lobe. D. Stamen. E. Apical portion of style. F. Mature capsule. G. Seed.
202 Flora Neotropica<br />
0 200 no so. 10no<br />
oI<br />
m O<br />
a10~~~ 7~FG0 D o to ma f 0 o<br />
5 mm long, terete, glabrous; style branches clavate,<br />
reflexed, 0.5-1 mm long, stigmatic surface minutely<br />
papillose. Capsules sessile to short-pedicellate,<br />
strongly bilobed the two sides globose, rounded at<br />
base and at apex, 3-4.5 x 4.5-6 mm, without lenticels,<br />
rust-brown, glabrous throughout; disk not exceeding<br />
the calyx; disk septicidal dehiscence absent. Seeds<br />
hemi-elliptic in outline,-2.2-4.5 x 1.5-2.5 mm, with<br />
lateral wing with entire margin; seed embryo about<br />
one half the size of the seed outline.<br />
Distribution (Fig. 89) and ecology. Primary and<br />
secondary rain forests of lowland, seasonally inun-<br />
dated, Amazonian rain forest of Colombia, Ecuador,<br />
Peru, and Brazil. Flowering specimens were collected<br />
in December. One specimen at late flowering-early<br />
fruiting stage was collected in March. Fruiting speci-<br />
mens were collected in January, March, April, July,<br />
and December.<br />
Specimens examined. COLOMBIA. AMAZONAS:<br />
Parque Nacional Cahuinari, Rio Bernardo, 01?28'S,<br />
70?45'W, 7 Mar 1991 (fl-fr), Fundaci6n Biol6gica Puerto<br />
Rastrepo 5052 (NY).<br />
PERU. LORETO: Rio Mazan, above La Libertad, ca. 35<br />
km above Mazan, 150 m, 10 Jul 1976 (fr), Gentry & Revilla<br />
16645 (NY); Maynas, Dtto. Iquitos, Rio Mom6n, ca. 2 km<br />
FIG. 89. Distribution of Parachimarrhis breviloba.<br />
above Rio Nanay, 13 Mar 1976 (fr), McDaniel & Rimachi<br />
20503 (NY); Maynas, Dtto. Puchana, Rio Amazonas,<br />
Caserio de Santa Maria de Ojeal, Quebrada de Sinchicuy,<br />
90 m, 13 Apr 1993 (fr), Rimachi 10501 (NY).<br />
BRAZIL. PAA: Rio Tapajoz, vic. of Francez, 19 Dec<br />
1919 (fr), Ducke s.n. (RB 15400)(B* [photo-B at F, MO,<br />
NY], F, RB, US, paratypes). RorND6N: Rio Machado, curso<br />
inferior, Jan 1981 (fr), Goulding 1048 (MG).<br />
Local names and uses. Peru: Huacamayo micuna<br />
(Rimachi 10501), quillobordon (McDaniel & Rimachi<br />
20503). Ducke (1922) reported that this species has<br />
good wood, but no report about its use has been found.<br />
Molopanthera breviloba is easily distinguishable<br />
by its small (3-4.5 x 4.5-6 mm), bilobed capsules,<br />
and minute (2.2-4.5 x 1.5-2.5 mm) seeds that are<br />
laterally attached and hemi-elliptic in outline. It is also<br />
typical in having flowers with small (7-13 x 3-5 mm)<br />
calycophylls covered with a thick mat of glandular<br />
hairs. The function of this matt of hairs is unknown.<br />
Molopanthera breviloba is a little-known species<br />
represented by mid-canopy trees, known from a few<br />
collections from Amazonian forests that have been<br />
little collected. In the last decades a few more collec-<br />
tions have shown its wide distribution throughout the<br />
Amazon basin.
Literature Cited 203<br />
ACKNOWLEDGMENTS<br />
My personal gratitude goes to Billie Turner (TEX),<br />
who was my doctoral supervisor during my Austin<br />
years (1991-1996); not only has he assisted me with<br />
his supervision, but he also helped sponsor my travel<br />
expenses to Costa Rica in 1991 and Ecuador in 1992,<br />
and the direct expenses for SEM studies of seeds and<br />
pollen in 1994. My extreme appreciation goes to<br />
Grady Webster (DAV), the first botanical teacher who<br />
guided my first steps through the cloud forests of<br />
Ecuador, and who was the first to inspire me to study<br />
<strong>Rubiaceae</strong> (sitting on a log, at sunset, in the outdoors<br />
near Austin). Larry Gilbert (TEX) showed me the<br />
beauty and diversity of tropical forests in Corcovado<br />
National Park, Costa Rica; from him I learned the<br />
attitude of wonder and curiosity that is present inside<br />
of me. The contribution by Paul Fryxell (TEX) toward<br />
the writing of this monograph has been among the<br />
most valuable and influential, helping me to solve<br />
problems ranging from taxonomic history to typification,<br />
from nomenclatural problems to species concepts,<br />
and carefully proofreading my entire doctoral<br />
dissertation. My most sincere thanks go to Joseph<br />
Kirkbride (USDA), who read every single word of the<br />
final version of the manuscript, contributing many<br />
valuable suggestions and helping to solve nomenclatural<br />
problems. My particular thanks go to Charlotte<br />
Taylor (MO) and John Dwyer (MO), who, in fall 1990,<br />
suggested that I study <strong>Rustia</strong> or Chimarrhis; following<br />
their suggestions the original study terminated in<br />
the ongoing monographic treatment of the entire Condamineeae<br />
(sensu Hooker). I am grateful to Charlotte<br />
Taylor for reviewing the manuscript and for helpful<br />
comments. I am also grateful to Rupert Bareby (NY),<br />
who improved the Latin diagnoses of the new taxa<br />
described here and elsewhere, and for helping me<br />
solve many nomenclatural and taxonomical problems.<br />
U, UB, UC, US, USCH, and W (acronyms according<br />
to Holmgren et al., 1990).<br />
I owe final thanks to the many colleagues and<br />
friends, some of them no longer with us, who contributed<br />
to my research on the Condamineeae: Linda<br />
Albert de Escobar, Lennart Andersson, Patricia<br />
Apreza, Alberto Areces, William Burger, John Clark,<br />
Hermes Cuadros, Todd Barkman, Brian Boom,<br />
Birgitta Bremer, Joan Busquets, Daryon Cardenas,<br />
Arthur Cronquist, Miles Davis, Ted Delevoryas,<br />
Bianca Dickerson, Imre Eifert, Javier Francisco-<br />
Ortega, Alina Freire-Fierro, Javier Fuertes, Fabio<br />
Garcia Cossio, Al Gentry, Steve Ginzbarg, Doug<br />
Goldman, Mario Gomes, Vera Gomes-Klein, Favio<br />
Gonzales, Barry Hammel, Jim Henrickson, Lauritz<br />
Holm-Nielsen, Otto Huber, Robert Jansen, Jaime<br />
Jaramillo, Dennis Kearns, Michael Kiehn, David<br />
Lorence, Jim Luteyn, Tom Mabry, Mark Mayfield,<br />
June McCaskill, Tim McDowell, John Mendenhall,<br />
Philip Morat, Scott Mori, Michael Nee, David Neill,<br />
Guy Nesom, Benjamin 0llgaard, Jose Panero, Cristina<br />
Panfet-Valdez, Henrik Pedersen, Claes Persson, Luis<br />
Poveda, Alan Prather, Christian Puff, Don Pullen,<br />
Montserrat Rios, Elmar Robbrecht, Katya<br />
Romoleroux, Johan Rova, Jan Saunders de Palacios,<br />
Beryl Simpson, Neil Snow, Bertil Stahl, Stella Sylva,<br />
Armen Takhtajahn, Ivan Valdespino, Renato<br />
Valencia, Angela Verduga, Justin Williams, and Tom<br />
Zanoni.<br />
During my collecting trips in the tropical forests<br />
of South America I was hosted by several native tribes,<br />
among them the Achuare, Embera, Kofan, Kogi,<br />
Secoya, Sharanahua, Shuar, Tikuna, Tukano, and<br />
Waorani. I regard the members of these tribes as my<br />
ultimate masters, and their ancestral wisdom should<br />
be the inspiration for the morals and ethics of modem<br />
societies.<br />
Jack Neff (TEX) identified all the insects that visited<br />
the flowers of the taxa presented in this study. Jim LITERATURE CITED<br />
Luteyn (NY) was always patient in answering all the<br />
editorial questions that came up as the manuscript was<br />
reaching its final form. The illustrations were all<br />
drawn by myself during the last three years in Austin<br />
and my first year in New York. The SEM and field<br />
photographs are all taken by myself, unless otherwise<br />
indicated in figure captions.<br />
I am grateful to the directors and staff of the following<br />
institutions for making their specimens available<br />
(a total of nearly 13,000), hosting me, and helping<br />
in the field: A, AAU, B, BH, BR, C, CAS, CAY,<br />
CEPEC, CHOCO, CLEMS, CM, COL, CR, DAV,<br />
DS, F, FI, FLAS, FSU, G, GA, GB, GH, GOET, HAC,<br />
HAJB, HRB, IAN, INPA, JBGP, JEPS, K, LL, M,<br />
MA, MG, MO, NY, P, QCA, QCNE, RB, S, SP, TEX,<br />
Adams, R. J. & J. K. Morton. 1972. An improved technique<br />
for examining pollen under the scanning electron<br />
microscope. Pollen & Spores 14: 203-212.<br />
Adanson, M. 1763. <strong>Rubiaceae</strong> [as "Aparines"]. Families<br />
des Plantes 140-147.<br />
Aiello, A. 1979. A reexamination ofPortlandia (<strong>Rubiaceae</strong>)<br />
and associated taxa. J. Arnold Arb. 60: 38-123.<br />
Andersson, L. 1992. A provisional checklist of Neotropical<br />
<strong>Rubiaceae</strong>. Scripta Bot. Belg. 1: 1-200.<br />
- . 1994. <strong>Rubiaceae</strong>-Cinchoneae. In: G. Harling & L.<br />
Andersson (eds.), Flora of Ecuador 50: 3-82.<br />
& C. Persson. 1991. Circumscription of the tribe<br />
Cinchoneae (<strong>Rubiaceae</strong>)-A cladistic approach. PI. Syst.<br />
Evol. 178: 65-94.<br />
Aublet, J. B. C. F. 1775. Histoire des plantes de la Guiane<br />
Francoise. P. F. Didot, Paris.<br />
Baillon, H. E. 1879. Les ailes seminales de certainnes<br />
Rubiacees. Adansonia 12: 296-302.
204 Flora Neotropica<br />
- . 1880. Rubiac6es-Dipsacacees. Histoire naturelle des<br />
plantes 7: 257-503. Leipzig (L. Hachette et Cie), Paris.<br />
Barbosa, M. R. de V. & A. L. Peixoto. 1989. As esp6cies<br />
de Simira (<strong>Rubiaceae</strong>, <strong>Rondeletieae</strong>) da Amaz6nia<br />
brasileira. Acta Amazon. 19: 27-46.<br />
Benoist, M. R. 1920. Bathysa difformis. Bull. Mus. Hist.<br />
Nat. Paris 26: 185-186.<br />
- . 1933. Pseudochimarrhis difformis. Bois de la<br />
Guyane. Arch. Bot. 5 (Mem. 1): 264-265.<br />
Bentham, G. 1844. The botany of the voyage H. M. S. Sulphur,<br />
under the command of Captain Edward Belcher,...<br />
during the years 1836-1842. Smith, Elder & Co., London.<br />
Blackmore, S. & S. H. Barnes. 1986. Harmomeghatic<br />
mechanisms in pollen grains. Linn. Soc. Symposium Ser.<br />
12: 137-149.<br />
Boom, B. M. 1984. A revision of Isertia (Isertieae: <strong>Rubiaceae</strong>).<br />
Brittonia 36: 425-454.<br />
& M. T. V. do A. Campos. 1991. A preliminary<br />
account of the <strong>Rubiaceae</strong> of a central Amazonian terra<br />
firme forest. Bol. Mus. Paraense Emilio Goeldi, n.s., Bot.<br />
7(2): 223-247.<br />
Borhidi, A. 1982 [1983]. Studies in <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)<br />
III: The genera Rogieria and Arachnotryx. Acta<br />
Bot. Acad. Sci. Hung. 28: 65-71.<br />
--- . 1987 [1989]. Studies in <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>)<br />
X. New combinations of Central American taxa. Acta<br />
Bot. Hung. 33: 301-303.<br />
--- . 1991. Phytogeography and vegetation ecology of<br />
Cuba. English translation by A. Borhidi, J. Podani & I.<br />
K. Kecskes. Szegedi Nyomda, Szeged, Hungary.<br />
& M. Fernandez Zequeira. 1981 a [1982]. Studies<br />
in <strong>Rondeletieae</strong> (<strong>Rubiaceae</strong>) I. A new genus: Roigella.<br />
Acta Bot. Hung. 27: 309-312.<br />
--- & .- 1981b [1982]. Studies in <strong>Rondeletieae</strong><br />
(<strong>Rubiaceae</strong>) II. A new genus: Suberanthus. Acta Bot.<br />
Hung. 27: 313-316.<br />
- & M. Jarai-Koml6di. 1983 [1986]. Studies in <strong>Rondeletieae</strong><br />
(<strong>Rubiaceae</strong>), IV. A new genus: Javorkaea. Acta<br />
Bot. Hung. 29: 13-27.<br />
, & M. Moncada. 1980 [1981]. Acunaeanthus,<br />
a new genus of <strong>Rubiaceae</strong>. Acta Bot. Acad. Sci.<br />
Hung. 26: 277-287.<br />
Bremekamp, C. E. B. 1934. Notes on the <strong>Rubiaceae</strong> of<br />
Surinam. Recueil Trav. Bot. Neerl. 31: 248-308.<br />
. 1940. On Urophyllum Wall. (<strong>Rubiaceae</strong>) and its<br />
nearest allies. Recueil Trav. Bot. Neerl. 37: 171-197.<br />
-- . 1954a. The identity of Simira tinctoria Aubl. Acta<br />
Bot. Neerl. 3: 150-153.<br />
1954b. Les sousfamilles et les tribus des <strong>Rubiaceae</strong>.<br />
Rapports et communications parvenus avant le congress<br />
aux section 2, 4, 5 et 6, Huitiem Congres International<br />
de Botanique, Paris, 1954, pp. 113-114.<br />
-- . 1966. Remarks on the position, the delimitation,<br />
and the subdivision of the <strong>Rubiaceae</strong>. Acta Bot. Neerl.<br />
15: 1-33.<br />
Bremer, B. 1992. Phylogeny of the <strong>Rubiaceae</strong> (Chiococceae)<br />
based on molecular and morphological data-Useful approaches<br />
for classification and comparative ecology. Ann.<br />
Missouri Bot. Gard. 79: 380-387.<br />
. 1996. Phylogenetic studies within <strong>Rubiaceae</strong> and<br />
relationships to other families based on molecular data.<br />
Opera Bot. Belg. 7: 33-50.<br />
& O. Eriksson. 1992. Evolution of fruit characters<br />
and dispersal modes in the tropical family <strong>Rubiaceae</strong>.<br />
Biol. J. Linn. Soc. 47: 79-95.<br />
& R. K. Jansen. 1991. Comparative restriction site<br />
mapping of chloroplast DNA implies new phylogenetic<br />
relationships within the <strong>Rubiaceae</strong>. Amer. J. Bot. 78:<br />
198-213.<br />
& L. Struwe. 1992. Phylogeny of the <strong>Rubiaceae</strong><br />
and the Loganiaceae: Congruence or conflict between<br />
morphological and molecular data? Amer. J. Bot. 79:<br />
1171-1184.<br />
, K. Andreasen & D. Olsson. 1995. Subfamilial and<br />
tribal relationships in the <strong>Rubiaceae</strong> based on rbcL sequence<br />
data. Ann. Missouri Bot. Gard. 82: 383-397.<br />
Brummitt R. K. & C. E. Powell (eds.). 1992. Authors of<br />
Plant Names. Royal Botanic Gardens, Kew.<br />
Burger, W. & C. M. Taylor. 1993. <strong>Rubiaceae</strong>. In: W. Burger<br />
(ed.), Flora Costaricensis. Fieldiana, Bot. n.s. 33: 1-333.<br />
Candolle, A. P. de. 1830. <strong>Rubiaceae</strong>. Prodromus Systematis<br />
Naturalis Regni Vegetabilis 4: 341-622. Treuttel &<br />
Wiirtz, Paris.<br />
Carrasco, L., A. Jimenez & D. Visquez. 1976. Specific<br />
inhibition of translocation by tubulosine in Eukaryotic<br />
polysomes. Europ. J. Biochem. 64: 1-5.<br />
Casaretto, G. 1843. Chrysoxylon. Novarum stirpium<br />
Brasiliensium decades. Decas 7: 59.<br />
Chamisso, L. A. & D. F. L. von Schlechtendal. 1829. De<br />
Plantis in Expeditionis speculatoria Romanzoffiana<br />
observatis. Linnaea 4: 129-202.<br />
Classen-Bockhoff, R. 1996. A survey on flower-like inflorescences<br />
in the <strong>Rubiaceae</strong>. Opera Bot. Belg. 7: 329-367.<br />
Condamine, C. M. de la. 1738. Sur l'arbre du quinquina.<br />
Mem. Acad. Roy. Sci. Paris 1738: 226-243, tabs. 5-6.<br />
--- . 1745. Relation abr6eg d'un voyage fait dans<br />
l'interieur de l'Amerique m6ridionale. Paris.<br />
Correll D. S. & D. H. Correll. 1982. Flora of the Bahama<br />
Archipelago. J. Cramer, Vaduz.<br />
Darwin, S. P. 1976a. The subfamilial, tribal and subtribal<br />
nomenclature of the <strong>Rubiaceae</strong>. Taxon 25: 595-610.<br />
. 1976b. The genus Lindenia (<strong>Rubiaceae</strong>). J. Arnold<br />
Arb. 57:426-449.<br />
Davis, P. H. & V. H. Heywood. 1963. Principles of angiosperm<br />
taxonomy. Van Nostrand, New York.<br />
Dean, W. 1995. With broadax and firebrand: The destruction<br />
of the Brazilian Atlantic forest. University of California<br />
Press, Berkeley.<br />
Delprete, P. G. 1995a. Proposed circumscription of the tribes<br />
Chiococceae, Condamineeae, and Catesbaeeae using<br />
morphological characters. Second International <strong>Rubiaceae</strong><br />
Conference (13-15 September 1995, Meise, Belgium).<br />
Programme and abstracts. Scripta Bot. Belg. 11: 80.<br />
. 1995b. Three new species of <strong>Rustia</strong> from Panama<br />
and Ecuador. Novon 5: 133-139.<br />
. 1996a. Studies on pendulous-flowered species of<br />
<strong>Rustia</strong> (<strong>Rubiaceae</strong>, Condamineeae), including a new<br />
species, <strong>Rustia</strong> schunkeana. Brittonia 48: 29-34.<br />
- . 1996b. Notes on calycophyllous <strong>Rubiaceae</strong>, part I.<br />
Morphological comparison of the genera Chimarrhis,<br />
Bathysa, and Calycophyllum, with new combinations and<br />
anew species, Chimarrhis gentryana. Brittonia 48: 35-44.<br />
- . 1996c. Systematics, typification, distribution, and<br />
reproductive biology of Pinckneya bracteata (W.<br />
Bartram) Raf. (<strong>Rubiaceae</strong>). PI. Syst. Evol. 201: 243-261.<br />
- . 1996d. Evaluation of the tribes Chiococceae, Condamineeae,<br />
and Catesbaeeae (<strong>Rubiaceae</strong>) based on morphological<br />
characters. Opera Bot. Belg. 7: 165-192.<br />
. 1996e. Notes on the taxonomic position of the<br />
monotypic Brazilian genus Kerianthera (<strong>Rubiaceae</strong>).<br />
Opera Bot. Belg. 7: 271-275.<br />
. 1996f. Revision of the tribe Condamineeae (sensu<br />
Hooker, 1873) (<strong>Rubiaceae</strong>). Ph.D. dissertation, University<br />
of Texas at Austin.
Literature Cited205<br />
. 1997a. Revision and typification of Brazilian Au- Gray, A. 1859. Notes upon some <strong>Rubiaceae</strong>, collected in<br />
gusta (<strong>Rubiaceae</strong>, <strong>Rondeletieae</strong>), with ecological obser- South Sea Exploring Expedition under Captain Wilkes.<br />
vation on riverine vegetation of Cerrado and Atlantic Proc. Amer. Acad. Arts 4: 306-324.<br />
Forest. Brittonia 49: 487-497.<br />
Grisebach, A. H. R. 1861. <strong>Rubiaceae</strong>. In: Flora of the Brit-<br />
- . 1997b. Notes on calycophyllous <strong>Rubiaceae</strong>. <strong>Part</strong> ish West Indian Islands 316-351. Lovell Reeve & Co.,<br />
II. Morphological comparison of the genera Bathysa and London.<br />
Schizocalyx. Brittonia 49: 480-486.<br />
- . 1862 [1860-1862]. Plantae wrightianae, e Cuba<br />
. 1998a. A new species of <strong>Rustia</strong> (<strong>Rubiaceae</strong>) from orientali. Cambridge, Boston.<br />
Bilsa Biological Station, Esmeraldas Province, Ecuador. .1866. Phyllomelia. Catalogus plantarum cubensium<br />
Novon 8: 231-234.<br />
exibens collectionem wrightianam aliasque minores ex<br />
. 1998b. Notes on calycophyllous <strong>Rubiaceae</strong>. <strong>Part</strong> insula Cuba missas, 139-140. G. Engelmann, Lipsia.<br />
III. Systematic position of the monotypic Mexican ge- Hall6, F. 1961. Contribution a l'6tude biologique et taxononus<br />
Cosmocalyx and notes on the calycophyll develop- mique des Mussaendeae (Rubiacees) d'Afrique tropicale.<br />
ment. Brittonia 50: 309-317.<br />
Adansonia, ser. 2, 1: 266-298.<br />
- . 1998c. Notes on calycophyllous <strong>Rubiaceae</strong>. <strong>Part</strong> .1967. Etude biologique et morphologique de la<br />
IV. The monotypic Brazilian genus Blandibractea tribu des Gardeni6es (Rubiacees). Mem. O.R.S.T.O.M.<br />
Wernham is a Simira (<strong>Rubiaceae</strong>, <strong>Rondeletieae</strong>). 22: 1-146.<br />
Brittonia 50: 318-323.<br />
- , R. A. A. Oldeman & P. B. Tomlison. 1978. Tropi-<br />
. 1999a. Morphological and taxonomical compari- cal trees and forests. An architectural analysis. Springer,<br />
son of Ariadne, Mazaea, Phyllomelia, and Eosanthe New York.<br />
(<strong>Rubiaceae</strong>, <strong>Rondeletieae</strong>). Brittonia 51: in press. Hemsley, W. B. 1881. <strong>Rustia</strong> occidentalis. Biologia centrali-<br />
. 1999b. <strong>Rubiaceae</strong>-Condamineeae. In: G. Harling americana. Botany 2: 14.<br />
& L. Andersson (eds.), Flora of Ecuador (in press). Hesse, M. 1984. An exine architecture model for viscine<br />
& M. Nee. 1997. The enigmatic genus Wernhamia threads. Grana 23: 69-75.<br />
S. Moore is a synonym of Simira (<strong>Rubiaceae</strong>, Rondele- ----. 1986. Nature, form and function of pollen-connecttieae).<br />
Brittonia 49: 303-308.<br />
ing threads in angiosperms. Linn. Soc. Symp. Ser. 12:<br />
Ducke, A. 1922. Plantes Nouvelles ou peu communes de la 109-118.<br />
region amazonienne (II <strong>Part</strong>ie). Arch. Jard. Bot. Rio Hiepko, P. 1987. The collections of the Botanical Museum<br />
Janeiro 3: 253-262.<br />
Berlin-Dahlem (B) and their history. Englera 7: 219-252.<br />
. 1925. Pseudochimarrhis. Plantes Nouvelles ou peu Hogeweg, P. & J. Koek-Noorman. 1975. Wood anatomicommunes<br />
de la region amazonienne (III <strong>Part</strong>ie). Arch. cal classification using interactive character weighing.<br />
Jard. Bot. Rio Janeiro 4: 177.<br />
Acta Bot. Neerl. 24: 269-283.<br />
. 1935. Dendrosipanea. Plantes Nouvelles ou peu Holmgren, P. K., N. L. Holmgren & L. C. Barnett. 1990.<br />
communes de la region Amazonienne (VIII Serie). Arq. Index herbariorum. <strong>Part</strong> I: The herbaria of the world.<br />
Inst. Biol. Veg. 2: 69-71.<br />
Ed. 8. Regnum Veg. 120. International Association for<br />
. 1945. Chimarrhis. New forest trees and climbers Plant Taxonomy, New York Botanical Garden, Bronx.<br />
of the Brazilian Amazon. Bol. Techn. Inst. Agron. Norte, Hooker, J. D. 1868. Limnosipanea spruceana. Hook. Icon.<br />
Para 4: 26-29.<br />
PI. 11: 38. PI. 1050.<br />
Duke, J. A. & R. Vasques Martinez. 1994. Amazonian --- . 1871a. Phitopis multiflora. Hook. Icon. PI. 11: 75.<br />
ethnobotanical dictionary. PCR Press, Boca Raton, FL. PI. 1093.<br />
Dwyer, J. D. 1980. Family 179. <strong>Rubiaceae</strong>. In: Woodson --. 187 lb. Phyllacanthus grisebachianus. Hook. Icon.<br />
E. et al. (eds.), Flora of Panama, part IX. Ann. Missouri PI. 11: 76-77. PI. 1095.<br />
Bot. Gard. 67: 1-522.<br />
.1873. Ordo LXXXIV. <strong>Rubiaceae</strong>. In: G. Bentham<br />
Endlicher, S. 1838. <strong>Rubiaceae</strong>. Genera plantarum, part 7: & J. D. Hooker (eds.). Genera Plantarum ad exemplaria<br />
520-566. Fr. Beck, Wien.<br />
imprimis in herbariis kewensibus servata definita, 2: 7-<br />
Esau, K. 1965. Plant anatomy. Ed. 2. John Wiley, New York. 151. Lovell Reeve & Co., London.<br />
Estrella, E. 1991. La Naturaleza Americana y las Expedi- Howard, R. A. & G. R. Proctor. 1958. Acrosynanthus<br />
ci6nes Botinicas Espafiolas del Siglo XVIII. In: I. C. jamaicense. J. Arnold Arb. 39: 101-102.<br />
Cevallos (ed.), Flora Huayaquilensis, La Espedici6n Hutchinson, J. 1910. Pogonopus febrifugus. Kew. Bull.<br />
Botanicas de Juan Tafalla a la Real Audiencia de Quito, 1910:200-201.<br />
1799-1808. Abya-Yala Editing, Quito, Ecuador. ----. 1926. The families of flowering plants. I. Dicotyle-<br />
Fagerlind, F. 1937. Embryologische, zytologische und dons. Arranged according to a new system based on their<br />
bestaubungs-experimentelle Studien in der Familie Ru- probable phylogeny. Macmillan, London.<br />
biaceae nebst Bemerkungen fiber einige Polyploiditats- ~--- & J. M. Dalziel. 1931. <strong>Rubiaceae</strong>. Flora of West<br />
probleme. Acta Horti Berg. 11: 195-470.<br />
Tropical Africa 2(1): 65-137. Whitefriars Press, London.<br />
Garcia Kirkbride, M. C. 1979. Review of the neotropical Jacquin, N.J. 1763. Selectarum stirpium americanum historia,<br />
Isertieae (<strong>Rubiaceae</strong>). Brittonia 31: 313-332.<br />
in qua ad linnaeanum systema determinatae descripti-<br />
. 1981. Duas novas tribos de <strong>Rubiaceae</strong> neotropicais. aeque situntur plantae illiae, quas in insula Martinica,<br />
Revista Brasil. Bot. 4: 119-123.<br />
Jamaica, Domingo. Ex officina Krausiana, Wien.<br />
.1982. A preliminary Phylogeny for the neotropical - . 1797. Macrocnemum speciosus. Plantarum rariorum<br />
<strong>Rubiaceae</strong>. PI. Syst. Evol. 141: 115-121.<br />
horti caesarei schoenbrunnensis 1: 19. C. F. Wappler, Wien.<br />
Germano Filho, P. 1993. Estudo taxon6mico do genero Jansen, M. E. 1979. A revision of the genus Kajewskiella<br />
Bathysa (<strong>Rubiaceae</strong>, <strong>Rondeletieae</strong>) no Brasil. Master's the- (<strong>Rubiaceae</strong>). Blumea 25: 283-294.<br />
sis, Universidade Federal do Rio de Janeiro, Rio de Janeiro. Jussieu, A. L. de. 1789. <strong>Rubiaceae</strong>. Genera Plantarum se-<br />
Goudot, J. 1909. <strong>Rustia</strong> mosenii. Bull. Soc. Bot. France cundum ordines naturales disposita, juxta methodum in<br />
56, mem. 3: 337.<br />
horto regio parisiensis exaratum, 196-210. Herissant &<br />
- . 1828. Cinchona muzonensis. Phil. Mag. n.s. 3: 132. Barrois, Paris.
206 Flora Neotropica<br />
--- . 1807. Septieme memoire sur les caracteres generaux<br />
des families, tires des graines, et confirm6 ou rectifies<br />
par les observations de Gaertner. Ann. Mus. Natl. Hist.<br />
Nat. 10: 307-332.<br />
- . 1820. Sur la famille des plantes Rubiacees. Mem.<br />
Mus. Hist. Nat. 6: 365-410.<br />
Karsten, H. 1858-1869. Florae Columbiae terrarumque<br />
adiacentium specimina selecta in peregrinatione duodecim<br />
annorum observata delineavit et descripsit H. Karsten.<br />
2 volumes. Ferdinandi Duemmleri successores, Berlin.<br />
. 1887. Bentham-Hooker's "Genera Plantarum" un<br />
Florae Columbiae specimina selecta. Bot. Jahrb. Syst.<br />
8: 354-360.<br />
Kiehn, M. 1995. Chromosome survey of the <strong>Rubiaceae</strong>.<br />
Ann. Missouri Bot. Gard. 82: 398-408.<br />
Kirkbride, J. H. 1985a. A revision of the genus Capirona.<br />
Acta Amazon. 15: 47-60.<br />
Amat. Bonpland et Alex. de Humboldt. [sumptibus<br />
librariae graeco-latini-germanicae] Paris [1815-1825].<br />
Lanjou, J. & F. A. Stafleu. 1954. Index herbariorum. <strong>Part</strong><br />
II. Collectors, A-D. Regnum Veg. 2: 1-174.<br />
Leppik, E. E. 1956. The form and function of numeral patterns<br />
in flowers. Amer. J. Bot. 43: 445-455.<br />
Lersten, N. R. 1974a. Colleter morphology in Pavetta,<br />
Neorosea and Tricalysia (<strong>Rubiaceae</strong>) and its relationships<br />
to the bacterial leaf nodule symbiosis. Bot. J. Linn.<br />
Soc. 69: 125-136.<br />
-. 1974b. Morphology and distribution ofcolleters and<br />
crystalsin relation to taxonomy and bacterial leaf nodule<br />
symbiosis of Psychotria (<strong>Rubiaceae</strong>). Amer. J. Bot.<br />
61: 973-981.<br />
--- . 1975. Colleter types in <strong>Rubiaceae</strong>, especially in<br />
relation to the bacterial leaf nodule symbiosis. Bot. J.<br />
Linn. Soc. 71: 311-319.<br />
- . 1985b. Manipulus rubiacearum IV. Kerianthera<br />
(<strong>Rubiaceae</strong>), a new genus from Amazonian Brazil.<br />
Brittonia 37: 109-116.<br />
.- 1997a. Manipulus rubiacearum VII. BioLlania,<br />
Edici6n Esp. 6: 393-406.<br />
. 1997b. Manipulus rubiacearum VI. Brittonia 49:<br />
354-379.<br />
Klotzsch, J. F. 1846. <strong>Rustia</strong>. In: F. G. Hayne, Getreue<br />
Darstellung und Beschreibung der Arzneykunde<br />
gebrauchlichen Gewachse 14: tabs. 14, 15 [555].<br />
- . 1853. Einige neue Gattungen der Rubiaceen.<br />
Monatsber. Koenigl. Preuss. Acad. Wiss. 1853: 494-502.<br />
Koek-Noorman, J. 1969a. Contribution to the wood<br />
anatomy of the South American <strong>Rubiaceae</strong>. I. Acta Bot.<br />
Neerl. 18: 108-123.<br />
---- . 1969b. Contribution to the wood anatomy of the<br />
South American <strong>Rubiaceae</strong>. II. Acta Bot. Neerl. 18:<br />
377-395.<br />
~--- . 1970. Contribution to the wood anatomy of the<br />
Cinchoneae, Coptosapelteae and Naucleeae (<strong>Rubiaceae</strong>).<br />
Acta Bot. Neerl. 19: 154-164.<br />
.1972. The wood anatomy of the Gardenieae, Ixoreae<br />
and Mussaendeae (<strong>Rubiaceae</strong>). Acta Bot. Neerl. 21:<br />
301-320.<br />
. 1976. Juvenile characters in the wood of certain<br />
<strong>Rubiaceae</strong> with special reference to Rubia fruticosa.<br />
IAWA-Bull. 3: 38-42.<br />
-- . 1977. Systematische Holtzanatomie einer Rubiaceen.<br />
Ber. Deutsch. Bot. Ges. 90: 183-190.<br />
---- . 1980. Wood anatomy and classification of<br />
Henriquezia Spruce, Platycarpum Hum. et Bonpl. and<br />
Gleasonia Standl. Acta Bot. Neerl. 29: 117-126.<br />
& P. Hogeweg. 1974. The wood anatomy of<br />
Vanguerieae, Cinchoneae, Condamineae and <strong>Rondeletieae</strong><br />
(<strong>Rubiaceae</strong>). Acta Bot. Neerl. 23: 627-653.<br />
& C. Puff. 1983. The wood anatomy of <strong>Rubiaceae</strong><br />
tribes Anthopermeae and Paederieae. PI. Syst. Evol. 147:<br />
17-45.<br />
Krause K. 1915. <strong>Rubiaceae</strong>. In: A. Engler & K. Prantl (eds.).<br />
Die Natiirlichen Pflanzenfamilien. Nachtrage [IV] zu den<br />
Teilen II-IV: 290-301. Engelmann, Leipzig.<br />
Krug, J. C. & I. Urban. Mazaea. 1897a. Ueber einigen<br />
Rubiaceen-Gattungen. Ber. Deutsche Bot. Ges. 15: 265-<br />
266, t. 9, fig. 10-21.<br />
& - . 1897b. Neomazaea. Ueber einigen<br />
Rubiaceen-Gattungen. Ber. Deutsche Bot. Ges. 15: 542.<br />
Kunth, C. S. 1818, 1820. Macrocnemum. In: F. H. A.<br />
Humboldt, A. J. de Bonpland & C. S. Kunth, Nova genera<br />
et species plantarum, quas in peregrinatione orbis<br />
novis collegerunt, descripserunt, partim adumbraverunt<br />
Linden, J. J. & J. E. Planchon. 1854. Pinckneya ionantha.<br />
Bot. Zeit. 365.<br />
Linnaeus, C. 1753. Species plantarum. Laurentii Salvii,<br />
Stockholm.<br />
Liogier, A. L. 1963. <strong>Rubiaceae</strong>. Flora de Cuba 5: 13-146.<br />
Industria Graficas Diario-Dia, Palencia de Castlla, Spain.<br />
. 1995. <strong>Rubiaceae</strong>. La Flora de la Espafiola. 7:<br />
207-439.<br />
Lorence, D. H. 1991. New species and combinations in<br />
Mexican and Central American Rondeletia (<strong>Rubiaceae</strong>).<br />
Novon 1: 135-157.<br />
- & J. D. Dwyer. 1988. A revision of Deppea (<strong>Rubiaceae</strong>).<br />
Allertonia 4: 389-436.<br />
Ma, W. W. 1991. Bioactive constituents of Pogonopus<br />
speciosus (<strong>Rubiaceae</strong>), Persea major (Lauraceae), and<br />
Endlicheria dysodantha (Lauraceae). Ph.D. dissertation.<br />
Dissertation Abstract International, Vol. 53/05-B.<br />
,J. E. Anderson, A. T. McKenzie, S. R. Byrn, J.<br />
L. McLaughlin, and M. S. Hudson. 1990. Tubulosine:<br />
An antitumor constituent ofPogonopus speciosus. J. Nat.<br />
Prod. (Lloydia) 53: 1009-1014.<br />
Maddison W. P. & D. R. Maddison. 1992. MacClade:<br />
Analysis ofphylogeny and character evolution. Version<br />
3.0. Sinauer Associates, Sunderland, Massachusetts.<br />
Marques-Souza, A. C., M. L. Absy, I. P. de A. Miranda<br />
& H. E. C. Kfichmeister. 1993. Caracteristica de flores,<br />
nectar y visitantes de Kerianthera preclara (<strong>Rubiaceae</strong>).<br />
Rev. Biol. Trop. 41: 483-489.<br />
Martius, C. F. P. de. 1837 [-1841]. Herbarium Florae<br />
Brasiliensi. Plantae brasilienses exsiccatae, quas<br />
denominatas, partim diagnosi aut observationibus<br />
instructas botanophilis offert Dr. C. Fr. Ph. de Martius.<br />
[Publisher unknown] Miinchen.<br />
McDowell, T. 1996. Exostema (<strong>Rubiaceae</strong>): Taxonomic<br />
history, nomenclature, position and subgeneric classification.<br />
Opera Bot. Belg. 7: 277-296.<br />
Merrill E. D. & L. M. Perry. 1947. Kajewskiella, a new<br />
rubiaceous genus from the Solomon Islands. J. Arnold<br />
Arb. 28: 331-332.<br />
Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the dicotyledons.<br />
Clarendon Press, Oxford.<br />
Michaux, F. A. 1817. Pinckneya. The North American sylva<br />
1: 180-181, pl. 49. Thomas Dobson, Philadelphia.<br />
Miquel, F. A. W. 1849. Coffeafloribunda. Linnaea 22: 804.<br />
Moore, S. 1922. Wernhamia. In: Herzog, T., Die von Dr.<br />
Th. Herzog auf seiner zweiten Reise durch Bolivien in<br />
den Jahren 1910 und 1911 gesammelten Pflanzen.<br />
Medel. Rijks. Herb. Leiden 46: 22-23.<br />
Muller, J. 1979. Form and function in angiosperm pollen<br />
grains. Ann. Missouri Bot. Gard. 66: 569-632.
Literature Cited 207<br />
Orsted, A. S. 1852. Macrocnemum exsertus. Vidensk. Robbrecht, E. 1988. Tropical woody <strong>Rubiaceae</strong>. Charac-<br />
Meddel. Naturhist. Foren. Kjobenhavn 2-4: 45. (reimpr. teristic features and progressions. Contributions to a new<br />
in Centralamerikas Rubiaceer. K0benhavn. 1853). subfamilial classification. Opera Bot. Belg. 1: 1-271.<br />
. 1863. Pogonopus exsertus. L'Amerique centrale 17, - . 1993 [1994]. Supplement to the 1988 outline of the<br />
Tab. 13. K0benhavn.<br />
classification of the <strong>Rubiaceae</strong>. Index to genera. In: E.<br />
Pacini, E. 1986. An approach to armomegathy. In: M. Cresti Robbrecht (ed.), Advances in <strong>Rubiaceae</strong> macro-<br />
& R. Dallai (eds.), Biology of reproduction and cell systematics. Opera Bot. Belg. 6: 173-196.<br />
motility in plants and animals. Siena University, Siena. & D. M. Bridson. 1993 [1994]. Nomenclatural notes<br />
. 1990. Harmomegathic characters of Pteridophyta on three Rubiaceeae genera. In: E. Robbrecht (ed.),<br />
spores and Spermatophyta pollen. PI. Syst. Evol. Suppl. Advances in <strong>Rubiaceae</strong> macrosystematics. Opera Bot.<br />
5: 53-69.<br />
Belg. 6: 197-200.<br />
Parrotta, J., J. K. Francis & R. R. de Almeida. 1995. & C. Puff. 1986. A survey of the Gardenieae and<br />
Trees of the Tapaj6s-A photographic field guide. Gen. related tribes (<strong>Rubiaceae</strong>). Bot. Jahrb. Syst. 108: 63-138.<br />
Tech. Rep. IITF - 1. U.S.D.A., Forest Service. International ---, --- & E. Smets. 1996. Second International<br />
Institute of Tropical Forestry, Rio Piedras, Porto Rico. <strong>Rubiaceae</strong> Conference. Proceedings. Opera Bot. Belg.<br />
Peixoto, A.L. 1981. Novas especies de Simira Aublet (Ru- 7: 1-432.<br />
biaceae) do norte do Espirito Santo. Bol. Mus. Bot. Rogers, G. K. 1987. The genera of the Cinchonoideae<br />
Munic. 44: 1-7.<br />
(<strong>Rubiaceae</strong>) in the southeastern United States. J. Arnold<br />
. 1982. As especies de Simira (<strong>Rubiaceae</strong>, Rondele- Arb. 68: 137-183.<br />
tieae) do Norte do Estado do Espirito Santo. Arq. Univ. Roig y Mesa, J. T. 1928. Diccionario botanico de nombres<br />
Fed. Rural Rio de Janeiro 5: 115-128.<br />
vulgares cubanos. [Boletin Estaci6n Experimental<br />
Planchon, J. E. 1849. Rogiera amoena. Fl. Serres Jard. Eur. Agron6mica, Santiago de las Vegas 54] Rambla, Bouza<br />
5: t. 442 (a-c).<br />
y Ca., La Habana.<br />
Pohl, J. E. 1828-1829. Augusta. Plantarum Brasiliae icones Rova, J. H. E., L. Andersson, P. G. Delprete & V. A.<br />
et descriptiones hactenus ineditae 2: 1-8, tabs. 101-105. Albert. 1997. Macrophylogeny of the <strong>Rubiaceae</strong> from<br />
[Publisher unknown] Wien.<br />
trnL-F sequence data. Abstract of the AIBS meetings,<br />
Poiret, J. L. M. 1811. Chimarrhis. In: J. B. A. P. M. de Canada. Amer. J. Bot. 84 (supplement): 227.<br />
Lamark, Encyclopedie methodique. Botanique. Suppl. Ruiz L6pez, H. 1778. Descripcion del Arbol de la Quina,<br />
2: 231. H. Agasse, Paris.<br />
hecha en el 1737 por Mr. de La Condamine de la Real<br />
Poiteau, P. A. 1804. Stevensia, nouveau genre de la famille Academia de Sciencias de Paris, entre cuyas memorias<br />
des Rubiacees. Ann. Mus. Natl. Hist. Nat. 4: 235-236, pertenecientes al afio 1738 se halla en la pagina 226.<br />
pi. 60.<br />
Traducida del Frances al Castellano, y aumentada con<br />
Prance, G. T. 1976. Inventario forestal de um hectare de algunas notas relativas a los arboles de Quina, nuevamata<br />
de terra firme km 30 de Estrada Manaus- mente descubiertos en Santa Fe de Bogota, Capital del<br />
Itacoatiara. Acta Amazon. 6: 9-35.<br />
Nuevo Reyno de Granada en la America Meridional.<br />
Presi, C. B. 1845. Bathysa. Abh. Konigl. B6hm. Ges. Wiss. [Publisher unknown] Madrid.<br />
3: 514- 515. (reimpr. Bot. Bemerk. 84. 1844 [1845]).<br />
& J. A. Pav6n y Jiminez. 1799. <strong>Rubiaceae</strong>. Flora<br />
Puff, C., A. Igersheim & U. Rohrhofer. 1993. peruviana et chilensis, sive descriptiones et icones plantarum<br />
Pseudomussaenda and Schizomussaenda (<strong>Rubiaceae</strong>): peruvianarum et chilensium, secundum systema linneanum<br />
Close allies of Mussaenda. Bull. Nat. Plantentuin. Belg. digestae, cum characteribus plurimum generum evulga-<br />
62: 35-68.<br />
torum reformatis. Vol. 2. Gabrielis de Sancha, Madrid.<br />
Radford, A. E., W. C. Dickison, J. R. Massey & C. R. Rusby, H. H. 1893. Malanea grandis. On the collections of<br />
Bell. 1974. Vascular plant systematics. Harper & Row, Mr. Miguel Bang in Bolivia. Mem. Torrey Bot. Club 3:<br />
New York.<br />
43-44.<br />
Raffauf, R. B. 1970. A handbook of alkaloids and alkaloid- . 1895. Elaeagia grandis. On the collections of Mr.<br />
containing plants. John Wiley, New York.<br />
Miguel Bang in Bolivia-<strong>Part</strong> II. Mem. Torrey Bot. Club<br />
Rafinesque, C. S. 1820. Tableau analytique des ordres 4: 208.<br />
naturels, families naturelles et genres, de la classe . 1896. Condaminea. On the collections of Mr.<br />
Endogynie, sous classe Corisantherie. Ann. Gen. Sci. Miguel Bang in Bolivia-<strong>Part</strong> III. Mem. Torrey Bot.<br />
Phys. 6: 76-89.<br />
Club 6: 45-46.<br />
Record, S. J. & R. W. Hess. 1943. Timbers of the New - . 1927. Flexanthera. Mulford exploration of the<br />
World. Yale University Press, New Haven.<br />
Amazon Valley. Mem. New York Bot. Gard. 7: 367-370.<br />
Reinders, E. 1935. Fiber-tracheids, libriform wood fi- Saldanha da Gama, J. de. 1872. Configuraqao e Estudio<br />
bers and systematics in wood anatomy. Trop. Woods Botanico dos Vegetaes Seculares da Provincia do Rio<br />
44: 30-36.<br />
de Janeiro e de Outros Pontos do Brasil. Laemmert, Rio<br />
Richard, A. 1830. Memoire sur la famille des Rubiacees. de Janeiro.<br />
Imprimerie de J. Tatsu. [First copies apparently distrib- Sandwith, N. Y. 1949. Contributions to the flora of tropical<br />
uted in July 1829]. Paris. (Reimpr. Mem. Soc. Hist. Nat. America: XLIX. Notes on <strong>Rubiaceae</strong>. Kew Bull. 1949:<br />
Paris, ser. 3, 5: 81-224. 1834).<br />
253-264.<br />
Richard, L. C. M. 1803. Pinckneya. In: A. Michaux, Flora Schumann, K. 1888a. <strong>Rubiaceae</strong>, trib. VII-IX. In: C. F. P.<br />
boreali-americana, vol. 1: 103-105, t. 13. Fratres von Martius, A. G. Eichler & I. Urban (eds.), Flora<br />
Levrault, Paris.<br />
brasiliensis enumeratio plantarum in Brasilia hactenus<br />
Ridsdale, C. E. 1982. A revision of Badusa (<strong>Rubiaceae</strong>, detectarum... 6(6): 1-124. Fleischer, Leipzig.<br />
Condamineeae, Portlandiinae). Blumea 28: 145-150. . 1888b. Uber einige verkannte oder wening gekannte<br />
Rizzini, C. T. 1947. Chimarrhis. Novitates rubiacearum Geschlechter der Rubiaceen Sfidamerikas. Bot. Jahrb.<br />
compositarumque. Rev. Brasil. Biol. 7: 275-277.<br />
Syst. 10: 302-363.
208 Flora Neotropica<br />
. 1889. <strong>Rubiaceae</strong>, trib. X-XIX. In: C. F. P. von<br />
Martius, A. G. Eichler & I. Urban (eds.), Flora brasili-<br />
ensis enumeratio plantarum in Brasilia hactenus<br />
detectarum... 6(6): 125-466. Fleischer, Leipzig.<br />
. 1891. <strong>Rubiaceae</strong>. In: A. Engler & K. Prantl (eds.).<br />
Die Natirlichen Pflanzenfamilien 4(4): 1-156. Engel-<br />
mann, Leipzig.<br />
& K. Krause. 1908a Chimarrhis. In: K. Krause,<br />
<strong>Rubiaceae</strong> andinae. Engl. Bot. Jahrb. 40: 312-313.<br />
- & -- . 1908b. Acrobotrys. In: I. Urban, Plantae<br />
novae andinae imprimis Weberbauerianae. III. Bot.<br />
Jahrb. 40: 316-318.<br />
& - . 1908c. Dolichodelphys. In: E. Ule, Verh.<br />
Bot. Vereins. Prov. Brandenburg 50: 102-104.<br />
Simpson, D. R. 1976. Studies in neotropical <strong>Rubiaceae</strong>. I.<br />
<strong>Rustia</strong>. Phytologia 33: 4-8.<br />
Skvarla, J. J., P. H. Raven, W. F. Chissoe & M. Sharp.<br />
1978. An ultrastructural study of viscin threads in<br />
Onagraceae pollen. Pollen & Spores 20: 5-143.<br />
Soejarto, D. D., P. G. Delprete, J. C. Regalado & D. A.<br />
Madulid. 1996. The true provenance and identity of<br />
Badusa philippica (<strong>Rubiaceae</strong>).<br />
Solereder, H. 1890. <strong>Rustia</strong> pauciflora. Studien uber die<br />
Tribus der Gaertnereen Bentham et Hooker. Ber.<br />
Deutsch. Bot. Ges. 8: 99-100.<br />
. 1893. Ein Beitrag zur anatomischen Charakteristik<br />
und zur Systematik der Rubiaceen. Bull. Herb. Boissier<br />
1: 167-183.<br />
. 1908. Systematic anatomy of the dicotyledons.<br />
Clarendon Press, Oxford.<br />
Spjut, R. W. 1994. A systematic treatment of fruit types.<br />
Mem. New York Bot. Gard. 70: 1-182.<br />
Standley, P. C. 1916. <strong>Rustia</strong>ferruginea. Systematic inves-<br />
tigations of tropical American plants. Contr. U.S. Natl.<br />
Herb. 18: 142.<br />
- . 1918, 1921, 1934a, 1934b. <strong>Rubiaceae</strong>. N. Amer.<br />
Fl. 32: 1-300.<br />
. 1926. Chimarrhis microcarpa. In: N. L. Britton,<br />
West Indian plants. Bull. Torrey. Bot. Club 53: 471.<br />
. 1929. Condaminea breviflora. Studies on American<br />
plants-I. Publ. Field Columbian Mus., Bot. Ser. 4: 264.<br />
. 1930a. The <strong>Rubiaceae</strong> of Colombia. Publ. Field<br />
Columbian Mus., Bot. Ser. 7(1): 1-176.<br />
. 1930b. Studies on American plants-IV, Publ. Field<br />
Columbian Mus., Bot. Ser. 8: 131-236.<br />
. 1931a. The <strong>Rubiaceae</strong> of Ecuador. Publ. Field<br />
Columbian Mus., Bot. Ser. 7(2): 177-178.<br />
. 1931b. The <strong>Rubiaceae</strong> of Bolivia. Publ. Field<br />
Columbian Mus., Bot. Ser. 7(3): 179-340.<br />
. 1931c. The <strong>Rubiaceae</strong> of Venezuela. Publ. Field<br />
Columbian Mus., Bot. Ser. 7: 341-485.<br />
-- . 193 Id. Phitopis sterculioides. Studies on American<br />
plants-V. Publ. Field Columbian Mus., Bot. Ser. 8: 341.<br />
-- . 1932. Holtonia, a new genus of trees of the family<br />
<strong>Rubiaceae</strong>. Trop. Woods 30: 37-38.<br />
.1936. <strong>Rubiaceae</strong>. In: J. F. Macbride, Flora of Peru.<br />
Field Mus. Nat. Hist., Bot. Ser. 13: 1-263.<br />
- . 1940. <strong>Rubiaceae</strong>. Studies on American plants. Publ.<br />
Field Mus. Nat. Hist., Bot. Ser. 22: 48-58, 107-125,<br />
171-218, 278-294, 382-390.<br />
. 1947. Stomandra. Studies on Central American<br />
plants-VII. Publ. Field Mus. Nat. Hist., Bot. Ser. 23:<br />
247-248.<br />
& J. A. Steyermark. 1953. Chimarrhis<br />
venezuelensis. Botanical exploration in Venezuela-III.<br />
Fieldiana, Bot. 28: 571-573.<br />
& L. O. Williams. 1975. <strong>Rubiaceae</strong>. In: Flora of<br />
Guatemala. Fieldiana, Bot. 24: 1-274.<br />
Stellfeld, C. 1948. Os novos generos e as novas species de<br />
Freire AllemAo. Pongetti, Rio de Janeiro.<br />
Stearn, W. T. 1957. Achille Richard's "M6moire sur la<br />
famille des Rubiac6es." Taxon 6: 186-188.<br />
Steyermark, J. A. 1963. Dioicodendron, a new genus of<br />
<strong>Rubiaceae</strong>. Bol. Soc. Venez. Ci. Nat. 25: 23-28.<br />
. 1964a. A new <strong>Tresanthera</strong> from Venezuela. Bol.<br />
Soc. Venez. Ci. Nat. 25: 248-249.<br />
. 1964b. Cuatrecasasiodendron. Novedades en las<br />
Rubiaceas Colombianas de Cuatrecasas. Acta Biol.<br />
Venez. 4: 29-35.<br />
-. 1964c. <strong>Rubiaceae</strong>. In: B. Maguire, J. J. Wurdak &<br />
Collaborators, Botany of the Guayana Highlands, <strong>Part</strong><br />
V. Mem. New York Bot. Gard. 10: 186-278.<br />
.1965. <strong>Rubiaceae</strong>. In: B. Maguire, J. J. Wurdak &<br />
Collaborators, Botany of the Guayana Highlands, <strong>Part</strong><br />
VI. Mem. New York Bot. Gard. 12: 178-285.<br />
---. 1967. <strong>Rubiaceae</strong>. In: B. Maguire, J. J. Wurdak &<br />
Collaborators, Botany of the Guayana Highlands, <strong>Part</strong><br />
VII. Mem. New York Bot. Gard. 17: 230-436.<br />
. 1971. Pogonopus speciosus. Rubiaceas nuevas de<br />
los Andes y Cordilleras de la costa de Venezuela. Acta<br />
Bot. Venez. 6: 108-110.<br />
. 1972. <strong>Rubiaceae</strong>. In: B. Maguire, J. J. Wurdak &<br />
Collaborators, Botany of the Guayana Highlands, <strong>Part</strong><br />
IX. Mem. New York Bot. Gard. 23: 227-832.<br />
-- . 1974. <strong>Rubiaceae</strong>. In: T. Lasser & J. A. Steyermark,<br />
Flora de Venezuela 9: 1-2070. Instituto BotAnico, Caracas.<br />
--- . 1975. Revision of the genus Joosia (<strong>Rubiaceae</strong>).<br />
Brittonia 27: 251-262.<br />
---- . 1986. Holstianthus, a new genus of <strong>Rubiaceae</strong> from<br />
the Guyana Highland. Ann. Misssouri Bot. Gard. 73: 495.<br />
Swofford, D. L. 1993. PAUP: Phylogenetic analysis using<br />
parsimony. Version 3.1.1. Computer program distributed<br />
by the Illinois Natural History Survey, Champaign, Illinois.<br />
Tange, C. 1995. The identity of Siderobombyx and a new<br />
species ofXanthophytum (<strong>Rubiaceae</strong>). Nordic J. Bot. 15:<br />
575-581.<br />
Taylor, C. M. 1995. First International Conference on the<br />
<strong>Rubiaceae</strong>: Introduction. Ann. Missouri Bot. Gard. 82:<br />
355-356.<br />
& B. E. Hammel. 1993. Notes on Elaeagia<br />
myriantha, Comb. Nov. (<strong>Rubiaceae</strong>). Novon 3:490-491.<br />
Troll, W. 1950. Uber den Infloreszenzbegriff und seine<br />
Anwendung aufdie krautige Region bliihender Pflanzen.<br />
Abh. Akad. Wiss. Lit. Mainz, Mat.-Nat. KI. Nr. 13.<br />
Tryon, A. F. 1963. Hermann Karsten, his collections and<br />
the Flora columbiae. Taxon 12: 103-105.<br />
Turczaninow, N. S. 1848. Molopanthera. Decades Quarta<br />
et Quinta, generum adhuc non descriptorum. Bull. Soc.<br />
imp. Natural. Moscou 21: 580-581.<br />
Urban, I. 1899, 1903,1912,1913,1923. In: I. Urban(ed.),<br />
Symbolae antillanae seu fundamenta florae Indiae<br />
occidentalis. 9 Volumes, 1898-1928. Berlin, Leipzig,<br />
Paris, London.<br />
. 1922. Ariadne. Sertum antillanum XIV. Feddes<br />
Repert. Spec. Nov. Regni Veg. 18: 25-26.<br />
- . 1931. <strong>Rustia</strong> haitiensis. Plantae haitienses novae<br />
vel rariores a cl. Ekman 1924-1930 lectae. Stokholm.<br />
Almqvist & Wiksells Botryckeri. [reimpr. Ark. Bot. 24A<br />
(4): 36].<br />
Verdcourt, B. 1958. Remarks on the classification of the<br />
<strong>Rubiaceae</strong>. Bull. Jard. Bot. Etat. 28: 209-281.
Numerical List of Taxa209<br />
Waha, M. 1984. Zur Ultrastruktur un Funktion pollenverbindender<br />
Faden bei Ericaceae und anderen Angiospermenfamilien.<br />
PI. Syst. Evol. 147: 189-203.<br />
Walpers, W. G. 1852. Henlea. Annales botanices<br />
systematicae 2: 788. F. Hofmeister, Leipzig.<br />
Weberling, F. 1977. Beitrige zur Morphologie der Rubiaceen-<br />
Infloreszenzen. Ber. Deutsche Bot. Ges. 90: 191-209.<br />
. 1992. Morphology of flowers and inflorescences.<br />
Cambridge Univ. Press.<br />
Weddell, H. A. 1849. Histoire Naturelle des Quinquinas,<br />
ou monographie du genre Cinchona suivie d'une description<br />
du genre Cascarilla et de quelques autres plantes<br />
de la meme tribu. Victor Masson, Paris.<br />
- . 1854. Howardia. Notice sur quelques Rubiacees<br />
de l'Amerique tropicale. Ann. Sci. Nat. 4: 66-71, pl. 10.<br />
Wernham, H. F. 1911. Pteridocalyx. A new genus of <strong>Rubiaceae</strong>.<br />
J. Bot. 49: 317-318.<br />
NUMERICAL LIST OF TAXA<br />
. 1912. Carmenocania. New <strong>Rubiaceae</strong> from tropical<br />
America-I. J. Bot. 50: 241-242. Tab. 520.<br />
- . 1913. Pteridocalyx minor. New <strong>Rubiaceae</strong> from<br />
tropical America-II. J. Bot. 51: 218.<br />
. 1916. Pseudomussaenda, a new genus of <strong>Rubiaceae</strong>.<br />
J. Bot. 54: 297-301.<br />
- . 1917. Blandibractea, Neobertiera. Tropical American<br />
<strong>Rubiaceae</strong>-VIII. J. Bot. 55: 170-171.<br />
Willaman, J. J. & B. G. Schubert. 1961. Alkaloid-bearing<br />
plants and their contained alkaloids. Techn. Bull.<br />
U.S.D.A. 1234: 1-287.<br />
Willdenow, C. L. von. 1801. Sickingia. Zhen Neue<br />
Gattungen von Gewichsen. Ges. Naturf. Freunde Berlin<br />
Neue Schriften 3: 445-448.<br />
Wodehouse, R. P. 1935. Pollen grains. McGraw-Hill, New<br />
York.<br />
1-1. <strong>Rustia</strong> costaricensis (Standley) Lorence<br />
5-3. P. tubulosus (A. Richard in A. P. de Candolle) K.<br />
1-2. R. occidentalis (Bentham) Hemsley<br />
Schumann<br />
1-3. R. dressleri Delprete<br />
1-4. R. alba Delprete<br />
6-1. Chimarrhis parviflora Standley<br />
1-5. R. bilsana Delprete<br />
6-2. C. latifolia Standley<br />
1-6. R. rubra Standley ex D. Simpson<br />
6-3. C. cubensis Steyermark<br />
1-7. R. viridiflora Delprete<br />
6-4. C. jamaicensis (Urban) Steyermark<br />
1-8. R. schunkeana Delprete<br />
6-5. C. cymosa Jacquin<br />
1-9. R. thibaudioides (Karsten) Delprete<br />
6-6. C. glabriflora Ducke.<br />
1-10. R. venezuelensis Standley & Steyermark<br />
6-7. C. hookeri K. Schumann<br />
1-11. R. formosa Klotzsch<br />
6-8. C. microcarpa Standley<br />
1-12. R. 6-9. C.<br />
simpsonii Delprete<br />
speciosa (Steyermark) Delprete<br />
1-13. R. angustifolia K. Schumann<br />
6-10. C. turbinata A. P. de Candolle<br />
1-14. R. gracilis K. Schumann<br />
6-11. C. barbata (Ducke) Bremekamp<br />
6-12. C. duckeana Delprete<br />
2-1. <strong>Tresanthera</strong> condamineoides Karsten<br />
6-13. C. brevipes Steyermark<br />
2-la. T. condamineoides var. condamineoides<br />
6-14. C. gentryana Delprete<br />
2-lb. T. condamineoides var. thyrsiflora (Steyermark) Delprete 7-1. Dioicodendron dioicum (K. Schumann & K. Krause)<br />
3-1. Condaminea corymbosa (Ruiz & Pav6n) A. P. de Candolle<br />
Steyermark<br />
3-2. C. elegans Delprete<br />
8-1. Molopanthera paniculata Turczaninow<br />
8-la. M. paniculata var. paniculata<br />
4-1. Picardaea cubensis (Grisebach) Britton ex Urban 8-lb. M. paniculata var. burchellii (J. D. Hooker) K.<br />
Schumann<br />
5-1. Pogonopus exsertus (Orsted) Orsted<br />
5-2. P. speciosus (Jacquin) K. Schumann<br />
9-1. Dolichodelphys chlorocrater K. Schumann & K. Krause<br />
5-2a. P. speciosus var. speciosus<br />
5-2b. P. speciosus var. sandwithianus (Steyermark) Steyermark 10-1. Parachimarrhis breviloba Ducke
210 Flora Neotropica<br />
Abbiatti, D. & L. Claps, 288 (5-3).<br />
Acevedo-Rodriguez, P. et al., 6774, 6900 (1-2).<br />
Acosta-Solis, M., 12635 (3-1).<br />
Acufia, J., 8917, s.n. (6-3).<br />
Adair, s.n. (6-12).<br />
Adams, C. D., 9289 (6-4).<br />
Aguilera, V., 17 (5-1).<br />
Alain, Bro. & J. Acufia, 7673 (4-1).<br />
Albert de Escobar, L., 1373 (3-1).<br />
Alexander, R. C., s.n. (6-4).<br />
Allard, H. A., 22129 (3-1).<br />
Allart, A., 236 (5-2a).<br />
Allen, C., 622 (5-2b).<br />
Allen, P. H., 860 (1-2); 2017 (5-1); 3217 (1-9); 5613 (6-2);<br />
5621 (5-1); 5762 (6-2); 5910 (3-1); 6302 (6-2).<br />
Almeda, F. et al., 4264 (3-1).<br />
Alvarado, A., 233 (1-6); 292 (6-7).<br />
Alverson, W. S. et al., 28 (10-1); 214 (5-1).<br />
Ancuash, E., 504 (6-7); 604 (3-1); 1256, 1372 (6-6); 1509<br />
(3-1).<br />
Anderson, s.n. (6-5).<br />
Anderson, W. R., 9927 (1-11).<br />
Andersson, L. et al., 2108 (7-1).<br />
Andre, E., 1955 (3-1).<br />
Antonio, T. M., 1935 (1-2).<br />
Antonio, T. M. & W. J. Hahn, 4240 (1-2).<br />
Apparicio, P. A., 919, 59161 (1-11).<br />
Argeli, C. & C. A. de Oliveira, 691 (1-11).<br />
Aristeguieta, L., 5414, 6635 (5-2a).<br />
Asplund, E., 7844, 8571, 12428, 18347 (3-1); 18686 (1-6).<br />
Assuncao, 369 (6-11).<br />
Aymard, G. et al., 5233 (7-1).<br />
Badcock, W. J., 107, 739 (5-3).<br />
Badillo, V. M., 335 (5-2a).<br />
BAFOG Service Forestiere, 166M, 1187, 1262, 7608 (6-10).<br />
Bailey, L. H. & E. Z. Bailey, 272 (6-5); 865 (5-2a).<br />
Baker, M. A. & D. Neil, 5556 (3-1).<br />
Baker, M. A. et al., 6000 (6-7).<br />
Balee, W. L. & B. G. Ribeiro, 431, 2826 (6-10).<br />
Bang, M., 610 (3-1); 1344(5-3); 1429(3-1); 1716, s.n. (5-3).<br />
Barclay, A. S. et al., 3424 (7-1); 3463 (3-1).<br />
Barfod, A., 41389 (3-1).<br />
Barreto, M., 1789 (JBBH16686), 8305 (JBBH26371) (8-1b).<br />
Barrett, G. E., s.n. (5-2a).<br />
Barrier, S., 4993, 5060 (6-10).<br />
Barroso, G. et al., 66 (RB 179015) (5-3).<br />
Bartlett, H. H., 20361, 20389 (5-3).<br />
Bautista, H. P. & R. P. Orlandi, 985 (8-la).<br />
Bayley, E., s.n. (5-2a).<br />
Beard, P., 1134 (6-5).<br />
Beck, H. T. et al., 1055 (6-14).<br />
Belanger, 195, 196 (6-5).<br />
Belem, P., 3677 (8-la).<br />
Belshaw, C. M., 3422 (3-1); 3514 (1-8); 3593 (9-1).<br />
Bena, P., 1066 (6-5); 1187, 1262 (6-10).<br />
Benavides, O., 5651 (3-1).<br />
Benitez de R., C. E., 1083 (5-2a); 1513 (3-1).<br />
Bennett, B. C. & P. G6mez Andrade, 3638 (6-6).<br />
Bennett, B. C. et al., 4028 (6-6).<br />
Benoist, R., 1190 (6-10).<br />
Bensman, R., 331 (6-6).<br />
Berg, 1227 (1-7).<br />
LIST OF EXSICCATAE<br />
Berlin, B., 1812(3-1).<br />
Berardi, A. L., 563, 2209 (3-1); 5829 (2-lb); 5943 (6-9).<br />
Berry, P., 1847 (5-2a).<br />
Besse, L. et al., 602 (5-3).<br />
Billiet, F. & B. Jadin, 5010 (6-5).<br />
Bisby, F. et al., P18095 (6-10).<br />
Black, G. A., F8-47-955 (6-10).<br />
Black, G. A. et al. F8-50-9831 (6-10).<br />
Blanchet, J., 2557, 3282, s.n. (8-la).<br />
Blanco, C., 822 (6-8).<br />
Blum, K. E., 1260 (5-1).<br />
Boeke, J. D., 1501 (3-1).<br />
Boeke, J. D. & S. Utzschneider, 2925 (5-1).<br />
Bonpland, M. A., 622 (3-1).<br />
Boom, B., 10698 (6-8).<br />
Boom, B. & S. Mori, 2037, 2041, 2043, 2050, 2067, 2069,<br />
2078, 2080, 2180, 2213, 2217, 2226, 2230, 2263, 2445<br />
(6-8).<br />
Boom, B. & D. Beardsley, 8443 (3-1).<br />
Boom, B. et al., 8756 (6-10).<br />
Borsini, O., s.n. (5-3).<br />
Brade, A. C., 10639 (1-11).<br />
Brandbyge, J. & E. Asanza C., 30431 (3-1); 31615 (6-7).<br />
Bravo & G6mez, 306 (6-7).<br />
Bredemeyer, F., s.n. (5-2a).<br />
Breedlove, D. E., 22535, 28572 (5-1).<br />
Breedlove, D. E. & R. F. Thore, 30715 (5-1).<br />
Breteler, F. J., 3612 (3-1).<br />
Bridges, s.n. (5-3).<br />
Bristan, N., 1330, 1346, 1425, 1519 (5-1).<br />
Britton, N. L., 610 (6-4).<br />
Broadway, W. E., 3076 (2-la); 6464, 10011 (6-8); s.n. (2-la).<br />
Bruijn, J. de, 1646, 1647, 1651 (6-8).<br />
Buchtien, 0., 761 (5-3); 1393 (3-1); 1397 (5-3); 8186, s.n.<br />
(3-1).<br />
Burchell, W. J., 3181 (1-11).<br />
Burger, W. & G. Matta, 4593 (3-1).<br />
Burger, W. et al., 12314 (1-1).<br />
Burkart, A., 17038 (5-2a).<br />
Busey, P., 464 (1-1).<br />
Cabrera, A. L., 4159, 14523 (5-3).<br />
Cabrera, A. L. et al., 22348, 30389 (5-3).<br />
Cabrera R., I., 3264 (1-2); 3701 (1-9).<br />
Calder6n, S., 1225 (5-1).<br />
Callejas, R. & O. Escobar, 3234 (3-1).<br />
Callejas, R. et al., 2512 (3-1); 4160(10-1); 5038 (1-2); 5824<br />
(3-1).<br />
Camp, W. H., A-C25, E-25, E-1016, E-1540, E-2384 (3-1).<br />
Campos, M. D. et al., 48 (1-11).<br />
Campos, M. T. V. do A. et al., 468 (INPA 185019) (6-11).<br />
Cardenas, D., 2593, 4261, 5895 (5-3).<br />
Cavalcante, P. & M. Silva, 1598 (6-10).<br />
Cazalet, P. C. D. & T. D. Pennington, 7789 (6-6).<br />
Castillo, A., 1966 (5-2a).<br />
Ceballos, A. et al., 36 (5-3).<br />
Cer6n, C., 1600 (3-1).<br />
Cer6n, C. E., 3661 (6-7).<br />
Cer6n, C. E. & J. Ayala, 9656 (6-7).<br />
Cer6n, C. E. et al., 4353 (3-1); 8458, 8462 (6-7).<br />
Chagas, A. S. L. et al. 2217 (6-10).<br />
Chagas, J., 486 (MG25131) (6-12).<br />
Chavez, F., 265 (6-7).
List of Exsiccatae211<br />
Chavez-Alfaro, R., 359 (5-3).<br />
Churchill, H. W. & G. de Nevers, 4446 (5-1).<br />
Cid, C. A. et al., 4896, 4939, 5022 (5-3).<br />
Claes, F., s.n. (1-9).<br />
Clark, J. L., 2979 (1-5).<br />
Clark, J. L. et al., 139 (1-5).<br />
Claussen, P., 669, s.n. (1-11).<br />
Clement, Bro. & P. Roca, 9941 (6-3).<br />
Constance, L. & O. Tovar, 2272 (3-1).<br />
Cook, O. F. & G. B. Gilbert, 913 (3-1); 1389, 1564 (5-3).<br />
Cooper, G. P., 38 (6-5).<br />
Cooper, G. P. & G. M. Slater, 3, 120 (6-1); 260 (6-2); 401<br />
(6-1).<br />
Cordeiro, 676 (7-13).<br />
Core, E. L. 566, 1307 (3-1); 1388 (7-1); 1416 (3-1).<br />
Cornejo, X., 582 (6-6).<br />
Correa, M. N. et al., 4577 (5-3).<br />
Cosson, E., 18 (6-5).<br />
Cowan, R. S., 1439 (2-la).<br />
Croat, T. B., 8959 (5-1); 9828 (1-2); 11244 (6-1); 12400,<br />
12465, 12511, 12652, 12895 (5-1); 21099 (3-1); 22191<br />
(1-1); 26547 (3-1); 27508, 27551, 27709 (1-1); 27755,<br />
36431 (6-1); 57632, 69930 (3-1).<br />
Cruz, R., WB-00482 (5-1).<br />
Cuadros V., H., 950 (1-2); 1243 (3-1); 1677 (5-2b).<br />
Cuadros V. J. H. et al., 3884 (5-1).<br />
Cuatrecasas, J., 2325 (3-1); 7370, 7819 (1-9); 8665 (7-1);<br />
8926 (9-1); 8947 (6-6); 11059 (3-1); 11076 (1-9); 13074<br />
(3-1); 14317 (1-2); 15192 (7-1); 15905, 17580, 19736<br />
(1-2); 22458, 22516 (7-1).<br />
Cuatrecasas, J. et al., 28672 (7-1).<br />
Cuezzo, A. R., 10.262 (5-3).<br />
Daly, D. C. et al., 4312 (6-11); 6496 (3-1).<br />
Daniel, Bro., 314 (3-1); 2787 (5-1); 5533 (1-9).<br />
Dannouse, I., 6946 (6-8).<br />
D'Arcy, W. & K. Sytsma, 14523 (5-1); 14584 (3-1).<br />
Davidse, G., 4174 (5-2a).<br />
Davidse, G. & G. Herrera C., 26263 (3-1).<br />
Davidson, C., 4805 (3-1).<br />
Davidson, M. E., 792 (3-1).<br />
Davis, E. W., 184 (1-9).<br />
Dayton, W. A. & W. R. Barbour, 3007 (6-1).<br />
Duarte, A. P. et al., 60854 (1-11).<br />
Delprete, P., 6196 (6-6); 6319,6329,6330 (1-2), 6526 (6-7),<br />
6527, 6528 (6-6).<br />
Delprete, P. & P. Apreza, 6359 (5-2b).<br />
Delprete, P. & G. Fagua, 6378, 6381, 6382 (1-9).<br />
Delprete, P. & L. Gilbert, 5267 (1-2); 5271 (6-2).<br />
Delprete, P. & J. Luteyn, 6205, 6207 (3-1).<br />
Delprete, P. & A. Verduga, 6412 (3-1); 6414, 6419, 6420<br />
(1-4); 6421, 6422 (6-7); 6423, 6424 (6-6); 6436 (3-1).<br />
Delprete, P. & G. L. Webster, 6104 (3-1).<br />
Delprete, P. et al., 6049, 6218 (3-1).<br />
Del Risco, E., 27343, 27395, 27458 (6-3).<br />
De Marco, N. E. et al., 10262 (5-3).<br />
Denslow, J., 2680 (9-1).<br />
De Paula, 3233 (1-11).<br />
Devia, W., 316, 399, 558, 559, 559A (3-1).<br />
Dezzeo, N., 73 (6-13).<br />
Diaz, V., 32 (5-1).<br />
Diaz Pedrahita, S., 3446, 3817 (7-1).<br />
Dick, 290 (6-7).<br />
D'Orbigny, A., 620 (5-3).<br />
Dodge, C. W. et al., 16977 (5-1).<br />
Dodson, C. H. & P. M. Dodson, 12001 (3-1).<br />
Dodson, C. H. & A. Gentry, 13726 (3-1).<br />
Dombey, J., 555 (3-1).<br />
Dorr, L. J. et al., 5719 (3-1).<br />
Dressier, R. L., 4207 (1-2); 4703, 4749 (1-3); 4912 (3-1).<br />
Drew, W. B., 546, E-607 (3-1).<br />
Dryander, E., 1976 (3-1); 2583 (1-2).<br />
Duarte, A. P., 989 (RB61684), 3037 (RB71834) (1-11); 4842<br />
(RB 110068) (8-lb).<br />
Duarte, A. P. et al., s.n. (RB60854) (1-11).<br />
Ducke, A., 38 (RB20719) (6-11); 860 (6-11); 1235 (MG19119)<br />
(6-10); 1618 (6-6); s.n. (MG15480), s.n. (MG15757) (6-10);<br />
s.n. (RB15400) (10-1); s.n (RB15685) (6-10); s.n.<br />
(RB15687) (10-1); s.n. (RB17355), s.n. (RB17356) (6-10);<br />
s.n. (RB17358) (6-11); s.n. (RB21691), s.n. (RB21692) (6-<br />
10); s.n. (RB22821) (1-11); s.n. (RB22845) (6-11, 6-12<br />
[mixed collection]); s.n. (RB22846) (6-12); s.n. (RB22847)<br />
(6-11); s.n. (RB35055) (6-11).<br />
Ducke, A. & J. C. Kuhlmann, s.n. (RB4620), s.n. (RB4688)<br />
(1-1 1).<br />
Dudley, T. R., 11488 (3-1).<br />
Duke, J. A., 4465, 5361, 6127 (5-1); 9653, 10567, 10605<br />
(1-2); 10621 (3-1); 14597 (5-1).<br />
Duke, J. A. & J. Idrobo, 11580 (1-2).<br />
Dunlap, V. P., 33 (6-1).<br />
Duss, R. P., 345, 1452, 1880, 1883, 2548, 3424 (6-5).<br />
Dwyer, J., 2231, 2290 (1-2); 3095 (5-1); 6178 (3-1); 8372,<br />
15265 (5-1).<br />
Dwyer, J. & B. MacBryde, 9626 (3-1).<br />
Dwyer, J. et al., 523 (3-1).<br />
E. L. C., s.n. (5-1).<br />
Edwall, G., 23938 (1-11).<br />
Eggers, F., 5812 (2-la); 13392 (5-2a).<br />
Elias et al., 583 (5-1).<br />
Elias, Bro., 450 (5-2a).<br />
Ekman, E. L., 3642, 4641, 5772, 7859, 7919, 9384 (4-1);<br />
9417, 9692 (6-3); 10408 (1-15); 10389 (6-4).<br />
Espina, J. et al., 2503 (1-2).<br />
Espinal S., T., 623, 633 (5-1).<br />
Eyerdam, W. J., 180 (4-1).<br />
Eyerdam, W. J. & A. A. Beetle, 22876 (5-3).<br />
Falcao, M. & D. Coelho, 209 (6-12).<br />
Fanshawe, D. B., F1249 (3985) (6-8).<br />
Famey, C. et al., 1237 (1-11).<br />
Fassett, N. C., 25427 (1-9); 25756 (3-1).<br />
Fendler, A., 289 (1-2); 584 (5-2a); 2366 (2-la).<br />
Fenmndez, A., 256 (1-2).<br />
Ferreira, V. F. et al., 285 (1-14).<br />
Ferreyra, R., 1112, 3612, 4208, 4542, 6823 (3-1).<br />
Fiebrig, K., 2705 (5-3).<br />
Field, A., 424 (6-9).<br />
Fishlock, W. C., 10, H338-16 (6-5).<br />
Flores-Paitan, S., 3B, 61 (6-7).<br />
Folsom, J. P., 6241 (5-1).<br />
Folsom, J. P. & R. Lantz, 1931 (5-1).<br />
Folsom, J. P. & R. Robinson, 2365 (6-1).<br />
Folsom, J. P. et al., 6581 (5-1).<br />
Fonnegra, R. & F. J. Roldan, 2278 (3-1).<br />
Fonnegra, R. et al., 2514 (3-1).<br />
Forero, E., 7689 (1-11).<br />
Forero, E. et al., 4418 (1-2); 6093 (3-1).<br />
Forest Department of British Guiana, 3985 (F1249), 5885<br />
(WB479), 6991 (F3427), 7892 (RB68) (6-8).<br />
Forestry Bureau Suriname, 1151, 1680, 2040, 2415, 2464,<br />
2585, 2907, 4952 (6-10).
212 Flora Neotropica<br />
Fors, A., 142 (6-3).<br />
Fosberg, F. R., 19440 (1-9); 19527 (7-1); 19880,20388 (3-1);<br />
21857 (7-1); 27750 (3-1); 28653 (5-3); 28672 (3-1).<br />
Fosberg, F. R. & L. R. Holdridge, 19409 (3-1).<br />
Foster, R. B., 1411 (1-2); 2877 (6-1); 4133 (1-2).<br />
Foster, R. B. & E. Vivar, 13333 (3-2).<br />
Frazod, A., s.n. (RB2316) (1-14).<br />
Freire, E. & J. Cerda, 461 (1-6).<br />
Freire-Fierro, A. et al., 2333 (3-1).<br />
Freire-Fierro, A. & J. Luteyn, 2159 (3-1).<br />
Freire-Fierro, A. & P. Yanez, 2697 (1-6).<br />
Freitas, C. A. A. et al., 83 (5-3).<br />
French Guiana Forest Service, 7608 (6-10).<br />
Friedrichshal, E. R. von, s.n. (1-2).<br />
Froes de Lemos, R., 20777 (6-10); 31049, 32097 (6-10);<br />
s.n. (RB22596) (6-12).<br />
Fuchs, H. P. & L. Zanella, 21703 (1-2).<br />
Fuentes, A. F., 471 (5-3).<br />
Fuertes, P., 649 (4-1).<br />
Funck, N., 372, 463 (5-2a).<br />
Funck, N. & L. J. Schlim, 1637 (3-1).<br />
Fundaci6n Biol6gica Puerto Rastrepo, 5052 (10-1).<br />
Garcia, 147 (6-9).<br />
Garcia-Barriga, H., 8197 (3-1); 18763 (7-1).<br />
Garcia-Barriga, H. & G. Stout, 18907 (3-1).<br />
Garwood, N. C., 719, 802, 1015, 1102, 1129 (3-1); 2531<br />
(6-1).<br />
Garwood, N. C. et al., 10 (1-2); 70 (3-1).<br />
Gaudichaud, C., 1830 (1-14).<br />
Gay, M. C., 919 (3-1).<br />
Gehrt, A., s.n. (1-11).<br />
Gentry, A., 1785, 5609, 6720 (5-1); 70741 (6-7).<br />
Gentry, A. & H. Cuadros, 55525 (5-2b).<br />
Gentry, A. & J. Aronson, 25307 (6-14).<br />
Gentry, A. & C. Diaz, 61118 (3-1); 58655 (5-3).<br />
Gentry, A. & S. Estensoro, 70562 (3-1); 70733 (6-7).<br />
Gentry, A. & M. Fallen, 17201 (3-1).<br />
Gentry, A. & E. Forero, 7195 (3-1); 7377 (1-2).<br />
Gentry, A. & N. Jaramillo, 28093 (6-6); 41321 (3-1).<br />
Gentry, A. & A. Juncosa, 40718 (3-1).<br />
Gentry, A. & W. Morawetz, 63183 (6-10).<br />
Gentry, A. & S. Mori, 13874 (5-1).<br />
Gentry, A. & G. Morillo, 10339 (5-2a).<br />
Gentry, A. & OTS class, 48572 (6-2).<br />
Gentry, A. & J. Revilla, 16645 (10-1).<br />
Gentry, A. & G. Shupp, 26475 (3-1).<br />
Gentry, A. et al., 27780, 31219,31384, 36636 (6-7); 37904<br />
(3-1); 40418A (1-2); 40125 (5-3); 40160 (3-1); 42942<br />
(6-6); 43018, 43027, 43135, 45661,46255 (6-7); 53411<br />
(1-2); 55625, 60913, 65564, 65621, 72152 (6-7).<br />
Gillis, W. T., 7535, 11168 (5-1).<br />
Giordano, L. C. et al., 1005 (1-11).<br />
Glaziou, A., Ibis, 148 (1-11); 1098 (1-12); 2607 (1-13);<br />
6564 (1-12); 12036 (1-14); 12038 (1-13); 13974, 13975,<br />
14892 (8-1); 14893 (1-14); 14937 (1-13); 18305 (5-2a);<br />
18306(1-13); 18309, 18905 (5-2a); 18906(1-13); 19437<br />
(1-14); 19438 (8-1); 21458a, s.n. (1-11).<br />
Gomes, M., 323 (1-13).<br />
G6mez-Laurito, J., 10694 (3-1); 11698 (1-1).<br />
G6mez P., L. D., 3307 (1-2).<br />
G6mez P., L. D. et al., 22770, 22944 (6-1), 23844 (3-1);<br />
24504 (5-1).<br />
Gosling, C., s.n. (5-3).<br />
Gottsberger, G. K., G213-13888 (3-1).<br />
Goudot, J., s.n. (1-9, 3-1).<br />
Goulding, M., 1048 (10-1).<br />
Grandez, G. et al., 1768 (6-13).<br />
Grant, M. L., 10620 (3-1); 10891 (5-2b).<br />
Grant, V., 630 (5-1).<br />
Granville, J. J. de, B-2671, B-2799, B-5167 (6-8); B-5377<br />
(6-10).<br />
Granville, J. J. de et al., 9041 (6-8).<br />
Grayum, M. et al., 4689 (3-1).<br />
Grenand, 1238, 1500(6-10).<br />
Grubb, P. J. et al., 1485 (1-8).<br />
Gudiflo, E., 1707 (6-6).<br />
Gudifio, E. & N. Andi, 2220 (6-7).<br />
Gudifio, E. & G. Grefa, 2336 (6-7).<br />
Gudifto, E. & C. Gualinga, 1640 (6-7).<br />
Guti6rrez, A., 32 (6-7).<br />
Haber, W. & E. Bello, 1735, 1736, 6837, 7152 (6-1).<br />
Hahn, M., 116 (5-1); 429 (6-5).<br />
Halton, J. & L. Besse, 101 (3-1).<br />
Hammel, B., 9753 (6-1); 12446 (6-1).<br />
Hammel, B. et al., 16858 (6-1); 17540 (3-1); 17726 (5-1);<br />
17719 (6-2).<br />
Harling, G., 3763 (1-6); 6138, 11016 (3-1); 11076 (1-6).<br />
Harling, G. & L. Andersson, 11860, 13755, 14065 (3-1);<br />
16423 (1-6); 16979 (3-1); 17003 (1-6); 23151,23944 (3-1).<br />
Harling, G. et al., 9916, 9769, 10079 (3-1).<br />
Harmon, E. W. & J. A. Fuentes, 4965 (5-1).<br />
Harris, W., 5810, 5828, 8783, 9449 (6-4).<br />
Harris, W. & N. L. Britton, 10576 (6-4).<br />
Hart, J., 109 (1-2).<br />
Hartshorn, G. S., 1211, 1500 (6-1); 1687 (6-7).<br />
Hartshorn, G. S. & J. Quijano, 2925 (6-6).<br />
Hartshorn, G. S. et al., 2906,2925,2932 (6-6); 62451 (5-3).<br />
Hatschbach, J. H., 23784 (5-3); 38796,42945,43851 (1-11).<br />
Haught, 0., 2236, 3705 (5-2b); 5427 (1-2).<br />
Hauman II, 5520 (5-3).<br />
Haymard, G. et al., 5233 (7-1).<br />
Hayes, S., 164 (5-1).<br />
H6guert, J. S. d', 642 (5-2a).<br />
Herbario Florestal Personnel, s.n. (8-lb).<br />
Heringer, E. P., 7065, 7681, 9156, 10468 (1-11).<br />
Heringer, E. P. et al., 1439, 1784, 3157, 3188, 3232, 4448,<br />
5082,6157(1-11).<br />
Hernmndez, J. J. & S. E. Hoyos, 182 (9-1).<br />
Herrera, G., 1976 (6-1); 3519 (3-1).<br />
Herrera, H., 1663 (1-3).<br />
Hill, S. R., 24770 (6-5).<br />
Hinds, 354, s.n. (1-2, 3-1).<br />
Hioram, Bro. & C. Maurel, 6066 (6-3).<br />
Hodge, W. H., 9 (6-7); 695 (6-5); 6048 (3-1).<br />
Hoene, F. C., s.n. (1-11).<br />
Hoffman, B. et al., 1532 (6-8).<br />
Holdridge, L. R., 6723 (6-1).<br />
Holm-Nielsen, L., 16311 (3-1).<br />
Holm-Nielsen, L. & J. Jaramillo, 28968 (3-1).<br />
Holm-Nielsen, L. et al., 3776, 20567, 20613 (3-1); 25235,<br />
26039, 26057 (1-2).<br />
Holton, I. F., 410 (7-1); 440 (3-1).<br />
Hoogte, L. v. d. & C. Roersch, 2791 (5-3).<br />
Hoover, W. S. & S. Wormsley, 1420 (3-1).<br />
Hoover, W. S. et al., 3047 (3-1).<br />
Horowitz, S., 663 (5-3).<br />
Howard, R. A., 11143 (6-5); 16044 (5-1).<br />
Howard, R. A. & L. I. Nevling, 16910 (5-1).<br />
Howard, R. A. & E. S. Howard, 18846, 19487 (6-5).<br />
Howard, R. A. et al., 19930 (6-5).
List of Exsiccatae213<br />
Howell, J. T., 10184A (1-2).<br />
Huashikat, V., 2136 (6-14).<br />
Huertas, G. & L. A. Camargo, 6282 (7-1).<br />
Humbert, H., 26973 (3-1).<br />
Humboldt, A., 567 (3-1).<br />
Hunziker, A. T., 2037 (5-3).<br />
Hurtado, F., 2365 (6-7).<br />
Hurtado, F. & A. Alvarado, 2240, 2261 (6-7).<br />
Hurtado L., R., 5, 111 (3-1).<br />
Hutchison, P. C., 1153 (3-1).<br />
Hutter, C., 1215 (7-1).<br />
Idrobo, J. M., 287 (3-1); 4874 (9-1).<br />
Idrobo, J. M. & R. E. Schultes, 570 (1-9); 1011 (9-1).<br />
Ijjasz-Madriz, E., 422 (3-1).<br />
INPA/WWF 1109.25 (6-10).<br />
Irvine, D., 634 (6-6).<br />
Irwin, H. S. & T. R. Soderstrom, 5369 (1-11).<br />
Irwin, H. S. et al., 8881, 14160, 25206(1-1 1); 48440, 50615,<br />
55443 (6-10).<br />
Isem, J., 2382 (3-1).<br />
Jack, J. G., 8173, 8430 (5-1).<br />
Jacobs, B., 2315 (6-1).<br />
Jahn, A., 1339 (6-9).<br />
Jangoux, J. & R. P. Bahia, 86 (6-10).<br />
Jansen-Jacobs, M. J. et al., 1899 (6-10).<br />
Jaramillo, J., 14167, 14290 (1-8); 14990 (1-2).<br />
Jaramillo, J. & F. Coello, 4322 (1-8).<br />
Jaramillo, J. et al., 13199 (3-1); 31361 (6-6).<br />
Jardim, A. & A. Cadden, 2153 (5-3).<br />
Jardim, A. & Mamami, 1423 (5-3).<br />
Jardim, J. G. et al., 711 (8-la).<br />
Jativa, C. & C. Epling, 771 (1-2); 870 (3-1); 2001,2201 (1-2).<br />
Jelski, de, 376 (7-1).<br />
Jimenez, Q. & R. Zuniga, 760 (6-2).<br />
Jim6nez, Q. et al., 751 (5-1).<br />
Jim6nez L., 0., 541 (5-1).<br />
Jim6nez M., A., 2451 (3-1); 2898, 3173 (1-2); 3418 (6-1);<br />
3728 (3-1).<br />
Johnson, H., 57, 1022 (5-1).<br />
Johnston, I. M., 1745 (1-2).<br />
Jorgensen, P. M. & S. Laegaard, 56452 (3-1).<br />
Jorgensen-Hansen, P., 35009 (5-3).<br />
Juncosa, A., 1589 (1-2); 1660, 2435 (3-1); 2543 (1-2).<br />
Juzepczuk, S., 6869 (3-1).<br />
Kalbreyer, W., 927 (5-2a); 1246 (1-2).<br />
Kanehira, R., 107 (3-1).<br />
Karsten, H., s.n. (1-9); s.n. (2-la).<br />
Kayap, R., 929, 2012 (3-1).<br />
Keman, H. S., 19, 89 (3-1); 167 (5-2b).<br />
Kiesling, R., 7109 (5-3).<br />
Killip, E. P., 3284 (5-1); 9735 (3-1); 11707 (1-2); 11199<br />
(3-1); 34986 (1-2).<br />
Killip, E. P. & J. Cuatrecasas, 38680, 38970, 39150 (1-2).<br />
Killip, E. P. & H. Garcia-Barriga, 32209, 33128, 33209,<br />
33338 (1-2); 33635 (3-1).<br />
Killip, E. P. & A. C. Smith, 22449, 23506 (3-1); 23811 (5-3);<br />
24130 (3-1).<br />
King, S. R. & M. Ramirez-Rengifo, 372 (3-1).<br />
Kirizawa, M. & S. A. C. Chiea, 2350 (1-1 1).<br />
Kirkbride, J. H., 2637, 2667 (5-2b); 5431 (1-11).<br />
Klawe, W. L., 1492 (1-2).<br />
Klug, G., 1804 (9-1); 3672 (3-1).<br />
Knapp, S., 1335 (1-2); 2713 (5-1).<br />
Knapp, S. & J. Mallet, 3035 (3-1); 3070 (1-2); 6764 (2-la).<br />
Knapp, S. et al., 4586 (6-1).<br />
Krapovickas, A. & A. Schinini, 31024,31193,31475,39017<br />
(5-3).<br />
Krapovickas, A. et al., 19143 (5-3).<br />
Krukoff, B. A., 1018 (6-10); 8782 (6-14); 10032 (3-1);<br />
10054, 10167 (5-3).<br />
Kuhlmann, M., 3769 (1-11).<br />
Kuhlmann, M. & S. Jimbo, 310, 329 (6-10).<br />
Kuhlmann, J. G., 82 (RB14083) (8-lb); 503 (RB139367)<br />
(1-11); 1221 (8-lb); 2353 (RB15366) (5-3); s.n.<br />
(RB16318)(1-11); s.n. (RB17357)(6-12); s.n. (RB17384)<br />
(6-11); s.n. (RB70222) (1-1 1).<br />
Kuntze, O., s.n. (5-3).<br />
Lamarck, W., s.n. (6-5).<br />
Lanjouw, J. & J. C. Lindeman, 2810 (6-10).<br />
Lansberge, J. G. van, 214 (5-2a).<br />
Lao Magin, R., 44, 51, 5107 (6-7).<br />
Lasser, T., 2264 (6-9).<br />
Lee, K., 49 (6-5).<br />
Legname, P. R. & A. R. Cuezzo, 5967, 6129, 7964, 9047<br />
(5-3).<br />
Lehmann, F. C., 473 (6-4); 5651 (3-1); 7459 (5-1); 7718<br />
(7-1); K-132 (1-2); s.n. (1-9).<br />
Leite, I., 2 (6-12).<br />
Lemos, C., 28678 (1-11).<br />
Lent, R. W., 359 (1-2).<br />
Le6n, Bro.,455 (5-1); 11011, 12813 (6-3).<br />
Le6n, Bro. & Matos, 19644 (4-1).<br />
Le6n, Bro. & M. Victorin, 17235 (4-1).<br />
Le6n, Bro. et al., 9941 (6-3).<br />
Le6n, J., 1484 (3-1); 2143 (6-1).<br />
Lescure, 2060 (6-6).<br />
Lewis, M., 88148, 88825, 88914 (3-1).<br />
Lewis, W. H. et al., 1 (5-1); 692 (3-1); 3392 (5-1); 13543<br />
(6-14).<br />
Liesner, R. & A. Gonzales, 10940 (1-9).<br />
Lillo, M., 10785, 22336 (5-3).<br />
Lima, J. & B. Zimmerman, 519 (6-10).<br />
Linden, J., 874 (3-1); 1604 (9-1); s.n. (5-2b).<br />
Lino, A. M., 75 (8-la).<br />
Liogier, A., 14353 (4-1).<br />
Liogier, A. & J. Acunia, 7673 (4-1).<br />
Liogier, A. & P. Liogier, 26174 (4-1).<br />
Little, E. L. Jr., 7186 (1-9); 8205 (7-1); 8490, 8757, 15872<br />
(3-1).<br />
Little, E. L. Jr. et al., 298 (3-1), 763 (6-6).<br />
L0jtnant, B. & U. Molau, 13513 (6-7).<br />
L6pez-Palacios, S., 1939, 1940 (5-2a).<br />
Lorentz & Hieronymus, 384 (5-3).<br />
Lourenco, s.n. (RB1042) (1-11).<br />
Lourteig, A., 714 (5-3).<br />
Lozano C., G. & J. Diaz, 3734 (7-1).<br />
Lozano C., G. et al., 2832 (5-2b); 4395, 4885 (7-1).<br />
Lugo, M., 111(3-1); 115(1-6); 141 (3-1).<br />
Lugo S., H., 1871 (3-1); 3674, 3946 (1-7); 4427,6034 (3-1).<br />
Luschnath, B., s.n. (1-11).<br />
M. A. V., 42 (6-7).<br />
Maas, P. J. M. & L. Y. T. Westra, 3912 (6-8).<br />
MacBride, J. F., 3662 (3-1); 5343 (5-3).<br />
MacBride, J. F. & J. Dwyer, 1408 (6-6).<br />
Macedo, A., 2402 (3-1).<br />
Machado, A. A., 11-1984 (5-2a).<br />
Macias & Burbano, 7788 (6-7).
214 Flora Neotropica<br />
Maciel, U. N. et al., 776 (6-10).<br />
Madison, M. T. et al., 4562 (3-1); 4808 (1-4); 5051 (3-1).<br />
Madsen, J., 85961 (7-1).<br />
MagalhAes, M., s.n. (8-lb).<br />
Maguire, B. & L. Politi, 27971 (6-13).<br />
Mamani, F., 436 (5-3).<br />
Manara, B., s.n. (5-2a).<br />
Marcano Berti, L., 241, 1426 (6-8).<br />
Marin, 2109 (3-1).<br />
Marin, E., 740 (6-13).<br />
Maroto, R. & E. Salsa, 123 (5-2a).<br />
Marques, M. C. et al., 285 (1-11).<br />
Marquete, R. et al., 832, 1084, 1371 (1-11).<br />
Martinelli, A. et al., 4133 (1-11).<br />
Martinelli, G., 6835 (6-10).<br />
Martinelli, G. et al., 13320 (RB28042) (1-11).<br />
Martinet, M., 1412, s.n. (3-1).<br />
Martius, K. F., 57 [in. obs.] (1-13); 111 [in. obs.] (1-11);<br />
2607, 2608 (1-9); 2964, 2965, 2966, 2967 (1-13); 3139,<br />
3201 (1-9).<br />
Mathias, M. E. & D. Taylor, 5932 (3-1).<br />
Mathews, A., 3030 (3-1).<br />
Mattheus, 1954 (3-1).<br />
Mattos, A. et al., 322 (1-11).<br />
Mattos Silva, L. A. et al., 2847 (8-la).<br />
Matuda, E., 273, 560, 16811, 18748 (5-1).<br />
McDaniel, S. & M. Rimachi Y., 20503 (10-1).<br />
McDowell, T. et al., 4411, 4472 (6-8).<br />
McPherson, G., 9146 (6-1).<br />
Mendonca, R.C. et al., 1292 (1-11).<br />
Meneces, E. & W. Terceros, 9 (6-6).<br />
Metcalf, R. D., 30676 (3-1).<br />
Mexia, Y., 6351 (3-1); 8035 (5-3).<br />
Meyer, T., 903, 4810, 4811, 8477 (5-3).<br />
Meyer, W. & S. Llamosa, 3627 (2-la).<br />
Meyer, W. et al., 2039 (2-la).<br />
Michel, R. de, 41 (5-3).<br />
Molina, F., s.n. (3-1).<br />
Molina R., A. et al., 1868, 2085 (1-2); 18005 (3-1).<br />
Montalvo, E. A., 3849 (5-1).<br />
Monteiro, M. T., 23498 (8-la).<br />
Moore, A., 10648 (6-4).<br />
Morales, J. F. et al. 1124 (6-2).<br />
Morawetz, W. & B. Walln6fer, 11-161085 (3-1).<br />
Moreno, L. M., 193 (5-2b).<br />
Moreno, P. P., 23868, 26317 (6-1).<br />
Mori, S., 421 (5-1).<br />
Mori, S. & J. Cardoso, 17128 (6-10).<br />
Mori, S. & C. Gracie, 18943 (6-8).<br />
Mori, S. & J. Kallunki, 3086 (1-1).<br />
Mori, S. & J. Pipoly, 15562 (6-8).<br />
Mori, S. et al., 10328 (8-la); 15016, 15109 (6-8); 15786,<br />
16248, 18691 (6-10); 20910 (6-8); 21567, 23098, 23645<br />
(6-10).<br />
Morillo, G. et al., 3298 (2-la).<br />
Moritz, K., 840 (5-2a).<br />
Mos6n, H., 3408 (1-11).<br />
Mulford Biological Exploration, 1016 (5-2a).<br />
Museu Goeldi Personnel, 9425, 9713 (6-10).<br />
Mutis, J. C., 2 (3-1); 3 (1-9); 1004 (3-1); 2194 (7-1); 4651<br />
(3-1).<br />
Nascimento, J. R. M. et al., s.n. (INPA/WWF 1109.302)<br />
(6-12).<br />
Nee, M., 8846, 9125 (5-1); 41570 (5-3).<br />
Nee, M. & G. Coimbra S., 35199 (5-3).<br />
Nee, M. & M. Saldias P., 36320 (5-3).<br />
Neill, D., 9817, 9840, 9848 (6-7).<br />
Neill, D. & A. Alvarado, 9040 (6-7).<br />
Neill, D. & W. Palacios, 7104 (6-7); 9718 (1-8).<br />
Neill, D. & J. Zaruma, 365, 7021 (6-7).<br />
Neill, D. et al., 6105 (3-1); 6204 (6-7); 6220 (6-6); 6515 (6-6);<br />
7394, 9145 (3-1).<br />
Nufiez, P., 6104 (3-2).<br />
Nufiez, P. & C. Mufioz, 5122 (3-1).<br />
Nufiez, P. et al., 10098 (5-3).<br />
Occhioni, P., 6655 (1-14); s.n. (RB15455) (5-1).<br />
Oldeman, R. A. A., B-674 (6-10).<br />
Oldeman, R. A. A. & Arevalo, 16 (6-7).<br />
Oliveira, A. de, s.n. (INPA 60564) (6-12).<br />
Oliveira, E. de, 3836, 3854, 4036, 4798 (6-10).<br />
Oliveira, R. P., 5 (3-1).<br />
011gaard, B. et al., 34627, 34667, 35520, 90375 (3-1).<br />
Orlandi, R., 223 (8-la).<br />
Orsted, A. S., 11536, 11537, s.n. (5-1).<br />
Otto, E., 901 (5-1).<br />
Ownbey, M., 2682 (1-6); 2686 (3-1); 2735 (1-8).<br />
Pab6n E., M., 960 (1-9).<br />
Pabst, G., 4667 (1-11).<br />
Palacios, W., 1994 (6-6); 2828 (6-7); 6616 (1-8).<br />
Palacios, W. & J. Clark, 12633 (7-1).<br />
Palacios, W. & D. Rubio, 7290 (7-1).<br />
Palacios, W. et al., 240, 365 (6-7); 3380 (6-6); 9663 (1-4).<br />
Parodi, L. R., 9220 (5-3).<br />
Patris, s.n. (6-10).<br />
Paula, J. E. de, 3233 (1- 1).<br />
Paulino Filho, H. F. & M. G. Silva, 93-96 (5-3).<br />
Pearce, R. W., s.n. (5-3).<br />
Pennell, F. W., 3503 (3-1).<br />
Peredo, Y., 525 (5-3).<br />
Pereira, B. A. S., 270 (1-11).<br />
Pereira, B. A. S. & D. Alvarenga, 2503 (1-11).<br />
Pereira, B. A. S. & V. V. Mecenas, 2442 (1-11).<br />
Pereira, E., 7363 (1-11).<br />
Pereira, E. & A. P. Duarte, 4484 (1-11).<br />
Pereira, E. et al., 3762 (RB266353) (1-11).<br />
Perez, P., s.n. (3-1).<br />
Perez-Arbelaez, E., 697 (3-1).<br />
Perez-Arbelaez, E. & J. Cuatrecasas, 5733 (3-1).<br />
Perrottet, 283 (6-5).<br />
Persson, C. et al., 43 (1-7).<br />
Pflanz, K., 937, 2053 (5-3).<br />
Philipson, W. R. & J. M. Idrobo, 1921 (1-9); 1955 (9-1).<br />
Philipson, W. R. et al., 1576 (9-1).<br />
Picarda, L., 1129(4-1).<br />
Pierotti, S. A., 76 (5-3).<br />
Pinto et al., 370 (1-9).<br />
Pipoly, J. J. et al., 12525 (6-7).<br />
Pirani, J. R. & R. Mello Silva, SPF 47733 (8-lb).<br />
Pires, J. M., 6826, 6922, 6969, 6978, 7107, 48902 (6-10).<br />
Pires, J. M. & R. P. Belem, 13006 (6-10).<br />
Pires, J. M. & N. T. Silva, 1742, 10879, 10880, 11895,<br />
11897, 11898 (6-10).<br />
Pires, J. M. et. al., 4997 (6-10); 9270, 9539 (1-11); 50615,<br />
50853, 51224, 51420 (6-10).<br />
Pittier, H., 58,236 (5-2a); 2396,2476 (5-1); 4166 (1-2); 4817<br />
(5-1); 5251 (3-1); 5507 (5-1); 7468 (5-2a); 9799, 10299,<br />
11672 (5-2a); 13854,14138,15275 (6-9); 15582 (5-2a).<br />
Pl1e, A., s.n. (6-5).<br />
Plowman, T. & W. Davis, 4762, 5164 (5-3).
List of Exsiccatae 215<br />
Plowman, T. & H. Kennedy, 5649 (5-3).<br />
Pohl, J. E., 809, 809d, 810, 2681, 3442, 3444, s.n. (1-11).<br />
Ponthieu, de, s.n. (6-5).<br />
Popenoe, W., 802 (5-1).<br />
Prance, G. T. et al., 4452 (6-8); 12462 (6-6).<br />
Prieto, F., ChuP-20 (1-7).<br />
Proctor, G. R., 19742, 19818, 21467, 22591 (6-4).<br />
Proctor Cooper III, G., 38, 38A (6-5).<br />
Pulle, A., 379 (6-8).<br />
Purdie, W., s.n. (1-9).<br />
Quevedo, R., 316 (5-3).<br />
Rabelo, B. V. & J. Cardoso, 2887 (6-10).<br />
Ragonesi & Coras, 37794, 37799 (5-3).<br />
Ramage, G. A., s.n. (6-5).<br />
Ramirez, J. G. & D. Cardenas Lopez, 179, 1017, 1501, 1620,<br />
1879 (9-1).<br />
Ramos, J. & G. Mota, 368 (5-3).<br />
Ramos, L. S. & C. Rosario S., 15 (6-10).<br />
Renteria, E. et al., 658 (3-1).<br />
Revilla, J. & C. H. Froehner, 2449 (6-6).<br />
Reynel, C. & J. Le6n, 4044 (3-1).<br />
Ribamar, J. & J. Ramos, 164 (6-10).<br />
Ribeiro, J. E. L. S. & P. A. C. L. Assuncao, 1776 (INPA<br />
185097) (6-12).<br />
Richard, L. C. M., s.n. (6-5).<br />
Riedel, L., 66,68 (1-13); 187 (1-11); 367 (1-14); 369 (5-3);<br />
424 (1-14); 480 (1-11); 1236 (1-11); 1229 (5-3); 1462<br />
(1-11); s.n. (1-11).<br />
Riedel, L. & B. Luschnath, 1062 (1-13).<br />
Riera, B. & D. Sabatier, 1844 (6-10).<br />
Rimachi Y., M., 2973 (6-6); 10178 (3-1); 10501 (10-1).<br />
Rizzini, C. T. & A. Mattos, 1126 (8-la).<br />
Robinson, W. & M. W. Lyon, s.n. (5-2a).<br />
Rodrigues, R. R. & N. Figueredo, 15745 (1-11).<br />
Rodrigues, W., 8573, 9954, 9976 (6-12).<br />
Rodrigues, W. & D. Coelho, 7600 (INPA 16903) (6-12).<br />
Rodriguez, H., 266 (5-2a).<br />
Rodriguez, W., 8098, 8099, 8692 (6-10); 9954 (6-12).<br />
Roig, F. A., 6419 (6-3).<br />
Rojas, T., 1145, 1546 (5-3).<br />
Romero-Castafieda, R., 132, 609 (5-2b); 2309 (3-1); 11198<br />
(5-2b); s.n. (1-2).<br />
Rosa, N.A., 1096 (6-10).<br />
Rosa, P., 145 (RB131708) (1-11).<br />
Rosario S., C. et al., 852, 858 (6-11); 1092 (6-10).<br />
Rose, J. N. & G. Rose, 21691 (5-2a); 23426 (3-1); s.n. (5-2a).<br />
Rudas, A. et al., 3016 (6-6).<br />
Ruiz L., H. & J. J. A. Pav6n, s.n (3-1).<br />
Ruiz Teran, L. E., 488 (6-9); 1204 (3-1).<br />
Ruiz Teran, L. E. & M. L6pez Figueras, 658 (3-1).<br />
Rusby, H. H., 106, 711, 1898 (3-1); 2090 (5-3).<br />
Rusby, H. H. & R. W. Squires, 126 (6-8).<br />
Sabatier, D., 606 (6-10).<br />
Sabatier, D. & M. F. Prevost, 2116, 3356, 3731 (6-10).<br />
Saer, d'H., J., 642 (5-2a).<br />
Sagastegui A., A., 5872 (3-1).<br />
Saint George Expedition, 538, 624 (1-2).<br />
Saint Hilaire, A., 348 (1-11).<br />
Saint John, H., 20813, 20871 (3-1).<br />
Salim, J. Filho, 148, 159 (8-1a).<br />
Sampaio, A., 612 (1-11).<br />
Sanchez Vega, I. & M. Dillon 9006, 9052 (3-1).<br />
Sandemer, 6055 (3-1).<br />
Sandino, J. C., 3294, 3355, 3412, 4842 (6-1).<br />
Sandwith, N. Y., 1757 (2-1a).<br />
Sanoja, R. E., s.n. (3-1).<br />
Santillan, P., 90436 (5-3).<br />
Santos, T. S., 920 (8-la).<br />
Saundeman, C., 6055 (3-1).<br />
Schlim, L., 755 (5-2a); s.n. (5-2b).<br />
Schnee, L., 1174 (5-2a).<br />
Schreiter, 220, 3536, 11168 (5-3).<br />
Schultes, R. E., 24356 (1-9).<br />
Schultes, R. E. & I. Cabrera, 18929 (6-6).<br />
Schultz, A. G., 978 (5-3).<br />
Schulz, J. P., 7289, 8581 (6-10).<br />
Schunke, J. M., 331 (6-6).<br />
Schunke Vigo, J., 1527 (6-6); 2072 (3-1); 2901 (6-6); 3205,<br />
3952 (1-8); 4364,4815 (3-1); 4877,5683 (1-8); 5807,5845<br />
(3-1); 6162 (1-6); 6214 (5-3); 6696 (1-8); 7267,7659,9494,<br />
10079(3-1); 10270(1-8); 10362,10512(3-1); 10653(6-6).<br />
Schwacke, C. A. W., 1441 (5-1).<br />
Scolnick, R., 1678 (5-3).<br />
Scolnick, R. & R. Luti, 497 (3-1).<br />
Seemann, B., 478 (5-1); 1593, s.n. (3-1), s.n. (5-1).<br />
Servicio Forestal do Brasil, Personnel, 1043 (1-11).<br />
Shafer, J. A., 4406 (6-3); 7850 (4-1).<br />
Shattuck, O. E., 688 (5-1).<br />
Shepherd, J. D., 459 (9-1).<br />
Sieber, F. W., s.n. (6-5).<br />
Silva, A. S. L. da, et al., 1553, 2217 (6-10).<br />
Silva, M., 2794, 5073 (6-10).<br />
Silva, M. F., 576 (6-10).<br />
Silva, M. G., 5517 (6-10); 6125 (5-3).<br />
Silva, M. G. & C. Rosario S., 2420, 2475, 3732 (6-10).<br />
Silva, N. T., 51, 628, 905, 999, 1246, 2561, 2568, 3421,<br />
3924 (6-10).<br />
Silverstone-Sopkin, P. et al., 3001 (7-1).<br />
Skutch, A. F., 3950 (5-1); 3981 (3-I); 4777 (1-1).<br />
Slane, V., 140 (6-5).<br />
Sleumer, H., s.n. (5-3).<br />
Smith, A., 1890 (6-1).<br />
Smith, A. C., 3433 (6-8).<br />
Smith, C. E. et al., 3282 (5-1).<br />
Smith, D. N., 1311 (6-6); 1415 (5-3); 2075 (1-6); 4019 (6-6).<br />
Smith, D. N. et al., 1415 (5-3); 1423 (3-1); 5605 (5-3); 6325<br />
(3-1).<br />
Smith, E. E., 396 (6-3).<br />
Smith, H. H., 100 (5-2b).<br />
Smith, H. H. & G. W. Smith, 889, 1245 (6-5).<br />
Smith, L. B. & E. L. McWilliams, 15364 (1-11).<br />
Smith, R. F., VI-280 (5-2a).<br />
Sobel, G. L. & J. Strudwick, 2430 (3-1).<br />
Sodiro, P. L., s.n. (1-6).<br />
Soejarto, D. D., 1086 (7-1); 3443 (5-1).<br />
Soejarto, D. D. et al., 1287 (9-1).<br />
Solanilla, O., 16 (5-1).<br />
Solomon, J. C., 3097 (5-3); 3153 (3-1); 7321 (5-3); 7322<br />
(3-1); 7359, 17641 (5-3); 18445 (3-1).<br />
Solomon, J. C. & J. Kuijt, 11585 (3-1).<br />
Sothers, C. et. al., 568 (INPA185102), 625 (INPA185105),<br />
750 (INPA185110) (6-11).<br />
Soukup, J., 597, 2281 (3-1).<br />
Spada, J., 31, 289, 293 (8-lb).<br />
Sparre, B., 14380 (3-1); 18300, 18340 (1-2).<br />
Spegazzini, C., 13814 (5-3).<br />
Spruce, R., 4579 (3-1); 4930 (6-7).<br />
Stahel, G., 261, 261a (6-10).<br />
Standley, P. C., 21672, 29972, 62055, 79549 (5-1).
216 Flora Neotropica<br />
Standley, P. C. & E. Padilla V., 2885 (5-1).<br />
StAhl, B. et al., 1822, 1866 (1-7); 2800 (1-6).<br />
Steere, W. C., 8016 (3-1); 8021 (1-6); 8060, 8337 (3-1);<br />
8339 (1-6).<br />
Steere, W. C. & W. H. Camp, 8228 (1-6); 8255 (3-1).<br />
Stehle, H., 987, 2298, 3015, 3652, 5068, 6626, 7178 (6-5).<br />
Stein, B. A. & D. D'Alessandro, 2776 (3-1).<br />
Stein, B. A. & C. Todzia, 2142, 2358 (3-1); 2372 (5-3).<br />
Steinbach, J., 9292 (3-1).<br />
Stem, W. L., & D. Wasshausen, 2582 (6-5).<br />
Stem, W. L., et al., 724 (5-1).<br />
Stevens, F. L., 662 (5-1).<br />
Stevens, W. D., 24597 (6-1).<br />
Stevens, W. D. et al., 24859 (6-1).<br />
Stewart, A., 315 (1-2).<br />
Steyermark, J. A., 54450 (7-1); 54732, 54865 (3-1); 54921<br />
(5-2a); 55220, 55223, 55408 (7-1); 56208, 56306 (3-1);<br />
56333 (1-10); 56659 (3-1); 57662 (5-2a); 61136, 62185<br />
(6-8); 65741 (5-2a); 87104 (6-9); 89514 (6-8); 89885,<br />
89961 (6-9); 91807 (5-2a); 100495 (9-1).<br />
Steyermark, J. A. & P. Berry, 112141 (5-2a).<br />
Steyermark, J. A. & C. B. Carias, 105353 (5-2a).<br />
Steyermark, J. A. & G. Davidse, 116278, 116885 (6-9).<br />
Steyermark, J. A. & V. C. Espinoza, 112706,108790 (5-2a).<br />
Steyermark, J. A. & M. Rabe, 96163 (2-la).<br />
Steyermark, J. A. & C. Steyermark, 95211 (6-9).<br />
Steyermark, J. A. et al., 111625 (6-8); 114008 (5-2a);<br />
121534 (2-la).<br />
Stork, H. E. et al., 10494 (5-3).<br />
Strier, K. B., 1007 (8- b)<br />
Sucre, D., 88 (5-3).<br />
Sucre, D. & E. P. Heringer, 624 (1-11).<br />
Svenson, H. K., 331 (1-2).<br />
Sytsma, K. & W. G. D'Arcy, 3391 (5-1); 3480 (3-1).<br />
Tafalla, J. J. & J. A. Manzanilla, s.n. (3-1).<br />
Tate, H. H., 488 (3-1); 543 (5-3).<br />
Tello, 87, 129,313 (6-6); 489, 862, 1284 (6-7); 1295, 1434,<br />
1995 (6-6).<br />
Tessmann, G., 4512 (3-1); 4668 (6-7).<br />
Thiel, J., 878 (6-10).<br />
Thomas, J. et al., 1437 (1-9).<br />
Thompson, S. A. & J. E. Rawlins, 816 (3-1).<br />
Thompson, S. A. et al., 9859 (4-1).<br />
Tonduz, A., 3947, 4671 (3-1).<br />
Toro, R. A., 421 (3-1); 540 (5-1).<br />
Tovar, O., 1421, 4629 (5-3).<br />
Triana, J., 1782 (3262.1) (3-1); 1784 (3262.3) (1-2); 1785<br />
(3262.4) (1-9); 1869 (7-1); s.n. (3262) (1-9).<br />
Trigoso, J. R., 1025 (3-1).<br />
Troll, C., 1514 (5-3); 1745 (3-1).<br />
Trujillo, B., 1413, 3416 (5-2a).<br />
Trujillo, B. & A. Fernmndez, 293 (5-2a).<br />
Tunqui, S., 205, 945 (6-7).<br />
Turpe, A. M. et al., 4690 (5-3).<br />
Tuxill, J. et al., 63 (6-5).<br />
Tyson, E. L., 2629 (5-1).<br />
Uferide, T., 13 (6-9).<br />
Ule, E., 6774 (9-1); 9848, 9848b (5-3).<br />
Uribe Uribe, L., 912, 1441, 1481 (5-1); 5468 (7-1); 6385 (3-1).<br />
Utley, J. & K. Utley, 5337 (6-1).<br />
Vaca, A. A., 18(5-3).<br />
Vaillant, P., 21 (8-la).<br />
Van der Werff, H. & E. Gudifio, 10874 (3-1); 11098 (7-1);<br />
11326 (9-1).<br />
Vargas C., C., 1719, s.n. (5-3).<br />
Vargas, H. & P. Grefa, 908 (1-6).<br />
Vargas, H. et al. 563 (6-7).<br />
Vargas V., V., 2575 (3-1).<br />
Vasquez, C. B., 14-CBV (6-6).<br />
Vasquez, R., 10631(6-6).<br />
Vasquez, R. & N. Arevalo, 8969 (6-14).<br />
Vasquez, R. & G. Criollo, 1855 (6-7).<br />
Vasquez, R. & N. Jaramillo, 2431 (6-6).<br />
Vasquez, R. et al., 5404 (6-13).<br />
Vauthier, 15, 205, s.n. (1-11).<br />
Ventenat, L., s.n. (1-11).<br />
Ventura, E. & E. L6pez, 424, 2856 (5-1).<br />
Venturi, S., 5207, 7648 (5-3).<br />
Vivas, J. M., 2 (5-2a).<br />
Vogel, S., 54 (1-9).<br />
Wagner, M., 3495, 3496 (5-1).<br />
Wagner, R. J., 711, 897 (5-1).<br />
Walker, F., 1277 (5-2b).<br />
Walln6fer, B., 26-19588 (3-1); 113-29788 (1-6).<br />
Warming, E., s.n. (8-1b).<br />
Warner, R. N., 316 (1-2).<br />
Warscewicz, J., 10, 11 (1-9).<br />
Weberbauer, A., 7940 (5-3).<br />
Webster, G. L., 13478 (6-5).<br />
Webster, G. L. & N. McCarten, 28816 (3-1).<br />
Webster, G. L. et al., 8341 (6-4); 9205, 9403 (6-5); 9805<br />
(2-la).<br />
Weddell, H. A., 2637, 2638 (1-11); 4190 (5-3); s.n. (5-3).<br />
Wedel, H. von, 2114 (1-2).<br />
Werling, L. & S. Leth-Nissen, 226 (3-1).<br />
West, J., 8261 (5-3).<br />
White, J. W. & R. H. Warner, 82 (1-2).<br />
Widgren, J., 491, 1036, s.n., s.n. (1-11).<br />
Wiggins, I. L., 10880 (8-1); 10947 (3-1).<br />
Wilbur, R. L., 37205 (6-1).<br />
Wilbur, R. L. et al., 8137 (6-5).<br />
Williams, LI., 369, 409 (6-7); 7794 (3-1); 10299, 10415,<br />
10646, 12408 (5-2a).<br />
Williams, L1. & A. H. G. Alston, 42 (5-2a).<br />
Williams, L. O., 5066 (8-Ib).<br />
Williams, L. O. & V. Assis, 6861 (8-la).<br />
Williams, L. O. et al., 28805 (3-1).<br />
Williams, R. O., 11881(6-8).<br />
Williams, R. S., 322 (3-1).<br />
Willink, 174(5-3).<br />
Wilson, F. G. & S. J. Browne, WB479 (5885) (6-8).<br />
Woods, C. W., 497 (5-2a).<br />
Woronow, G. J. N. & S. V. Jusepczuk, 6426 (9-1).<br />
Woytkowski, F., 6663 (3-1); 7380 (6-6).<br />
Wright, C., 1262, 1622 (6-3); 2661 (4-1).<br />
Wullschlagel, 1344bis (6-4).<br />
Wurdack, J. J., 1383 (7-1); 1831 (3-1); 2348 (9-1).<br />
Young, K., 942 (3-1).<br />
Young, K. & M. Eisenberg, 233 (5-3); 255, 342 (3-1).<br />
Young, K. & G. Sullivan, 610, 800, 864 (3-1).<br />
Young, K. et al., 493 (3-1).<br />
Yuncker, T. G., 4529 (5-1).<br />
Zabala, S., 165 (5-3).<br />
Zak, V., 3979 (6-7).<br />
Zak, V. & S. Espinoza, 5130 (6-6).<br />
Zak, V. & J. Jaramillo, 3577 (3-1).<br />
Zamora, N. et al. 1901 (6-2).<br />
Zanderij, I., 261 (6-10).
Nomenclatural List 217<br />
Zanoni, T. & J. Pimentel, 35991 (4-1).<br />
Zanoni, T. et al., 25634, 33125, 40662 (4-1).<br />
Zarucchi, J. L., 3265, 3338 (9-1).<br />
Zarucchi, J. L. et al., 5398 (3-1); 5465 (5-1); 6104 (3-1);<br />
7044 (5-1).<br />
NOMENCLATURAL LIST<br />
Zaruma, J., 802 (3-1).<br />
Zaruma, J. & A. Arguello, 451 (6-6).<br />
Zehntner, M., 38 (5-2a).<br />
Zuloaga, F. O. & N. B. Deginani, 331 (5-3).<br />
Zuluaga R., S., 1184 (5-1).<br />
Due to the complexity of the taxonomic problems and large number of transfers and rearrangements in the<br />
present treatment, a list of accepted taxa (boldface), orthographic variants, and synonyms, as recognized by<br />
the author, is given. Synonyms, orthographic variants, and unpublished names are followed by the corresponding<br />
accepted taxon. Authority abbreviations follow Brummitt & Powell, 1992.<br />
Allenathus Standl.<br />
A. erythrocarpus Standl.<br />
Acunaeanthus Borhidi, Jarai-Koml. & Moncada<br />
A. tinifolius (Griseb.) Borhidi<br />
Arachnotryx Planch. =Rondeletia<br />
Ariadne Standl. =Mazaea<br />
A. ekmanii Urb. =Mazaea shaferi<br />
A. shaferi (Standl.) Urb. =Mazaea shaferi<br />
Augusta Pohl (nomen cons.)<br />
Augusta subgen. Lindenia (Benth.) J.H.Kirkbr.<br />
Augusta subgen. Augusta (Pohl) J.H.Kirkbr.<br />
A. attenuata Pohl =Augusta longifolia var. parvifolia<br />
A. austro-caledonica (Brogn.) J. H. Kirkbr.<br />
A. glaucescens Pohl =Augusta longifolia var. longifolia<br />
A. lanceolata Pohl =Augusta longifolia var. longifolia<br />
A. longifolia (Spreng.) Rehder<br />
A. loagifolia (Spreng.) Rehder var. longifolia<br />
A. longifolia (Spreng.) Rehder var. parvifolia (Pohl) Delprete<br />
A. oblongifolia Pohl =Augusta longifolia var. longifolia<br />
A. parvifolia Pohl =Augusta longifolia var. parvifolia<br />
A. rivalis (Benth.) J.H.Kirkbr.<br />
A. vitiensis (Seem.) J.H.Kirkbr.<br />
Badusa A.Gray<br />
B. corymbifera (Forst.f.) A.Gray<br />
Bartramia W. Bartram =Pinckneya<br />
B. bracteata W.Bartram = Pinckneya bracteata<br />
(W.Bartram) Raf.<br />
Bartramia L. =Triumfetta Hedw. (Tiliaceae)<br />
Bartramia Hedw. (Bryophyta)<br />
Bartramia Salisb. =Penstemon Schmidel<br />
Bathysa Presl<br />
B. bathysoides (Steyerm.) Delprete<br />
B. bracteosa (Wedd.) Delprete<br />
B. difformis Benoist =Chimarrhis turbinata<br />
B. perijaensis (Steyerm.) Delprete<br />
B. pittieri (Standl.) Steyerm.<br />
Bignonia W. Bartram =Pinckneya<br />
Bignonia L. (Bignoniaceae)<br />
B. bracteata W.Bartram = Pinckneya bracteata<br />
(W.Bartram) Raf.<br />
Blandibractea Werham =Simira<br />
B. brasiliensis Wernham =Simira glaziovii<br />
Bonifacia Manso ex Steud. =Augusta<br />
B. riparia Manso ex Steud. =Augusta longifolia var. longifolia<br />
Buena muzonensis (Goudot) Wedd. =Ladenbergia<br />
muzonensis<br />
Calycophyllum DC.<br />
C. tubulosum Seeman =Pogonopus exsertus<br />
C. tubulosum (A.Rich. in DC.) DC. =Pogonopus tubulosus<br />
Capirona erythroxylon Standl. =Simira ecuatoriensis<br />
Carmenocania Wernham =Pogoaopus<br />
C. porphyrantha Wernham =Pogonopus speciosus var.<br />
speciosus<br />
Cascarilla muzonensis (Goudot) Wedd. =Ladenbergia<br />
muzonensis<br />
Chimarrhis Jacq.<br />
Chimarrhis Jacq. subgen. Chimarrhis<br />
Chimarrhis Jacq. subgen. Pseudochimarrhis (Ducke)<br />
Delprete<br />
C. barbata (Ducke) Bremek.<br />
C. bathysoides Steyerm. =Bathysa bathysoides<br />
C. brevipes Steyerm.<br />
C. clausicorollata J.H.Kirkbr. =Simira tinctoria<br />
C. cubensis Steyerm.<br />
C. cymosa Jacq.<br />
C. cymosa Jacq. subsp. genuina Urb. =Chimarrhis cubensis<br />
C. cymosa Jacq. subsp. jamaicensis Urb. =Chimarrhis<br />
jamaicensis<br />
C. cymosa Jacq. var. jamaicensis (Urb.) Standl. =Chimarrhis<br />
jamaicensis<br />
C. cymosa Jacq. subsp. microcarpa Urb. =Chimarrhis cubensis<br />
C. cymosa Jacq. var. microcarpa (Urb.) Standl. =Chimarrhis<br />
cubensis<br />
C. cyrrhosa Stend. (orth. var.) =C. cymosa<br />
C. decurrens Steyerm. =Allenanthus erythrocarpus<br />
C. dioica K.Schum. & K.Krause =Dioicodendron dioicum<br />
C. duckeana Delprete<br />
C. duckei Rizzini =Chimarrhis turbinata<br />
C. ferruginea (Standl.) Standl. =Bathysa sp.<br />
C. gentryana Delprete<br />
C. glabriflora Ducke<br />
C. goudotii Baill. =Simira goudotii<br />
C. hookeri K. Schum.<br />
C. jamaicensis (Urb.) Steyerm.<br />
C. latifolia Standl.<br />
C. longifolia (Willd.) Baill. =Simira longifolia<br />
C. longistipulata Bremek. =Chimarrhis microcarpa<br />
C. microcarpa Standl.<br />
C. microcarpa Standl. var. microcarpa =Chimarrhis<br />
microcarpa<br />
C. microcarpa Standl. var. speciosa Steyerm. =Chimarrhis<br />
speciosa<br />
C. paraensis Baill. =Simira paraensis
218 Flora Neotropica<br />
C. parviflora Standl.<br />
C. perijaensis Steyerm. =Bathysa perijaensis<br />
C. pisoniaeformis Baill. =Simira pisoniiformis<br />
C. pittieri Standl. =Bathysa pittieri<br />
C. speciosa (Steyerm.) Delprete<br />
C. turbinata DC.<br />
C. venezuelensis Standl. & Steyerm. =Dioicodendron dioicum<br />
C. williamsii Standl. =Chimarrhis hookeri<br />
Chrysoxylon Casaretto =Plathymenia (Mimosaceae)<br />
Chrysoxylon Wedd. =Pogonopus<br />
C. febrifugum Wedd. =Pogonopus tubulosus<br />
C. speciosum (Jacq.) Kuntze =Pogonopus speciosus var.<br />
speciosus<br />
C. tubulosum (A.Rich. in DC.) Kuntze =Pogonopus tubulosus<br />
Cinchona caroliniana Poir. =Pinckneya bracteata<br />
(W.Bartram) Raf.<br />
C. corymbifera Cav. =Badusa corymbifera<br />
C. muzonensis Goudot =Ladenbergia muzonensis<br />
C. philippica Cav. =Badusa corymbifera<br />
Coffea floribunda Miq., nom. superfl. =Molopanthera<br />
paniculata var. paniculata<br />
C. floribunda Martius = Ixora densiflora<br />
Condaminea DC.<br />
C. angustifolia Rusby =Condaminea corymbosa<br />
C. breviflora Standl. =Condaminea corymbosa<br />
C. corymbosa (R. & P.) DC.<br />
C. corymbosa (R. & P.) DC. var. pubescens Spruce ex<br />
K.Schum. =Condaminea corymbosa<br />
C. corymbosa (R. & P.) DC. var. subsessilis Regel, in herb.<br />
=Condaminea corymbosa<br />
C. elegans Delprete<br />
C. glabrata Bartl. ex DC. =Not Condaminea!<br />
C. macrophylla Poepp. & Endl. =Capirona decorticans<br />
C. microcarpa (R. & P.) DC. =Bathysa sp.<br />
C. petiolata Dwyer =Rondeletia panamensis<br />
C. tinctoria (Kunth in H.B.K.) DC. =Simira rubescens<br />
C. utilis Goudot =Elaeagia utilis<br />
C. venosa (R. & P.) DC. =Not Condaminea!<br />
Cuatrecasasiodendron Standl. & Steyerm.<br />
C. colombianum Standl. & Steyerm.<br />
C. spectabile Steyerm.<br />
Dioicodendron Steyerm.<br />
D. cuatrecasasii Steyerm. =Dioicodendron dioicum<br />
D. dioicum (K.Schum. & K.Krause) Steyerm.<br />
Dolichodelphys K.Schum. & K.Krause<br />
D. cholorocrater K.Schum. & K.Krause<br />
Elaeagia brasiliensis Standl. =Chimarrhis turbinata<br />
E. miryantha (Standl.) C.M.Taylor & Hammel<br />
Exandra Standl. =Simira Aubl.<br />
E. rhodoclada Standl. =Simira rhodoclada<br />
Exostema corymbifera (Forst.f.) Roem. & Schult. = Badusa<br />
corymbifera<br />
E. formosum Cham. & Schlecht. =<strong>Rustia</strong> formosa<br />
E. formosum var. laeve Cham. & Schlecht. =<strong>Rustia</strong> formosa<br />
E. formosum var. leprosum Cham. & Schlecht. =<strong>Rustia</strong><br />
formosa<br />
E. leonis Standl. =Acunaeanthus tinifolius<br />
E. (Pseudostemma) occidentale Benth. =<strong>Rustia</strong> occidentalis<br />
E. philippica (Cav.) Roem. & Schult. =Badusa corymbifera<br />
Flexanthera Rusby =Simira Aubl.<br />
F. fragrans Rusby =Simira fragrans<br />
F. subcordata Rusby =Simira cordifolia<br />
Forsteronia panniculata Casar. (nom. nud.) =Molopanthera<br />
paniculata var. paniculata<br />
Henlea Griseb. (Malpighiaceae)<br />
Henlea H.Karst. =<strong>Rustia</strong><br />
H. muzonensis (Goud.) Klotzsch & H.Karst. ex Walp.<br />
=Ladenbergia muzonensis<br />
H. rosea (Goudot) Klotzsch & H.Karst. ex Walp.<br />
=Ladenbergia muzonensis<br />
H. splendens H.Karst. =<strong>Rustia</strong> thibaudioides<br />
H. thibaudioides H.Karst. =<strong>Rustia</strong> thibaudioides<br />
Holtonia Standl. =Elaeagia<br />
H. myriantha (Standl.) Standl. =Elaeagia miryantha<br />
Howardia Wedd. =Pogonopus<br />
H. caracasensis Wedd. =Pogonopus speciosus var. speciosus<br />
H. febrifuga (Wedd.) Wedd. =Pogonopus tubulosus<br />
H. grandiflora Wedd. =Pogonopus speciosus var. speciosus<br />
H. richardii Wedd. =Pogonopus tubulosus<br />
Ixora L.<br />
I. densiflora (Martius) Mull. Arg.<br />
Javorkaea Borhidi & Jarai-Koml. =Rondeletia<br />
Kajewskiella Merrill & Perry<br />
K. polyantha M.Jansen<br />
K. trichantha Merrill & Perry<br />
Kerianthera J.H.Kirkbr.<br />
K. preclara J.H.Kirkbr.<br />
Ladenbergia muzonensis (Goudot) Standl.<br />
Lindenia Benth. =Augusta subgen. Lindenia<br />
L. austro-caledonica =Augusta austro-caledonica<br />
L. rivalis Benth. =Augusta rivalis<br />
L. vitiensis Seem. =Augusta vitiensis<br />
Macrocnemum P. Br.<br />
M. corymbosum R. & P. =Condaminea corymbosa<br />
M. cubense Griseb. =Picardaea cubensis<br />
M. exsertus 6rst. =Pogonopus exsertus<br />
M. glabratum Bartl. =Not Condaminea!<br />
M. longifolium A. Rich. =Chimarrhis cymosa<br />
M. microcarpon R. & P. =Bathysa sp.<br />
M. speciosus Jacq. =Pogonopus speciosus var. speciosus<br />
M. tinctorium Kunth in H.B.K. =Simira rubescens<br />
M. tubulosum A.Rich. in DC. =Pogonopus tubulosum<br />
M. venosum R. & P. =Not Condaminea'<br />
Mazaea Krug & Urb.<br />
M. phialanthoides (Griseb.) Krug & Urb.<br />
M. pungens (Urb.) Alain = Rondeletia sp.?<br />
M. shaferi (Standl.) Delprete<br />
M. tinifolia (Griseb.) Alain =Acunaeanthus tinifolius<br />
Molopanthera Turcz.<br />
M. burchellii J. D.Hook. =Molopanthera paniculata var.<br />
burchellii<br />
M. colombiana Standl., in herb. =Dioicodendron dioicum<br />
M. paniculata Turcz.<br />
M. paniculata Turcz. var. paniculata<br />
M. paniculata Turcz. var. burchellii (J. D.Hook.) K.Schum.<br />
M. panniculata K.Schum. (hort. var.) =Molopanthera<br />
paniculata<br />
Mussaenda bracteata (W.Bartram) Raf. =Pinckneya<br />
bracteata (W.Bartram) Raf.<br />
M. speciosa (Jacq.) Poir. =Pogonopus speciosus var. speciosus<br />
Neobertiera = Pteridocalyx?<br />
N. gracilis Wernham = Pteridocalyx gracilis (Wernham)<br />
Delprete, ined.(?)<br />
Neomazaea Krug & Urb. =Mazaea<br />
N. phialanthoides (Griseb.) Krug & Urb. =Mazaea<br />
phialanthoides
Nomenclatural List 219<br />
N. shaferi Standl. =Mazaea shaferi<br />
N. pungens Urb. =Rondeletia sp.?<br />
N. tinifolia (Griseb.) Krug & Urb. =Acunaeanthus tinifolius<br />
R. brandegeeana Lorence<br />
R. phialanthoides Griseb. =Mazaea phialanthoides<br />
R. tinifolia Griseb. =Acunaeanthus tinifolius<br />
<strong>Rustia</strong> Klotzsch<br />
Otocalyx Brandegee =Rondeletia<br />
0. chiapensis Brandegee =Rondeletia brandegeeana<br />
Parachimarrhis Ducke<br />
P. breviloba Ducke<br />
Picardaea Urb.<br />
P. cubensis (Griseb.) Britt. ex Urb.<br />
P. haitiensis Urb. =Picardaea cubensis<br />
Pinckneya Rich. in A.Michx.<br />
P. acroma Freire & Allem?o =Simira rubra<br />
P. bracteata (W.Bartram) Harper =Pinckneya bracteata<br />
(W.Bartram) Raf.<br />
P. bracteata (W.Bartram) Raf.<br />
P. erubescens Glaz. =Simira glaziovii<br />
P. ionantha Planch. & Lind. =Pogonopus speciosus var.<br />
sandwithianus<br />
P. officinalis Lavalle (nom. nud.) =Pinckneya bracteata<br />
(W.Bartram) Raf.<br />
P. pubens L.C.M. Rich. in A.Mich. =Pinckneya bracteata<br />
(W. Bartram) Raf.<br />
P. rubescens Allemao =Simira rubra<br />
R. alba Delprete<br />
R. angustifolia K.Schum.<br />
R. bilsana Delprete<br />
R. costaricensis (Standl.) Lorence<br />
R. dressleri Delprete<br />
R. ferruginea Standl. =Bathysa sp.<br />
R. formosa (Cham. & Schltdl. ex DC.) Klotzsch<br />
R. haitiensis Urb. = not <strong>Rustia</strong> (?)<br />
R. isernii Standl. =Alibertia isernii<br />
R. longifolia Standl. =Dolichodelphys chlorocrater<br />
R. mosenii K.Schum. ex Glaziou =<strong>Rustia</strong> formosa<br />
R. occidentalis (Benth.) Hemsl.<br />
R. panamensis Dwyer =<strong>Rustia</strong> costaricensis<br />
R. pauciflora K.Schum. ex Soler. =<strong>Tresanthera</strong> pauciflora<br />
R. pohliana Klotzsch =<strong>Rustia</strong> formosa<br />
R. rosea (Klotzsch & H.Karst. ex Walp.) K.Schum<br />
=Ladenbergia muzonensis<br />
R. rubra Standl. ex D. Simpson<br />
R. schunkeana Delprete<br />
R. secundiflora K.Schum. =<strong>Rustia</strong> thibaudioides<br />
R. sellowiana Klotzsch =<strong>Rustia</strong> formosa<br />
P. viridiflora Allemao & Saldanha =Simira viridiflora R. simpsonii Delprete<br />
Pinknea Persoon (orth. var.) =Pinckneya<br />
R. splendens (H.Karst.) Standl. =<strong>Rustia</strong> thibaudioides<br />
Pinknea pubescens Person (orth. var.) =Pinckneya R. thibaudioides (H.Karst.) Delprete<br />
bracteata (W.Bartram) Raf.<br />
R. venezuelensis Standl. & Steyerm.<br />
Pinkneya Raf. (orth. var.) =Pinckneya<br />
R. viridiflora Delprete<br />
Pogonopus A.Rich. in DC.<br />
R. warscewicziana Klotzsch, in herb. = <strong>Rustia</strong> thibaudioides<br />
P. amarus Hutch., in herb. =Pogonopus tubulosus<br />
P. bolivianus Britt., in herb. =Pogonopus tubulosus Schizocalyx =Bathysa<br />
P. erythroxylon (Standl.) J.H.Kirk. =Simira ecuatoriensis S. bracteosus Wedd. =Bathysa bracteosa<br />
P. febrifugus (Wedd.) J. D.Hook. =Pogonopus tubulosus S. hirsutus Standl. =Bathysa bracteosa<br />
P. febrifugus (Wedd.) J. D.Hook. var. macrosema Hutch. S. pohliana Berg (unpubl.?) =Faramea sp.<br />
=Pogonopus tubulosus<br />
Schreibersia Kuntze =Augusta<br />
P. exsertus (Orst.) Orst.<br />
S. longifolia (Spreng.) Kuntze =Augusta longifolia var.<br />
P. moritzii Klotch., in herb. =Pogonopus speciosus var. longifolia<br />
speciosus<br />
Semaphyllanthe L. Andersson =Calycophyllum<br />
P. ottonis Klotch. =Pogonopus speciosus var. speciosus Sickingia =Simira Aubl.<br />
P. speciosus (tubulosus) var. ottonis K. Schum., in herb. S. catappiifolia Standl. =Simira macrocrater<br />
=Pogonopus speciosus var. speciosus<br />
S. cordifolia J. D.Hook. =Simira cordifolia<br />
P. speciosus (Jacq.) K.Schum.<br />
S. ecuadorensis Standl. =Simira ecuadorensis<br />
P. speciosus (Jacq.) K.Schum. var. speciosus<br />
S. erythroxylon sensu Oliv. =Simira glaziovii<br />
P. speciosus (Jacq.) K.Schum. var. sandwithianus Steyerm. S. erythroxylon Willd. =Simira erythroxylon<br />
P. speciosus (Jacq.) K.Schum. subsp. sandwithianus Steyerm. S. fragrans (Rusby) Standl. =Simira fragrans<br />
= Pogonopus speciosus var. sandwithianus<br />
S. goudotii (Baill.) Standl. =Simira goudotii<br />
P. speciosus (Jacq.) K.Schum. subsp. speciosus var. S. longifolia Willd. =Simira longifolia<br />
sandwithianus Steyerm. =Pogonopus speciosus var. S. myriantha Standl. =Elaeagia myriantha<br />
sandwithianus Steyerm.<br />
S. paraensis (Baill.) K.Schum. =Simira paraensis<br />
P. speciosus (Jacq.) K.Schum. subsp. speciosus var. S. pisoniiformis (Baill.) K.Schum. =Simira pisoniiformis<br />
speciosus =Pogonopus speciosus var. speciosus S. tinctoria (Aubl.) Lem6e =Simira tinctoria<br />
P. speciosus (Jacq.) K. Schum. subsp. exsertus (Orst.) S. tinctoria (Kunth) K.Schum. =Simira rubescens<br />
Steyerm. =Pogonopus exsertus<br />
Simira Aubl.<br />
P. speciosus [tubulosus] (Jacq.) K.Schum. var. panamensis S. catappiifolia (Standl.) Steyerm. =Simira macrocrater<br />
K.Schum., in herb. =Pogonopus exsertus<br />
S. cordifolia (J. D.Hook.) Steyerm.<br />
P. tubulosus (A.Rich. in DC.) K.Schum.<br />
S. ecuadorensis (Standl.) Steyerm.<br />
Pseudochimarrhis Ducke =Chimarrhis subgen. S. erythroxylon (Willd.) Bremek.<br />
Pseudochimarrhis<br />
S. fragrans (Rusby) Steyerm.<br />
P. barbata Ducke =Chimarrhis barbata<br />
S. glaziovii (K.Schum.) Steyerm.<br />
P. difformis (Benoist) Benoist =Chimarrhis turbinata S. goudotii (Baill.) Steyerm.<br />
P. turbinata (DC.) Ducke =Chimarrhis turbinata S. longifolia (Willd.) Steyerm.<br />
S. macrocrater K.Schum.<br />
Rogiera Planch. =Rondeletia<br />
Roigella Borhidi & Z. M. Femrnndez =Rondeletia<br />
Rondeletia L.<br />
S. myriantha (Standl.) Steyerm. =Elaeagia myriantha<br />
S. paraensis (Baill.) Steyerm.<br />
S. pisoniiformis (Baill.) Steyerm.
220 Flora Neotropica<br />
S. rhodoclada (Standl.) Steyerm.<br />
S. rubescens (Benth.) Bremek. ex Steyerm.<br />
S. rubra (Mart.) Steyerm.<br />
S. tinctoria Aubl.<br />
S. viridiflora (Allemao & Saldanha) Steyerm.<br />
Simira Raf. =Ornithogalum L. (Liliaceae)<br />
Sprucea Benth. =Simira Aubl.<br />
S. rubescens Benth. =Simira rubescens<br />
Stomandra Standl. =<strong>Rustia</strong><br />
S. costaricensis Standl. =<strong>Rustia</strong> costaricensis<br />
Suberanthus Borhidi & Z. M. Fernmndez =Rondeletia?<br />
S. pungens (Urb.) Borhidi & Z. M. Fernandez<br />
=Rondeletia sp.?<br />
agua fresca (6-8)<br />
amarelinha (6-12)<br />
amarilla (6-8)<br />
anime chiquita (3-1)<br />
aromuhe (6-7)<br />
bois chapelle (6-10)<br />
bois de riviere (6-5)<br />
bois des bams-jaunes (6-5)<br />
bois pagaie (6-10)<br />
bois riviere (6-5)<br />
bois riviere r6solu (6-5)<br />
bouganvillea (5-3)<br />
caapeba (1-11)<br />
cacahuito (3-1)<br />
cambur6n (2-lb)<br />
carapanauba (6-10)<br />
carapana6ba cinzeiro (6-10)<br />
carutillo (6-8)<br />
cascarilla (5-3)<br />
cascarilla masha (6-6; 1-8)<br />
cascarillita (1-8)<br />
caspi blanco (3-1)<br />
cera (6-3)<br />
chibigui (5-1)<br />
chollachaqui caspi blanco (6-7)<br />
chorcha de gallo (5-1)<br />
chullachasqui (3-1)<br />
citronelle (6-10)<br />
citronelle rouge (6-10)<br />
coralito (5-2a)<br />
doncello (1-9)<br />
flor de hoja (5-3)<br />
guamo (3-1)<br />
guapeba branca (1-11)<br />
INDEX OF LOCAL NAMES<br />
higueron (6-1)<br />
hilacho (6-3)<br />
huacamayo micuna (10-1)<br />
huacap6 masha (6-7)<br />
intacchi (6-7)<br />
itahuba (6-7)<br />
itauba (6-6, 6-7)<br />
itauba amarilla (6-6, 6-7)<br />
jagua amarilla (6-2)<br />
jatun mincha caspi (6-6)<br />
jilacho (6-3)<br />
kaire-e<br />
(6-10)<br />
koemaramaraballi<br />
(6-10)<br />
larangui (3-1)<br />
lengua de vaca (3-1)<br />
llema de huevo (6-1)<br />
madrofio (5-1)<br />
man wood (6-1)<br />
mecha caspi (6-6)<br />
mincha caspi (6-6)<br />
mukugd (6-7)<br />
mun baikuanin (3-1)<br />
naranjillo montafiero (5-2a)<br />
pablo manchana (6-6; 6-7)<br />
palo marfil (5-3)<br />
palo palillo (6-6)<br />
palo perro (6-7)<br />
pampa remo caspi (6-7)<br />
pampa remocaspi (6-6)<br />
papelillo caspi (6-7)<br />
paraka'y (6-10)<br />
paraku-'iran-'y (6-10)<br />
pau amarelo (6-12)<br />
pau de remo (6-10)<br />
<strong>Tresanthera</strong> H.Karst.<br />
T. condamineoides H.Karst. var. condamineoides<br />
T. condamineoides H.Karst. var. thyrsiflora (Steyerm.)<br />
Delprete<br />
T. pauciflora K.Schum. ex Soler. =<strong>Tresanthera</strong><br />
condamineoides var. thyrsiflora<br />
T. thyrsiflora Steyerm. =<strong>Tresanthera</strong> condamineoides var.<br />
thyrsiflora<br />
Ucriana Spreng. =Augusta<br />
U. longifolia Spreng. =Augusta longifolia var. longifolia<br />
Wernhamia S. Moore =Simira Aubl.<br />
W. boliviensis S. Moore =Simira macrocrater K.Schumann<br />
pejiballito (6-1)<br />
penda (6-3)<br />
pesquim (8- b)<br />
purma-caspi (6-7)<br />
quillo-bordon (6-7)<br />
quillobordon (10-1)<br />
quina (5-1; 5-2a; 5-3)<br />
quina blanca (7-1)<br />
quina do Rio de Janeiro (1-11)<br />
quina morada (5-2b)<br />
quinoyer (6-5)<br />
ramal do pau rosa (6-10)<br />
river wood (6-5)<br />
resolu de montagne (6-5)<br />
resolu de riviere (6-5)<br />
roble amarillo (6-3)<br />
sacha cascarilla (1-8)<br />
sachajagua (6-7)<br />
sacha quina (5-3)<br />
saco de mono (8-la)<br />
samaati-palioudou (6-10)<br />
santonino (6-9)<br />
sienjahoedoe (6-10)<br />
sinjahoedoe (6-10)<br />
sobrasil (1-11)<br />
totumo amarillo (6-8)<br />
tuwara (6-7)<br />
walalu (6-10)<br />
waterwood (6-5)<br />
white calabash (6-8)<br />
wild fiddle wood (6-4)<br />
yacu-caspi(6-7)<br />
yema de huevo (6-1; 6-2)<br />
yerno prueba (6-6; 6-7)
Index of Scientific Names 221<br />
INDEX OF SCIENTIFIC NAMES<br />
Synonyms are in italics. Names in [square brackets] are nomen nuda or unpublished names. Page numbers in<br />
boldface indicate primary references. Page numbers with an asterisk (*) indicate pages with illustrations or maps.<br />
Acrobotrys, 8, 14, 22<br />
Acrosynanthus, 14, 16, 21<br />
Acunaeanthus, 8, 13, 14, 21<br />
Aleisanthia, 3, 9, 18<br />
Alibertia, 96<br />
isernii, 58, 96<br />
Allenanthus, 12*<br />
erythrocarpus, 141, 185<br />
Angiosperms, 7<br />
Anthospermeae, 53<br />
Antirheoideae, 8, 10<br />
Aparines, 4<br />
Aparinia, 5<br />
Arachnothryx, 9, 10, 13<br />
Ardisia, 80<br />
Ariadne, 13<br />
shaferi, 13<br />
Aristolochia, 121<br />
Aristolochiaceae, 121<br />
Augusta, 6, 9, 11, 12*, 13, 14, 16, 18<br />
subgen. Augusta, 22<br />
subgen. Lindenia, 16, 22<br />
longifolia, 16<br />
Augusta group, 13<br />
Badusa, 8, 12*, 18<br />
Bathysa, 6,9,12*, 13,14,21,58,96,105,116,138,186,187<br />
autralis, 58<br />
bathysoides, 141, 185<br />
bracteosa, 13, 20<br />
cuspidata, 58<br />
difformis, 140, 169, 175<br />
perijaensis, 186<br />
pittieri, 140, 187<br />
Bathysa group, 13<br />
Bikkia, 3, 6, 8, 12*, 29, 42, 52, 53, 54<br />
campanulata, 53<br />
palauensis, 53<br />
Blandibractea, 9, 17<br />
brasiliensis, 17<br />
Bradea, 17<br />
Buena muzonensis, 96<br />
Calycophylleae, 9, 138<br />
Calycophyllum, 9, 29, 53, 122, 138, 175, 185<br />
candidissimum, 140<br />
obovatum, 185<br />
tubulosum (A. Rich.) DC., 122, 123, 133, 136<br />
tubulosum Seeman, 123<br />
Capirona, 53, 137<br />
decorticans, 116<br />
erythroxylon, 136, 137<br />
Carmenocania, 121, 122<br />
porphyrantha, 121, 130, 131<br />
Carphalea, 6<br />
Carphalia, 6<br />
Cascarilla, 6<br />
muzonensis, 96<br />
sect. muzonia, 96<br />
Catesbaea, 3, 9, 11, 12*<br />
Catesbaeeae, 3, 5*, 9, 10, 11, 12*, 18, 42, 53<br />
Catesbaeeae-Chiococceae complex, 3, 16<br />
Cephalanthia, 5<br />
Ceratina, 54, 84<br />
Ceuthocarpus, 3, 8, 12*, 29<br />
Chalepophyllum, 6, 9, 14, 19, 22<br />
[Chimarhidia], 18<br />
[Chimarhis], 137<br />
[Chimarrheae], 7, 18<br />
[Chimarrhidoideae], 18<br />
Chimarrhis, 4,5,6,7, 8, 10, 11, 12*, 14, 16, 17, 18, 19,20,<br />
23, 24, 25, 26*, 27, 29, 30, 31*, 32, 33, 38, 39*,<br />
40*,48,50*, 51*, 52,53,54,55,56,137,138,139*,<br />
140, 141,150, 155, 158, 159, 169, 175, 177, 177*,<br />
183, 185, 186, 187, 188, 200<br />
subgen. Chimarrhis, 25, 138, 140, 141, 142<br />
subgen. Pseudochimarrhis, 25,137,138,140,141,169,175<br />
barbata, 23, 24, 27, 29, 32, 38, 48, 55, 56, 138, 139*,<br />
141,142, 159, 175, 176, 177, 177*, 178*, 181<br />
bathysoides, 141, 185<br />
brevipes, 24,29,38,40*, 137, 138, 139*, 141,142,177*,<br />
181,182*, 183, 185<br />
[corymbosa], 140, 152, 155<br />
clausicorollata, 185<br />
cubensis, 23, 24, 32, 38, 39*, 48, 50*, 55, 138, 139*,<br />
141,147,148*, 149*, 150, 152, 164<br />
cymosa, 23, 25, 26*, 27, 28, 32, 38, 39*, 48,50*, 53, 56,<br />
137,138,139*,140,141,147,149*,150,152,153*,<br />
154,155,166<br />
subsp. genuina, 140, 152, 155<br />
subsp.jamaicensis, 140, 141, 150<br />
subsp. microcarpa, 140, 141, 147, 164<br />
var.jamaicensis, 150<br />
var. microcarpa, 147, 164<br />
[cyrrhosa], 152<br />
decurrens, 141, 185<br />
dioica, 140, 141, 185, 187, 192<br />
duckeana,27,32,38,40*,51*, 138,142,177, 177*, 179,<br />
180*, 181<br />
duckei, 141, 169, 175, 179<br />
ferruginea, 140, 141, 186<br />
gentryana,24,29,32,38,40*, 137,138,139*, 142,177*,<br />
183,184*,185<br />
glabriflora, 23, 24, 26*, 27, 28, 31*, 38, 39*, 54, 55, 56,<br />
138, 139*, 141,155, 156*, 157*, 158, 159, 162*,<br />
163, 177*<br />
goudotii, 141, 186<br />
hookeri, 23, 24, 27, 38, 39*, 48, 51*, 53,54, 55, 56, 138,<br />
139*, 140, 141, 142, 157, 158, 159, 160*, 161*,<br />
162*, 163, 164, 177*<br />
jamaicensis, 23,24,32,48, 50*, 56, 137, 138, 139*, 141,<br />
149*,150,151*,152<br />
latifolia, 23, 24, 32, 38, 39*, 48, 50*, 54, 55, 56, 138,<br />
139*, 142, 144, 145*, 146*<br />
longifolia, 186<br />
longistipulata, 53, 141, 164, 166<br />
microcarpa, 23, 28, 32, 38, 48, 53, 56, 138, 139*, 140,<br />
141,144, 164, 165*, 166, 169, 169*, 183<br />
var. microcarpa, 141, 164<br />
var. speciosa, 141, 167<br />
paraensis, 186
222 Flora Neotropica<br />
Chimarrhis (continued)<br />
Cubanola, 3, 8, 12*, 29<br />
parviflora, 22, 28, 32, 28, 39, 48, 50*, 56, 139*, 141,<br />
142, 143*, 146*, 147, 169<br />
perijaensis, 186<br />
pisoniaeformis, 141, 186<br />
pittieri, 140, 141, 187<br />
[sessile], 187<br />
speciosa, 32, 38, 39*, 48, 51*, 141, 164, 167, 168*, 169,<br />
169*, 183<br />
Dendrosipanea, 9, 14, 22<br />
Deppea, 6<br />
Dilaria, 5<br />
Dioicodendron, 4, 8, 10, 11, 12*, 14, 18, 20, 23, 24, 25,<br />
26*, 27, 28, 29, 30, 31*, 32, 33, 38, 41*, 48, 51*,<br />
55, 140, 141,187, 188, 189*, 190*, 192<br />
cuatrecasasii, 188, 192<br />
turbinata, 23,25,26*, 27,29,32, 38,40*, 48,51*, 53, 55,<br />
56, 138, 139*, 140, 141, 142, 158, 159, 169, 170*,<br />
171,172*, 174*, 175, 177, 177*, 179, 181, 183<br />
venezuelensis, 141, 183, 187, 188, 192<br />
williamsii, 141, 159, 164<br />
dioicum,26*,31*,41*,48,51*, 140,183,186,187,188,<br />
189*, 190*, 191*<br />
Dolichodelphys, 4, 11, 12*, 14, 20, 23, 24, 25, 26*, 27,<br />
28, 29, 30, 32, 33, 38, 43*, 48, 52*, 55, 191*,<br />
196, 197, 198*<br />
Chiococca, 9, 12*<br />
chlorocrater, 26*, 43*, 48, 52*, 58, 85, 96, 196, 197,<br />
Chiococceae, 4, 5*, 9, 11, 12*, 13, 16<br />
198*, 199<br />
Chione, 12*<br />
Dolicholobium, 42<br />
Chrysoxylon Casaretto, 121<br />
Chrysoxylon Weddell, 121, 122<br />
Eizia, 9, 17<br />
febrifugum, 121, 122, 133, 136<br />
speciosum, 130<br />
tubulosum, 133<br />
Cigarilla, 8, 9<br />
Cinchona, 9, 11, 12*, 53, 105<br />
muzonensis, 96<br />
philippica<br />
Cinchona Series, 140<br />
Elaeagia, 6, 9, 14, 16, 17, 18, 19, 38, 53, 116, 117, 187<br />
brasiliensis, 169, 175<br />
cubensis, 19<br />
grandis, 187<br />
myriantha, 16<br />
Endogynie Rafinesque, 5<br />
Eosanthe, 16<br />
Ericaceae, 42<br />
Cinchonaceae, 6<br />
Erithalis, 9, 13*<br />
Cinchoneae, 6, 8, 9, 11, 17, 18, 28, 42, 53, 54, 117, 122, Eucondamineae, 3, 6, 18, 140<br />
138, 140, 175, 193<br />
Cinchoninae, 7<br />
Cinchonoideae, 7, 8, 9, 10, 11, 197<br />
Euglossa, 54, 64*<br />
Eupinckneya, 122<br />
Exandra, 16<br />
Clusia, 80<br />
rhodoclada<br />
Coffea, 9, 11<br />
Exostema, 8, 9, 12*, 53, 88<br />
floribunda Martius, 193<br />
australe, 58<br />
floribunda Miquel, 193, 195<br />
stipulata, 13<br />
Coffeae, 9, 11<br />
Coleoptera, 55, 81<br />
Condaminea, 4, 6, 8, 9, 10, 11,12*, 13, 14,18, 19,23,24,25,<br />
cuspidatum, 58<br />
formosum, 58, 85, 86, 88<br />
var. laeve, 85, 88, 89<br />
var. leprosum, 58, 85, 88<br />
occidentale, 58, 62, 65<br />
26*, 27, 28, 29, 30, 31*, 32, 33, 34, 36*, 44,47*, 48, souzanum, 58<br />
53,54,55,57,58,98,103,104*, 105,114,116,117 Exostemma, 58, 88<br />
angustifolia, 105<br />
breviflora, 105<br />
corymbosa, 23, 24, 26*, 31*, 33, 36*, 47*, 48, 53, 54,<br />
sect. Pseudostemma, 58, 62, 65<br />
formosum, 58<br />
occidentale, 25, 58, 62, 65<br />
55,98,99,103,104*,105,106*,107*,108*, 109*,<br />
110, 112*, 113, 114, 116,117,140<br />
var. pubescens, 105, 114<br />
elegans, 33, 36*, 48, 103, 105, 112*, 114, 115*<br />
glabrata, 105, 116, 159<br />
macrophylla, 105, 116<br />
microcarpa, 105, 116<br />
petiolata, 105, 116<br />
tinctoria, 105, 116<br />
Ferdinandusa, 53<br />
Ficus, 80<br />
Flexanthera, 8, 10, 16<br />
fragrans, 16<br />
subcordata, 16<br />
[Forsteronia], 192, 193<br />
[panniculata], 193<br />
utilis, 116<br />
venosa, 105, 117<br />
Gardenieae, 11, 38, 53, 197<br />
Geonoma, 80<br />
Condamineeae, 3, 4, 5*, 6, 7, 8, 9, 10, 11, 12*, 16, 17, 18, Gleasonioideae, 10<br />
19, 23, 25, 26*, 29, 31*, 38, 42, 52, 54, 122, 138,<br />
140, 193<br />
Glionnetia, 3, 9, 18<br />
Gloneria, 6, 9, 18<br />
Condamineinae, 3, 4, 6, 7, 10, 11, 18, 42, 140<br />
Condaminieae, 6, 18<br />
Couepia, 80<br />
Coutaportla, 3, 8, 12*, 29<br />
Coutarea, 3, 8, 9, 12*, 29, 53<br />
Cuatrecasasiodendron, 9, 12*, 14, 18, 20, 22, 41*, 42<br />
colombianum, 41*<br />
Greenea, 6, 9, 18<br />
Greenia, 6<br />
Grias, 80<br />
Guarea, 80<br />
Guettardeae, 53<br />
Guettardoideae, 7, 10<br />
Gyranthera caribensis, 101
Index of Scientific Names 223<br />
[Hamelia albopurpurea], 84<br />
Hamelieae, 6, 17, 53<br />
Hedyoteae, 6<br />
Hedyotideae, 6, 8, 17, 18, 42, 53, 140<br />
Hedyotidinae, 6<br />
Henlea Grisebach, 57<br />
Henlea H. Karsten, 6, 7, 25, 57, 58, 84, 96, 97<br />
muzonensis, 96<br />
rosea, 58, 96<br />
splendens, 25, 58, 81, 84, 85<br />
thibaudioides, 57, 58, 81, 84<br />
Hillieae, 53<br />
Hillioideae, 10<br />
Hintonia, 3, 8, 12*, 29<br />
Hippotis, 13<br />
Hippotieae, 13<br />
Holstianthus, 9, 14, 22<br />
Holtonia, 16<br />
Howardia Klotzsch, 121<br />
Howardia Weddell, 121, 122, 136<br />
caracasensis, 122, 130<br />
febrifuga, 56, 121, 122, 133, 136<br />
grandiflora, 122, 130, 131<br />
richardi, 122, 133, 136<br />
Hymenocnemis, 25<br />
Iriartera fusca, 101<br />
Isertieae, 8, 9, 10, 18, 53, 197<br />
Isidorea, 3, 6, 8, 12*, 29<br />
Ixora, 186, 187, 193<br />
densiflora, 193<br />
podocarpa, 186, 187<br />
Ixoreae, 13, 53<br />
Ixoroideae, 3, 8, 9, 10, 11<br />
Javorkaea, 9, 10, 13<br />
Joosia, 9, 11, 12*<br />
Kajewskiella, 8, 10, 12*, 18, 26*, 27, 42, 43*, 48, 52*<br />
polyantha, 48, 52*<br />
trichantha, 26*, 42,43*<br />
Kerianthera, 8, 10, 12*, 18, 26*, 27, 31*, 41*, 49*<br />
preclara, 26*, 31*, 41*, 42, 48, 49*<br />
Ladenbergia, 6, 58<br />
muzonensis, 58, 96<br />
Larospermia, 5<br />
Lerchea, 18<br />
Limnosipanea, 9, 14, 17, 19<br />
schomburgkii, 19<br />
Lindenia, 6, 9, 11, 12*, 13, 16, 18<br />
Mabea, 80<br />
Macbrideina, 9, 14, 21, 117<br />
Machaonia, 13<br />
Macrocnemum P. Browne, 32, 105, 117, 140, 155<br />
Macrocnemum sensu A. Richard, 140<br />
candidissimum, 140<br />
corymbosum, 105, 109*, 113, 140<br />
cubense, 118, 120, 121<br />
exsertum, 122, 123<br />
glabratum, 116<br />
jamaicense, 140<br />
longifolium, 140, 152, 155<br />
microcarpon, 116<br />
speciosus, 122, 130, 131<br />
tetrandrum, 140<br />
tinctorium, 105, 116<br />
tubulosum, 122, 133, 136<br />
venosum, 117<br />
Maguireothamnus, 9, 14, 22<br />
Malanea grandis, 187<br />
Malpighiaceae, 57<br />
Mazaea, 4, 10, 13, 14, 19, 21<br />
shaferi, 13<br />
Mazaea group, 13<br />
Melastomataceae, 30<br />
Melipona, 54, 64*, 84<br />
Mimosaceae, 121<br />
Molopanthera,4, 9, 11, 12*, 14, 18, 20,23,24,25,26*, 27,<br />
28,29,30,31*,32,33,38,41*,55,192,193<br />
burchellii, 195<br />
[colombiana], 192<br />
paniculata, 26*, 32, 41*, 192, 193, 194*, 195, 196*<br />
var. paniculata, 26*, 31*, 194*, 195, 196*<br />
var. burchellii, 41*, 195, 196*<br />
panniculata, 193, 195<br />
var. [scabrida], 196<br />
Morierina, 6, 8, 12*, 53<br />
montana, 53<br />
Morindeae, 53<br />
Mussaenda, 9, 18<br />
Mussaenda speciosa, 130<br />
Mussaendeae, 53, 122<br />
Mussaendopsis, 53<br />
Naucleeae, 4, 53<br />
Neblinathamnus, 9, 14, 22<br />
Neobertiera, 9, 10, 17<br />
Neomazaea, 13<br />
Nerstia, 3, 8, 9, 12*, 29<br />
Ochroma, 147<br />
Onagraceae, 42<br />
Osa, 3, 8, 29<br />
Otocalyx, 13<br />
Paederieae, 53<br />
Pallasia, 6<br />
Parachimarrhis, 4, 8, 10, 12*, 14, 19, 21, 28, 29, 30, 32, 33,<br />
38,42,43*, 48,51*, 55,56,199,200,201*, 202*<br />
breviloba, 43*, 48, 51*, 199, 200, 201*, 202*, 202<br />
Phialanthus, 13, 16<br />
Phitopis, 7, 13, 14, 22<br />
Phyllacanthus, 3, 11, 12*<br />
Phyllomelia, 4, 13, 16, 19<br />
Picardaea, 4, 8, 10, 11, 12*, 14, 19,20,23,24,25,26*,27,<br />
28, 29, 30, 31*, 32, 33, 37*, 47*, 48, 52, 55, 117,<br />
118, 120<br />
cubensis,26*,31*,37*,47*,48,117,118,119*, 120*, 121<br />
haitiensis, 117, 118, 120, 121<br />
Pinckneya L.C.M. Richard, 3, 6, 8, 9, 10, 11, 12*, 14, 17,<br />
18, 19, 25, 26*, 27, 29, 30, 31*, 32, 37*, 41, 48,<br />
49*, 52, 53, 54, 55, 121,122<br />
Pinckneya sensu Saldanha & Allemao, 7<br />
bracteata (W. Bartram) Rafinesque, 18,23,26*, 31*, 37*,<br />
49*, 53<br />
erubescens, 17<br />
ionantha, 131, 132, 133<br />
pubens, 53<br />
rubescens, 17
224 Flora Neotropica<br />
Pinckneya sensu Saldanha & Allemao (continued)<br />
viridiflora, 17<br />
Pinckneyinae, 3, 4, 6, 11, 18, 122<br />
Pinckneyeae, 3, 18<br />
Pinkneya, 6<br />
Placocarpa, 12*<br />
Plathymenia, 121<br />
Pogonopus, 4,6,8,9, 10, 11, 12*, 14, 19,23,24,25,26*, 27,<br />
28,29,30,31*,32, 33,34,37*,38,42, 48,49*, 52,<br />
53,54,55,56, 117,121,122,123, 131,136, 137<br />
[amarus], 136<br />
[bolivianus], 136<br />
caracasensis, 130<br />
erythroxylon, 136<br />
febrifugus, 133, 136<br />
var. macrosema, 133, 136<br />
exsertus, 30, 37*, 38, 48, 49*, 55, 56, 122, 123, 124*,<br />
125*, 126, 127<br />
[moritzii], 131<br />
ottonis, 121, 122, 130<br />
speciosus, 23, 24, 36*, 30, 31*, 37*, 38, 48, 49*, 53, 54,<br />
55,56,121,122,123,125*, 126,127,128*, 129,133<br />
subsp. exsertus, 122, 123<br />
subsp. sandwithianus, 122<br />
subsp. speciosus, 122, 132, 133<br />
var. sandwithianus, 122, 132<br />
var. speciosus, 130<br />
var. speciosus, 37*, 38, 121, 125*, 127, 128*, 130,<br />
131,133<br />
var. sandwithianus, 23, 24,26*, 31*, 37*, 38,48,49*,<br />
54, 55, 125*, 127, 128*, 129*, 130, 131,132<br />
[var. ottonis], 131<br />
[var. panamensis], 127<br />
tubulosus, 30, 37*, 38, 48, 49*, 55, 56, 121, 122, 123,<br />
125*, 126, 133, 134*, 136<br />
[var. ottonis], 131<br />
[var. panamensis], 127<br />
[var. boliviensis], 136<br />
Polarnia, 5<br />
Polyspia, 5<br />
Pomazota, 18<br />
Pomazotoideae, 10<br />
Portlandia, 3, 6, 8, 9, 12*, 29, 53, 54, 193<br />
grandiflora, 53<br />
Portlandia-group, 9<br />
Portlandia series, 6<br />
Portlandieae, 6, 10<br />
Portlandiinae, 3, 6, 8, 9, 10, 11, 18, 29, 42<br />
Pseudochimarrhis, 53, 137, 140, 141, 169, 175<br />
barbata, 140, 141, 175, 176<br />
difformis, 140, 141, 169<br />
turbinata, 137, 140, 141, 169<br />
Pseudomussaenda, 8, 10, 18<br />
Psychotria, 6, 9, 18<br />
Psychotrieae, 18, 53<br />
Pteridocalyx, 9, 14, 17, 20<br />
Raritebe, 197<br />
Retiniphyllum, 197<br />
Rogiera, 9, 10, 13<br />
Roigella, 9, 13<br />
Rondeletia, 4, 6, 9, 10, 12*, 13, 18, 42, 116<br />
brandegeeana, 13<br />
panamensis, 105, 116<br />
Rondeletia-complex, 13, 14, 20, 22<br />
<strong>Rondeletieae</strong>, 3,4,5*, 6, 7,8,9, 10, 11, 12*, 13, 16, 17, 18,<br />
19, 29,38, 42, 53, 54, 116, 117, 138, 140, 197<br />
Rondeletiinae, 6, 18, 140<br />
<strong>Rubiaceae</strong>, 3, 4, 5, 6, 7, 8, 9, 10, 24, 25, 27, 29, 30, 32, 33,<br />
52,53,54,56,57, 97, 116, 117,138, 150, 187, 193<br />
Rubieae, 53<br />
Rubioideae, 3, 7, 8, 9, 10, 11<br />
<strong>Rustia</strong>, 3,4,6,7,8,10,11,12*, 14,16,19,20,23,24,25,26*,<br />
27, 28, 29, 30, 31*, 32, 33, 34*, 35*, 38,42,44,45*,<br />
46*,53,54,55,56,57,58,59,78,84,85,88,89,91,<br />
96,97,98,99, 103,114,116,117,141,186, 199<br />
alba, 23, 24, 26*, 27,28, 30,33, 35*, 44,45*, 57, 58, 59,<br />
67, 68*, 69*, 70, 74*, 88<br />
angustifolia, 24, 25, 28, 30, 33, 35*, 44, 46*, 58, 59, 91,<br />
92, 92*, 93*, 95<br />
bilsana, 23, 24, 28, 29, 30, 54, 57, 58, 70, 71*, 72*, 74*<br />
costaricensis, 23, 28, 30, 31*, 33, 34*, 44, 45*, 54, 57,<br />
58,59,60*, 61*, 62*<br />
dressleri, 23, 24,28,29,30,44,54,57,58,59,61*, 65,66*<br />
ferruginea, 58, 96, 140, 186<br />
formosa, 23,24,25,26*, 27,28,30,33, 34*, 44,46*, 53,<br />
57,58,59,70,85, 87*, 88, 89, 91, 92*<br />
gracilis, 25, 28, 30, 33, 34*, 44, 46*, 57, 58, 59, 89, 91,<br />
92, 92*, 94*, 95<br />
haitiensis, 58, 95<br />
isernii, 58, 96<br />
longifolia, 58, 85, 96, 197, 199<br />
mosenii, 86<br />
occidentalis, 23, 24,25, 26*, 27,28,29,30,31 *, 33, 34*,<br />
44, 45*, 54, 55, 57, 58, 59, 62, 63*, 64*, 65, 70,<br />
74*,98<br />
panamensis, 58, 59, 62<br />
pauciflora, 25, 96, 99<br />
pohliana, 58, 86, 88<br />
rosea, 58, 96<br />
rubra, 23, 28, 30, 33, 34*, 44, 45*, 54, 57, 59, 72, 74*,<br />
75*, 76, 80<br />
schunkeana, 28, 30, 33, 35*, 44, 45*, 54, 55, 57, 58, 59,<br />
74*,76,78,79*,80,84<br />
secundiflora, 58, 81, 84<br />
sellowiana, 58, 85, 88<br />
simpsonii, 24, 28, 30, 32, 44, 46*, 57, 59, 88, 89, 90*,<br />
91,92*, 95<br />
splendens, 76, 81, 82, 84, 199<br />
thibaudioides, 23, 24, 25, 28, 30, 31 *, 33, 35*, 44,45*,<br />
54, 55, 57, 58, 59, 74*,76,80,81,82*,83*,84,<br />
85, 199<br />
venezuelensis, 23, 24, 28, 33, 35*, 58, 59, 74*, 85<br />
[veraguensis], 62<br />
viridiflora, 23, 28, 30, 33, 35*, 44, 46*, 54, 57, 58, 59,<br />
74*, 76, 77*, 78, 80<br />
[warscewicziana], 84, 85<br />
Sambucidia, 5<br />
Schizocalyx, 13<br />
Schizolobium, 147<br />
Schizomussaenda, 18<br />
Schmidtottia, 3, 8, 12*, 16, 29<br />
Scolosanthus, 13*<br />
Sickingia, 7, 16, 105, 117, 140, 141, 186<br />
ecuadorensis, 137<br />
erythroxylon, 16, 136<br />
goudotii, 186<br />
longifolia, 16, 186<br />
myriantha, 16
Index of Scientific Names 225<br />
paraensis, 186<br />
pisoniiformis, 186, 187<br />
rubra, 17<br />
tinctoria (Kunth) K. Schumann, 116<br />
viridiflora, 17<br />
Siemensia, 3, 8, 29<br />
SimiraAublet, 7,9,10,12*, 14,16,17,18,19,21,38,42,43*,<br />
105,117,137,138,140,141,185,186,200<br />
catappiifolia, 17<br />
cordifolia, 16<br />
ecuadorensis, 136, 137<br />
erythroxylon, 136<br />
fragrans, 16<br />
glaziovii, 17<br />
goudotii, 186<br />
longifolia, 43*, 186<br />
macrocrater, 17, 20<br />
paraensis, 186<br />
pisoniiformis, 186<br />
rhodoclada, 16<br />
rubescens, 105, 116<br />
rubra, 17<br />
tinctoria Aublet, 116, 185<br />
viridiflora, 18<br />
Simira group, 16, 17<br />
Simireae, 5*, 10, 16, 18<br />
Sipanea, 4, 9, 14, 17, 18, 19<br />
biflora, 19<br />
Sipanea group, 17, 19<br />
Sipaneeae, 4, 5*, 7, 9, 10, 11, 13, 17, 18<br />
Sipaneopsis, 8, 9, 14, 17, 19, 20, 21<br />
Solanaceae, 30<br />
Spathichlamys, 3, 9, 18<br />
Sphanidia, 5<br />
Sprucea, 140<br />
Standleya, 17<br />
Steenisia, 9, 18<br />
Stevensia, 9, 14, 16, 21<br />
Steyermarkia, 9, 14, 17, 19<br />
Stomandra, 8, 10, 57, 58<br />
costaricensis, 57, 58, 59, 62<br />
Suberanthus, 13<br />
Synisoon, 197<br />
Syringantha, 9<br />
Thogsennia, 3, 8, 12*, 29<br />
Tocoyena, 197<br />
<strong>Tresanthera</strong>, 3, 4,6, 7, 8, 10, 11,12*, 14,19,20,23,24,25,<br />
26*, 27, 28, 29, 30, 31*, 33, 36*, 42, 44, 47*, 54,<br />
55, 57, 58, 96, 97, 98, 99<br />
condamineoides, 25, 26*, 28, 31*, 33, 36*, 44, 47*, 58,<br />
96, 97, 98, 99, 100*, 101, 103, 199<br />
var. condamineoides, 26*, 28, 31 *, 33, 36*, 44, 47*,<br />
96, 99,100*, 101<br />
var. thyrsiflora, 28, 33, 36*, 44, 47*, 99, 101, 102*<br />
pauciflora, 25, 97, 98, 99<br />
thyrsiflora, 97, 99, 101<br />
Trigona, 54, 84<br />
Urophylloideae, 10<br />
Vanguerieae, 53<br />
Viburidia, 5<br />
Warszewiczia, 6, 9, 14, 20, 53, 116, 187<br />
schwackei, 20<br />
Wendlandia, 3, 9, 18<br />
Wernhamia, 9, 17<br />
boliviensis, 17<br />
Wittmackanthus, 6<br />
Xanthophytum, 18, 42