Citation
Lankesteriana

Material Information

Title:
Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Creator:
Jardi´n Bota´nico Lankester
Jardín Botánico Lankester
Place of Publication:
Cartago Costa Rica
Publisher:
Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Publication Date:
Frequency:
Three times a year[2002-]
Irregular[ FORMER 2001]
three times a year
regular
Language:
English
Physical Description:
v. : ill. (some col.) ; 25 cm.

Subjects

Subjects / Keywords:
Botany -- Periodicals -- Costa Rica ( lcsh )
Epiphytes -- Periodicals -- Costa Rica ( lcsh )
Orchids -- Periodicals -- Costa Rica ( lcsh )
Plantkunde ( gtt )
Botanische tuinen ( gtt )
Genre:
periodical ( marcgt )
serial ( sobekcm )
Spatial Coverage:
Costa Rica

Notes

Language:
In English and Spanish.
Dates or Sequential Designation:
No. 1 (mayo 2001)-
Numbering Peculiarities:
Issues for May 2001-Oct. 2003 designated no.1-8; issues for Apr. 2004- designated vol. 4, no. 1-
General Note:
Latest issue consulted: Vol. 4, no. 1 (abr. 2004).
General Note:
International journal on orchidology.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright Jardín Botánico Lankester, Universidad de Costa Rica. Permission granted to University of Florida to digitize and display this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
48491453 ( OCLC )
2001240973 ( LCCN )
1409-3871 ( ISSN )

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Full Text
BIAI ESTERI AN



























I, i. issue I Lankesteriana is dedicated to the memory
I the great Mexican botanist and orchidologist,
1. 1 i LL, ANGEL SOTo ARENAS
I *" 32009),
I i.... II murdered in Mexico
on i .... i 9, 20(--







ISSN 1409-3871



LANESTERIANA

VOL. 9, No. 3 JANUARY 2010

Miguel Angel Soto Arenas (1963-2009)
EDUARDO A. PEREZ GARCIA & ERIC HAGSATER 269
In memorial: Miguel Angel Soto Arenas (1963-2009)
ERIC HAGSATER 273
Miguel Angel Soto Arenas: publications and main conferences (1983-2009) 276
A new species of Vanilla from South America
MIGUEL A. SOTO ARENASt 281
A revision of the mexican and Central American species of Vanilla
Plum. ex Mill. with a characterization of their ITS region of the nuclear
ribosomal DNA
MIGUEL A. SOTO ARENASt & ROBERT L. DRESSLER 285
A new infrageneric classification and synopsis of the genus Vanilla Plum. ex
Mill. (Orchidaceae, Vanillinae)
MIGUEL A. SOTO ARENASt & PHILLIP CRIBB 355
Comparettia sotoana (Orchidaceae: Oncidiinae), a new Ecuadorian species
FRANCO PUPULIN & GILBERTO MERINO 399
Confusion in Epidendrum brenesii Schltr., and a new Costa Rican species:
Epidendrum sotoanum (Orchidaceae: Laeliinae)
ADAM P. KARREMANS & ERIC HAGSATER 403
Oncidium ornithorhynchum, una especie mal interpretada y un nombre
para una vieja especie: Oncidium sotoanum (Orchidaceae)
ROLANDO JIMENEZ MACHORRO & ERIC HAGSATER 411
Two new species ofLepanthes from Costa Rica close to L. schizocardia
(Orchidaceae: Pleurothallidinae)
FRANCO PUPULIN, DIEGO BOGARIN & CHRISTINA M. SMITH 423
Stanhopeinae Mesoamericanae, V. Las Stanhopea de Mexico
GONTER GERLACH 431
_A continues


INTERNATIONAL JOURNAL ON ORCHIDOLOGY







LANKESTERIANA


Crossoglossa sotoana (Orchidaceae: Malaxideae), a new species honoring
the Mexican botanist, Miguel Angel Soto Arenas 443
FRANCO PUPULIN & ADAM P. KARREMANS

Dos species nuevas de Pleurothallidinae (Orchidaceae) para M6xico
RODOLFO SOLANO GOMEZ 447

Masdevallia sotoana (Orchidaceae: Pleurothallidinae), a new species
from Ecuador
HUGO MEDINA & FRANCO PUPULIN 455

New species of Porroglossum (Orchidaceae: Pleurothallidinae) from Ecuador
GILBERTO MERINO, ALFONSO DOUCETTE & FRANCO PUPULIN 459

On the identity of Myoxanthus scandens (Orchidaceae: Pleurothallidinae),
with a new species
FRANCO PUPULIN, DIEGO BOGARIN & MELANIA FERNANDEZ 467
Some new Sobralieae from Costa Rica and Panama
ROBERT L. DRESSLER & DIEGO BOGARIN 475

Lepanthes arenasiana (Pleurothallidinae: Orchidaceae), a new species from
Costa Rica
DIEGO BOGARIN & MELANIA FERNANDEZ 487

Sotoa, a new genus of Spiranthinae (Orchidaceae) from Mexico and southern
United States
GERARDO A. SALAZAR & CLAUDIA BALLESTEROS-BARRERA 491

A new species of Odontoglossum (Orchidaceae: Oncidiinae) from Ecuador
STIG DALSTROM & GILBERTO MERINO 505

A New Ornithidium (Orchidaceae: Maxillariinae) from the Massif
de la Hotte of Haiti
JAMES D. ACKERMAN & W. MARK WHITTEN 509

Novelties in the orchid flora of Venezuela II --- Cranichideae
GUSTAVO A. ROMERO-GONZALEZ, GERMAN CARNEVALI FERNANDEZ-CONCHA
& PAUL ORMEROD 513

A new species ofLophiaris Raf. (Orchidaceae) from the Pacific Coastal
of Mexico
RICARDO BALAM NARVAEZ, WILLIAM CETZAL IX
& GERMAN CARNEVALI FERNANDEZ-CONCHA 521

A la tercera se gana: the validation ofBenzingia (Orchidaceae:
Zygopetalinae)
GUSTAVO A. ROMERO-GONZALEZ & CALAWAY H. DODSON 526


IANKESTERIANA 9(3), January 2010. 0 Umversidad de Costa Rica, 2010.







INDEX


Epidendrumjalcanse (Orchidaceae), a new species from Northern Peru
MIGUEL CHOCCE, STIG DALSTROM, ERIC HAGSATER & JORGE ARNAIZ 529

AnAlisis de la selecci6n de sustrato por parte de Dendrophylax lindenii
(Lindl.) Benth. & Rolfe (Orchidaceae) en Cabo San Antonio, Peninsula de
Guanahacabibes, Pinar del Rio, Cuba
ERNESTO MUJICA, JOSEP RAVENTOS & ELAINE GONZALEZ 533

Orquideofl6rula de un sector de Serrania de La Cuchilla, municipio Caripe,
estado Monagas, Venezuela
CARLOS LEOPARDI 541

El redescubrimiento de Mexipedium xerophyticum (Soto Arenas, Salazar &
Higsater) V.A. Albert & M.W. Chase
EDUARDO A. PEREZ-GARCIA 557
Book Review 564
The Slide collection of Karlheinz Senghas now in "Word Orchid Iconography"
SAMUEL SPRUNGER 566

La colecci6n de dibujos de Erich Nelson en "Word Orchid Iconography"
SAMUEL SPRUNGER 567
Reviewers of the manuscripts submitted to Lankesteriana, vol. 8-9 568


IANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.











LANKESTERIANA
INTERNATIONAL JOURNAL ON ORCHIDOLOGY







Copyright C 2010 Lankester Botanical Garden, University of Costa Rica
Effective publication date: January 21, 2010

Layout: Jardin Botanico Lankester.
Cover: Vanilla helleri A.D.Hawkes. Drawing by M. Lopez and M.A. Soto Arenas.
Printer: Litografia Ediciones Sanabria S.A.
Printed copies: 500

Printed in Costa Rica / Impreso en Costa Rica


R Lankesteriana/ International Journal on Orchidology
No. 1 (2001)-- -- San Jose, Costa Rica: Editorial
Universidad de Costa Rica, 2001--
V.
ISSN-1409-3871

1. Botanica Publicaciones peri6dicas, 2. Publicaciones
peri6dicas costarricenses



0








LANKESTERIANA9(3) 269-272 2010


MIGUEL ANGEL SOTO ARENAS

(JULY 12, 1963 AUGUST 27, 2009) '

EDUARDO A. PEREZ GARCIA2 & ERIC HAGSATER3

2 Departamento de Ecologia y Recursos Naturales, Facultad de Ciencias,
Universidad NacionalAut6noma de Mexico, Ciudad Universitaria, Coyoacan, Mexico D.F.,
eduardo.perez-garcia@ciencias.unam.mx
3 AMO Herbarium, Instituto Chinoin, Mexico City, herbamon@prodigy.net.mx


Miguel Angel Soto was bor in the city of Torreon,
Coahuila, on July 12, 1963. In that city he studied
elementary and secondary school at the Colegio
Cervantes (1969-1978) whilst he finished high-school
at the Universidad Autonoma del Noreste (1978-
1980). Parallel with his first studies, Miguel Angel
became involved in the world of plants. His parents
loved plants and he inherited great part of this culture.
Living in a semi-arid region allowed him to get to
know closely one of his favorite groups of plants,
about which he acquired ample knowledge: the cati
and succulents. In spite of living in the desert, Miguel
Angel already knew and cultivated his favorite plants,
the orchids. Numerous visitors came to his house,
even journalists from the local communication media,
to see his orchid house.
Later on, Miguel Angel moved to Mexico City
to follow university studies; he majored in Biology
at the Faculty of Sciences /UNAM) from 1982-1987.
When he finished the credits, he began to write his
undergraduate thesis on the orchids of Bonampak,
Chiapas. It is suitable to point out that in those times
Bonampak was a very remote place, of difficult
access and with great extensions of the poorly known
Lacandon Jungle. From there, Miguel Angel and Jorge
Meave had to flee in view of the apocalyptic scenes
generated by the eruption of the "El Chichon" (or
Chichonal) volcano. Miguel Angel was a perfectionist
and, maybe because of this, he did not finish that
thesis, although he did publish an article on that results
of that sojourn in the heart of the Lacandon Jungle:
Soto Arenas, M.A., (1986), Orquideas de Bonampak,
Chiapas. Orquidea (Mex.) 10(1): 113-132. Miguel
graduated some-time later with a new thesis topic, this


1 Translation by Carlos Ossenbach.


time on the genus Lepanthes. He wrote the thesis with
Gerardo Salazar, and their work served as the basis
for the publication of the book: The genus Lepanthes
Sw. in Mexico, where they published 32 new species,
doubling the number of known species for Mexico.
Between finishing his undergraduate studies and
finishing his thesis, Miguel Angel developed numerous
projects and publications on orchids. During this
period, Miguel spent his days between the Ecology
Laboratory (of the Faculty of Sciences, UNAM) and
the AMO Herbarium since its inauguration (at that
time it was located in La Herradura, in the outskirts
of Mexico City, at Eric Hagsater's house), and his
innumerable field trips. He also traveled to the main
herbaria of the United States and Europe, studying all
the Mexican material on orchids; these voyages being
very fruitful in the search of types of numerous species
native to Mexico.
Miguel taught some undergraduate courses of the
biology, in the Faculty of Sciences, UNAM. Among
them: Bio-geography on 10 times (from 1983-1993),
General Ecology twice (1987-1988), Systematics
(1999), and Natural Resources (2000-2001). He was
also professor of the field biology: "Phytogeography of
the Montane Cloud Forest of the Sierra Madre in the
South of Guerrero" (1983-1984) and "Sinecological
Analysis of the Montane Cloud Forest of the State Park
of Omiltemi, Guerrero." (1984-1985).
He directed or co-directed around ten students both
of undergraduate as well as graduate levels. Some of
them received recognition for the quality of their
thesis; like Rodolfo Solano Gomez whose thesis "The
Genus Stelis Sw. (Orchidaceae: Pleurothallidinae) in
Mexico" (E.N.E.P Iztacala, UNAM, written in 1993
in co-direction with E. Hagsater, AMO Herbarium)
earned an Honors Mention in the Undergraduate








LANKESTERIANA


Thesis Contest of the Botanical Society of Mexico.
Similarly, Mariana Hemandez Apolinar received
First Place in the Undergraduate Thesis Contest
of the Botanical Society of Mexico with the thesis
"Population Dyanmics of Laelia speciosa (HBK)
Schltr. (Orchidaceae)" (Faculty of Sciences, UNAM,
written in 1992 with the co-direction of Irene Pisanty).
In October of 1993 he organized very successfully
the 5th Latin American Meeting of Orchideology in
Xalapa, Veracruz, with the participation of the best
known specialists and scholars of orchids of tropical
America. In this event an important international
exhibition of orchids was also held and of which he was
President of the Organizing Committee (Exporquidea
Xalapa '93). Miguel Angel was Vice-President of the
Latin American Orchideology Commission (C.L.O.,
1991-1993) and President of the same in 1993.
Miguel Angel was admitted into the Post-
Graduate Program in Ecology of the Ecology
Institute in 1994. He approved all the credits
and passed the doctoral candidacy examination.
However, due to his perfectionism, he postponed
sitting for his graduation examination until he had
the publications he considered were necessary.
The thesis he developed was titled "Evolution in
Vanilla (Orchidaceae): Phylogeny, biogeography and
evolution of characters" and he prepared it under the
direction of Dr. Elena Alvarez-Buylla (Laboratory of
Molecular Genetics and Evolution, Ecology Institute,
UNAM). Miguel Angel's studies on vanillas were
not limited to the aspects mentioned in the title of
his thesis, since, for example, he also included the
study of diversity, genetic variation and the uses
historic of the vanillas. From this work interesting
results derived, such as the routes and dates of the
human dispersion of the cultivation of Vanilla
planifolia around the world. Unfortunately he did
not see the publication of the formal description of
several new species and varieties of this genus, like
Vanilla costarricensis (ined), V cribbiana (ined), V
dessleri (ined), V martinezii (ined), V sarapiquiensis
(ined), V pompona subsp. grandulitnor, (Lindl.)
Soto Arenas (ined), and V espondae (ined). which
are now published in this issue of Lankesteriana,
and the corresponding species are included in his
contribution to Flora Mesoamericana, which had
already been delivered to the editors at Missouri

LANKESTERIANA9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


Botanical Gardens.
From his studies on vanillas the following were
published or were left to be published:

* Soto Arenas, M.A. (2006). La vanilla: Retos y
perspectives de su cultivo. (Vanilla: Challenges
and Perspectives of its cultivation). Biodiversitas
66: 1-9.
* Soto Arenas, M.A. (in press) A new species of
Vanilla from South America. Lankesteriana.
* Soto Arenas, M.A., & R.L. Dressier. (in press)
A revision of the Mexican and Central American
species of Vanilla Plum. ex Mill: Conspectus of
morphological and molecular data. Lankesteriana.
* Soto Arenas, M.A., K.M. Cameron & E. R.
Alvarez-Buylla. (in preparation) Phylogenetic
analysis of Vanilla Plum. ex Mill. (Orchidaceae:
Vanillianae) from congruent morphological and
molecular data.
* Soto Arenas, M.A. & P Cribb. (in preparation).
Annotated checklist, identification guide, and a
proposal for a new infrageneric classification of
the genus Vanilla Plum. ex Miller (Orchidaceae,
Vanillinae).
* Soto Arenas, M.A. & E. R. Alvarez-Buylla.
(in preparation) Notes on the floral biology of
Mexican Vanilla (Orchidaceae) and the evolution
of pollination systems in the genus.
* Soto Arenas, M.A. & E. R. Alvarez-Buylla.
(in preparation) Bio-geographic history of the
Pantropical genus Vanilla and the history on the
Gondwanic tropical biota.
* Soto Arenas, M.A., J. Cibrian, A., E. R. Alvarez-
Buylla, P Delgado & D. Pifiero. (in preparation)
Intraspecific variation of Vanilla planifolia: what
morphology, isozimes, RAPD's, and nuclear DNA
sequences indicate.

Miguel Angel was one of the most knowledgeable
people on the orchid flora of Mexico and, in general,
of all of Tropical America. He described, alone or
as co-author, many new orchids, among many others
are: Phragmipedium xerophyticum, Barkeria fritz-
halbingerii, Rossioglossum hagsaterianum, Sobralia
macdougalii, Stanhopea dodsoniana, S. whittenii,
Stelis greenwoodii, Elleanthus teotepecensis, Encyclia
calderoniae, E. rzedowskiana and Oncidium leleui.
Also, he formalized the intra-specific systematization








PEREZ-GARCIA & HAGSATER -Miguel Angel Soto Arenas (1963-2009)


of Laelia anceps, and reclassified (alone or as co-
author) numerous species of different genera, among
them Barkeria, Elleanthus, Rhynchostele, Prosthechea,
Dichromanthus, etc. He had ample knowledge on the
Pleurothallidiinae (Stelis, Acianthera, Pleurothallis,
etc.). According to the Missouri Botanical Garden
database (W3TROPICOS), there are more than 160
species and sub-species described by him, including
new descriptions and reclassifications of some
previously published taxa.
Miguel collected more than 11,000 different
samples (collection numbers) of plants in Mexico,
Guatemala, Costa Rica, Panama and Brazil, which
include almost 150 type collections. The main set of
his collections as well as his collecting notebooks and
personal notes are deposited in the AMO Herbarium
(Chinoin Institute, Mexico City). Miguel was a tireless
traveler and few people knew the natural habitats of
orchids like he did. This is one of the reasons why
he was one of the authors that wrote most in the book
"The Orchids of Mexico", and personally supervised
its design and edition; the work describes a journey
through the ecosystems of Mexico and its orchids,
including cultural and conservation chapters. Together
with the digital Catalogue (CD) of The Orchids of
Mexico it is the most complete popular work there is
about Mexican orchids.
One of his already classic works was the
publication: Updated Listing of the Orchids of Mexico
[Orquidea (Mexico City.) 11: 231-275 (1989)]; this
list was the basis of the most recent lists about the
orchids of Mexico. Jointly with Federico Halbinger
he coauthored the book Laelias of Mexico, which is
one of the most widely cultivated genera by those fond
of orchids. Miguel Angel was executive editor of the
journal Orquidea (Mexico City.) from 1985 to date.
He was also executive editor of some volumes of the
Icones Orchidacearum (Mexico), which is probably
the best technical reference of the Mexican orchids.
He had two other volumes in preparation. Other
issues deal with the genus Epidendrum throughout
the neotropics.
Also, Miguel Angel collaborated with several of
the great current orchideologists including Robert
Dressier, Gerardo Salazar, Eric Hagsater, German
Camevali, Mark Chase, Cassio van den Berg, Mark
Whitten, Phil Cribb, and Ed Greenwood, among


many others. Miguel's publications are a mandatory
reference for orchid scholars, but are also very useful
for bio-geographers and evolutionists of the flora of
the Neotropics. Due to his great work as a botanist,
several species have been dedicated in his honor like:
Lepanthes sotoi Archila, Maxillaria sotoana Camevali
et G6mez-Juirez, Mormodes sotoana Salazar, Stelis
sotoana R. Solano, and some others that are being
published in this issue of Lankesteriana.
Miguel was a conservationist since an early age; for
example, he was one of the most participative students
in the creation of the Reserve of the Pedregal of San
Angel in Mexico City. More recently, he published
one of the most complete works on the current situation
of orchid conservation in Mexico and participated in
many forums related with the conservation of orchids.
He was a prominent member of the "Orchid Specialist
Group, Species Survival Commission, IUCN" (1993-
1997; 1998-2009) and a member of the "Conservation
ex-situ Committee" of the same commission
(2000-2003). He also participated as counselor of
various government agencies like SARH (Ministry
of Agriculture and Water Resources), SEDESOL
(Ministry of Social Development), SEMARNAT
(Ministry of Natural Resources), CONABIO (National
Commission of Biodiversity), establishing the most
important criteria for the national orchid conservation
strategies of Mexico.
Miguel Angel was a botanist who loved plants and
this was reflected in his very good hand in cultivating
various families, among which stand out the
Crassulaceae, the Cycadaceae and of course, orchids.
He was able to form the most important collection
of live plants of Mexican orchid species, much of
which is now located in the live collection at AMO
Herbarium. From the cultivated plants and the field
samples, he helped form a DNA bank for research in
molecular biology of almost 500 orchid species, and in
addition 500 Vanilla samples.
In collaboration with E. Hagsater and Cassio van
den Berg, he was preparing a phylogeny of the genus
Epidendrum based both on molecular data as well
as morphological and on the vegetative architecture;
for that he had sequenced more than 300 species
from throughout the neotropics, mostly collected by
Hagsater and cultivated at AMO, always careful to
have voucher specimens prepared.
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


He stood out for his attitude, always cooperating
with hobbyists and other biologists, which led him
to share his knowledge in his field trips, in numerous
sessions of the Mexican Orchid Association, and
with farmers, particularly those that cultivate vanilla
in the states of Veracruz, Oaxaca and Chiapas. He
was a proficient lecturer at conferences and scientific
seminars, he delivered more than 150 conferences in
universities, in botanical gardens, and in botanical and
horticultural associations.
Those of us who had the good fortune of having
access to his field notebooks, could observe the
encyclopedic knowledge he had about the flora and
vegetation of Mexico. His acute vision allowed him
to find orchids even when driving the car at high speed
or in the darkness of the closed forest canopy. It is
suitable to point out that he performed as a human
altimeter, since he could calculate very exactly the
altitude where we found ourselves by only looking at
the vegetation around us.
Nora Esponda, the associate secretary of Instituto
Chinoin, worked with Miguel for 25 years and
developed a close friendship/professional relationship,
and describes him as passionate in his work and
everything he did, his conversations, his friendship, he
was also fun, and worked late into night, sending mails
and instructions in the early morning. She claims he


described himself as "neurotic and ill-tempered", but
she never saw him loose control, his character was
strong but affable, very demanding of himself, but also
very sensitive.
His sister Miriam describes him as a great
Chef, it was Miguel who invited the family over
for Christmas or some special occasion. He enjoyed
cooking a chicken in "mole", or a leg of venison with
herbs in white wine for his friends. His specialty,
vanilla ice-cream, was delicious. He moved back to
Torreon a couple of years ago in search of tranquility
to continue writing about orchids of Mexico, and
spend much more time with his parents and two
sisters. He had recently traveled with his sisters on a
month-long trip through the Huasteca region of San
Luis Potosi, Veracruz and into Chiapas; they were
amazed how well he knew the country and the most
spectacular landscapes. A couple of weeks before his
assassination, he had discovered what appeared to be
a new species of Pine-tree, on an excursion with his
sister Miriam and other friends.
His tragic death occurred on August 27, at his
home in Torreon, Coahuila. Miguel Angel Soto was
a controversial character, but without a doubt, he was
an extraordinary human being who leaves behind a
great void that will be very difficult to fill. We all miss
him.


LANKESTERIANA9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








LANKESTERIANA9(3) 273-275 2010


INMEMORIAM: MIGUEL ANGEL SOTO ARENAS

(1963-2009)

ERIC HAGSATER

Herbario AMO, Montafias Calizas 490, Mexico D.F. 11000, Mexico


We probably first met at one of the monthly
gatherings of the Asociaci6n Mexicana de
Orquideologia in the early 1980's. He and Gerardo
A. Salazar were studying biology at the Faculty of
Sciences at the UNAM, the National University in
Mexico City. Both made their social service mounting
specimens and looking after the live orchid collection
at the recently founded Herbario AMO (then a part
of the Asociaci6n Mexicana de Orquideologia). They
both proved to have a deep interest in orchids, and
decided to pursue a career in orchid research.
Miguel chose to specialize in ecology and
taxonomy, and one of his first projects was the
epiphytic flora of the region of the archaeological site
of Bonampak, Chiapas, were he became particularly
well acquainted with the orchids of this tropical,
lowland rain-forest. He later participated in several
studies of the mesophilous forests of Omiltemi,
Guerrero, another site of high orchid diversity, but
this time in the higher mountains of southern Mexico,
overlooking the Pacific Ocean.
In was in 1985 that, after finishing their formal
undergraduate studies, that they both became formally
employed as a research associates in Mexican orchids
by the Instituto Chinoin, A.C., in particular in the
AMO Herbarium as a base. The objective was to
further the knowledge of Mexican orchids from every
viewpoint.
Through the years Miguel was able to travel to
the most important herbaria in the Americas and
Europe, spending sometimes weeks or months at a
time, studying and photographing Mexican orchid
material. The information contained in the images
was transferred to the electronic database at AMO,
now covering some 130 thousand records. Most of
the slides have been digitalized for easier access.
Miguel thus obtained an encyclopedic knowledge
of the Mexican orchids. One of his goals was to
finish Orchids of Mexico, a treatise covering all


the taxonomic and ecological information of all
the known Mexican orchid species. This work was
done in collaboration with Hagsater, who focused
on the genus Epidendrum, Gerardo A. Salazar, on
terrestrial orchids, and Rodolfo Solano in relation to
the Pleurothallidinae; Rolando Jimenez worked on
Oncidium. Unfortunately, this work, now covering
some 1,900 pages remains unfinished.
From the early beginning he stood out as someone
willing to learn from others, doing team work and
sharing his information with others. This can be seen
through the number and variety of thesis he directed,
not only with students in Mexican universities, but
also at Oxford, England and Riverside, California, as
well as in the many papers he co-authored.
Miguel tackled many groups of mainly epiphytic
genera, and had a special interest in the Mexican
Laelia, where he worked together with Federico
Halbinger and produced a magnificent revision of all
the species and lower taxa, having personally visited
with Federico practically each and every locality, to
understand each entity.
During several years he worked with Mariana
Hernandez in the state of Michoacan studying the
population dynamics ofLaelia speciosa, studying how
the local farmers cropped the plants for the flower
market in ways that could be sustainable. They also
followed seedlings on oak trees for several years,
recording how many survived each successive year,
and how many years it took adult plants to flower for
the first time. At the time there were a number of such
studies with terrestrial orchid species in Europe and
the United States, but very few with epiphytic species
in tropical countries. Mariana's thesis was recognized
as the best thesis of the year by the Mexican Botanical
Society.
Once, while studying the material of
Phragmipedium exstaminodium he exclaimed "you
collected this specimen the day I was born!" Indeed








LANKESTERTANA


that June day in 1963 I was visiting the lakes of
Monte Bello, a two day jeep-ride from Comitan,
through cattle ranches and virgin forests in the
highlands of Chiapas. The clear-water, colored-
lakes where surrounded by Pine-Oak forests covered
with epiphytes, including many orchids, and the
amazing lady-slipper with two-foot long petals in
full bloom! Miguel and I were deeply aware of the
destruction of many natural habitats, mainly due
to their transformation into farm-land. That forest
however was mostly destroyed by the fires of the
very hot spring of 1998, after a deep-cold winter the
previous year had killed many of the epiphytes in
the trees. He made several field trips to Chiapas to
evaluate the state of conservation of Phragmipedium
exstaminodium for the IUCN. A couple score mature
plants were found, but no young seedlings, there
appeared to be no natural reproduction.
Miguel, with his interest in ecology, participated
in numerous conservation conferences, was a
member of the IUCN Orchid Specialist Group, and
collaborated with numerous governmental agencies
regarding orchid conservation. He prepared the
revision of the list of endangered orchids for the
Mexican government, prepared monographs of each
species, the illustrations prepared by several other
botanists at AMO.
Curiously, we never went into the field together
during these 25 years of working together. We
often discussed plans for the future and worked on
many projects, one of them being a new system of
infrageneric classification in Epidendrum, combining
floral, vegetative and molecular data. He sequenced
over 300 species of the genus throughout most of its
range, most of the samples from plants which I had
collected during field trips to Costa Rica, Panama,
Colombia and Ecuador. We came to have a good
understanding of the variation. His sudden departure
leaves this project in chaos.
One of our dreams, since we began working
together, was to eventually publish an illustrated book
on the Orchids of Mexico. We agreed that it should
not be a catalogue, nor a scientific reference book, but
rather a narration of the many diverse habitats where
orchids can be found, a walk through these forests and
savannas. Through the years we gathered material,
and he became intimately acquainted with each and

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


every habitat. The problem was how to finance the
edition. I discussed the issue with several prominent
editors without finding any way out. Suddenly, in
late 2004, at the pharmaceutical company of which
Institute Chinoin is a part, I found myself discussing
how we could commemorate the upcoming 80'h
anniversary of the introduction of its products to the
Mexican market. All things came together; we could
publish a book of interest to a cultivated public,
which was the result of the accumulated knowledge
financed by the Institute. Thus we rushed to put a
long cherished dream into action. Miguel was at
first in charge of putting everything in writing with
the help of Gerardo Salazar. The photographs were
to be made mainly by Rolando Jimenez Machorro
and Marco Lopez Rosas, the illustrators and
photographers at AMO. We contracted one of the
top scientific publishers in Mexico for the publishing
work and asked them to find a suitable translator. As
it was evident that we could not include photographs
of all the species known to Mexico, we decided to
add a CD with as many as we could get. We asked
for and got collaboration from all quarters of the
world for suitable slides, three dozen photographers
participated. Miguel dropped everything else he was
working on, including his doctoral thesis on Vanilla.
He got involved in everything, including the balance
and quality of the color selections, and the lay-out of
the entire book. We could not find a suitable translator
into English, so he made a first translation which
Gerardo Salazar and I then worked on with a final
revision by Robert Dressier. The book was printed in
Japan and arrived in time for the anniversary of the
pharmaceutical company. Miguel had put a full year's
work into it, often working until mid-night.
It was an immediate success, within a year the
25,000 copies printed in Spanish were distributed and the
edition was sold out; those who received the book did
not let go of it. It had, however, a surprising unintended
effect; it became the standard show-book for sellers of
wild-collected orchid species in the local markets in
Mexico City and elsewhere. We were dismayed by
this. After years of efforts by Miguel Angel to curb this
illegal market which depleted populations of mostly
desirable horticultural species, growing and collecting
species orchids became popular again, with very few
sources of propagated plants.








HAGSATER In memorial: Miguel Angel Soto Arenas


Probably one of Miguel's most cherished projects
was Vanilla. Aside from being the subject of his
doctoral thesis, for which he sequenced some 500
samples from around the world, he discovered that
the plant in commerce for its flavor was basically a
single clone probably selected by the Totonaca people
of the State of Veracruz. He had special interest in
working with local communities in recuperating this
valuable crop. He searched for the few individual
plants with diverse genetic structures. Working in
collaboration with several specialists throughout
the world he prepared a proposal for a new generic
classification, and described a number of species
new to science. Seven papers were prepared for
publication, unfortunately only one was sent to the
publisher. Others pending details are being finished


by the coauthors, and will hopefully be published
sometime soon, some in this issue dedicated to
Miguel Angel.
Miguel was working at his home in Torreon,
Coahuila, late at night on August 27th. His sister had
been helping him till eight o'clock that evening.
Suddenly, an assailant entered his house, and after
a fierce battle assassinated him. The reasons are still
unknown. His body was discovered the following
day. He was 47 years old.
His sisters and parents, the whole orchid
community, is shocked, as we have all lost a great
friend and collaborator, and one of the greatest
contributors to the knowledge of orchids in Mexico,
not only their diversity, but their ecology and
conservation. May he rest in peace.


LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA 9(3) 276-2802010


MIGUEL ANGEL SOTO ARENAS:

PUBLICATIONS AND MAIN CONFERENCES, 1983-2009


PEER REVIEWED PUBLICATIONS (PAPERS AND CHAPTERS IN BOOKS)

Lorenzo S.A., L., A. Ramirez, M1A. Soto Arenas, A. Breceda, M. del C. Calderon, H. Cortez, C. Puchet, M.
Ramirez, R. Villal6n & E. Zapata. 1983. Notas sobre la fitogeografia del bosque mes6filo de montafia en la
Sierra Madre del Sur, Mexico. Bol. Soc. Bot. MIx. 44: 97-102.
Soto Arenas, M.A. 1982. El Pedregal de San Angel: un refugio natural de orquideas. Ocelote 1(1): 9-11.
Soto Arenas, M.A. 1986. Orquideas de Bonampak, Chiapas. Orquidea (Mex.) 10(1): 113-132.
Soto Arenas, M.A. 1986. El g6nero Elleanthus en Mexico y una nueva especie de Guerrero, Elleanthus
teotepecensis Orquidea (Mex.) 10(1): 161-190. 1986.
Soto Arenas, M.A. 1987. Una revision de las species mexicanas de Trichosalpinx subgen Trichosalpinx.
Orquidea (Mex.) 10(2): 247-296.
Hagsater, E. & MVA. Soto Arenas. 1998. Orchid conservation in Mexico. Selbyana 19(1): 15-19.
Salazar, G.A. & M.A. Soto Arenas. 1989. Lepanthes hagsateri, una nueva especie de Guerrero, Mexico.
Orquidea (Mex.) 11: 15-22. 1989.
Soto Arenas, M.A. & E.W. Greenwood. 1989. Undesirable technical terminology --a current example. Orch.
Research Newsletter 13: 8-9.
Soto Arenas, M.A. & E. Hagsater. 1990. Algunas ideas acerca de la conservaci6n de orquideas mexicanas. Pp.
155-172 in J.L. Camarillo y F. Rivera (eds.) Areas Naturales Protegidas y Especies en Extincidn. UNAM.
Mexico, D.F.
Soto Arenas, M1A., G.A. Salazar & E. Hagsater. 1990. Phragmipedium xerophyticum, una nueva especie del
sureste de Mexico. Orquidea (Mex.) 12(1): 1-10. (German translation: Neue Frauenschuhorchideen aus
Mexiko, Phragmipedium xerophyticum Soto, Salazar & Hagsater. Die Orchidee 42(5): 253-262. 1991).
Salazar, G. & M1A. Soto Arenas. 1990. Una nueva especie de Malaxis (Orchidaceae) de flores grandes del norte
de Chiapas. Acta Botinica Mexicana 10: 45-49.
Soto Arenas, M.A. 1990. Una nueva orquidea de Morelos, Mexico: Ponera dressleriana. Orquidea (Mex.) 12:
117-126.
Jimenez, R. & M.A. Soto Arenas. 1990. Oncidium leleui, una nueva especie de la costa pacifica mexicana.
Orquidea (MAx.) 12(1): 57-64. (French translation: Oncidium leleui, une espece nouvelle de la c6te pacifique
du Mexique. Orchidees. Culture et Protection 10:3-9. 1992).
Soto Arenas, M.A. 1991. Studien fber die Population und die Moeglichkeit der Erhaltung des Phragmipedium
exstaminodium (Orchidaceae), ein gefehrdeter Arten. pp 132-134. In E. Hagsater (ed.) Phragmipedium
exstaminodium Castafio, Hagsater & Aguirre. Erst vor wenigen Jahren entdeckt -heute fast von Aussterben
bedroth. Die Orchidee 42(3): 127-134.
Salazar, G. & M.A. Soto Arenas. 1992. Lepanthes williamsii (Orchidaceae), una nueva especie de Mexico y
Guatemala. Orquidea (Mex.) 12(2): 139-143.
Jimenez, R. & M.A. Soto Arenas. 1992. El complejo Oncidium maculatum. Orquidea (Mex.) 12(2): 297-316.
Meave, J., M1A. Soto Arenas, L. Calvo, H. Paz y S. Valencia. 1992. Analisis sinecol6gico del bosque mes6filo
de montafia de Omiltemi, Guerrero, Mexico. Bol. Soc. Bot, MIxico 52: 31-77.
Soto Arenas, M1A. 1992. Una nueva especie de Chiapas: Ornithocephalus obergiae. Orquidea (Mex.) 12(2):
193-198.
Soto Arenas, M.A.1992. Maxillaria histrionica y M. tonsoniae. Orquidea (Mex.) 12(2): 244-250.
Soto Arenas, MVA. 1992. Maxillaria hagsateriana, una nueva especie similar a Maxillaria densa. Orquidea
(Mix.) 12(2): 251-260.


IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








Miguel Angel Soto Arenas: publications and conferences (1983-2009)


Soto Arenas, M1A. 1992. Draculla pusilla an addition to the Mexican orchid flora. Orquidea (Mfx.) 12(2): 277-
280.
Soto Arenas, MVA. & F. Chiang. 1992. Maxillaria lexarzana, un nuevo nombre para Psitacoglossum atratum
Orquidea (MAx.) 12(2): 237-243.
Soto Arenas, M.A. (1993). Population studies in Mexican orchids. pp. 153-160 in A.M. Pridgeon. [ed.]
Proceedings of the 14th World Orchid Conference, Glasgow. HMSO Publ., Edinburgo.
Soto Arenas, M.A. (1994). Orchidaceae pp. 170-176 en Martinez, E., C.H. Ramos y F. Chiang. Lista Floristica
de la Lacandona, Chiapas. Bol. Soc. Bot. MIxico 54: 99-177.
Soto Arenas, M.A. (1994). Barkeria melanocaulon y Barkeria whartoniana. Orquidea (MAx.) 13(1-2): 233-244.
Soto Arenas, M.A. (1994). Barkeriafritz-halbingeriana. Orquidea (MAx.) 13(1-2): 245-248.
Soto Arenas, M.A. (1994). Eurystyles, a new generic record for the Mexican orchid flora. Orquidea (MAx.)
13(1-2): 269-274.
Soto Arenas, M.A. (1994). Vanilla odorata. Orquidea (MAx.) 13(1-2): 290-294.
Soto Arenas, M.A., G.A. Salazar y A. Rojas. (1994). Nomenclatural changes inRhynchostele and Lemboglossum
(Orchidaceae, Oncidiinae). Orquidea (MAx.). 13(1-2): 145-152.
Jimenez, R. y M Soto Arenas. (1994). Oncidium hagsaterianum, una nueva especie de Mexico y Guatemala.
Orquidea (Mfx.) 13(1-2): 159-164.
Cribb, P y M.A. Soto Arenas. (1994). The genus Cypripedium in Mexico and Central America. Orquidea (Mfx.)
13(1-2): 125-144. (Cribb, P und M. Soto Arenas. 1994. Die Gattung Cypripedium in Mexiko und Zentral-
America. Die Orchidee: 1995).
Solano, R. y MVA. Soto Arenas. (1994). Stelis aeolica y S. vespertina en R. Solano, El g6nero Stelis en Mexico.
Orquidea (Mfx.) 13(1-2): 18-21; 106-108.
Salazar, G. y MVA. Soto Arenas. (1996). A new species of Ornithocephalus (Orchidaceae) from Oaxaca, Mexico.
Brittonia 48(2): 209-212.
Soto Arenas, M.A. (1996). The importance of research. pp. 33-38. en IUCN/SSC Orchid Specialist Group.
Orchids Status Survey and Conservation Action Plan. IUCN, Gland, Suiza.
Soto Arenas, MVA. (1996). Mexico (Regional Account). pp. 53-58 en IUCN/SSC Orchid Specialist Group.
Orchids Status Survey and Conservation Action Plan. IUCN, Gland, Suiza.
Soto Arenas, M.A. (1997). Introduction aux orchid6es mexicaines. Orchidees. Culture et Protection. 31: 18-21.
Hagsater, E. y M.A. Soto Arenas. (en prensa). Notes for a proposal of a infrageneric phylogenetic classification
of Epidendrum, the largest Neotropical orchid genus. Proceedings of the 18th World Orchid Conference,
Miami, Florida. Estados Unidos.
van der Berg, C., W.E. Higgins, R.L. Dressier, W.M. Whitten, MVA. Soto Arenas, A. Culham, y M.W. Chase.
(2000). Aphylogenetic analysis of Laeliinae (Orchidaceae) based on sequence data from Internal Transcribed
Spacers (ITS) of nuclear ribosomal DNA. Lindleyana 15(2): 96-114.
Salazar, G.A. y M.A. Soto Arenas. (2001). Anew species of Stanhopea (Orchidaceae) from Mexico. Lindleyana
16(3): 144-148.
Salazar, G.A. y MVA. Soto Arenas. (2001). Nomenclatural changes in Mexican Orchidaceae. Lindleyana 16(3):
149-150.
Salazar, G.A., M.W. Chase y M.A. Soto Arenas. (2002). Galeottiellinae, a new subtribe and other nomenclatural
changes in Spiranthinae (Orchidaceae: Cranichideae). Lindleyana 17(3): 172-176.
Salazar, G.A., M.W. Chase, M.A. Soto Arenas y M. Ingrouille. (2003). Phylogenetics of Cranichideae with
emphasis on Spiranthinae (Orchidaceae, Orchidoideae): Evidence from plastid and nuclear DNA sequences.
Am. J. Bot. 90(5): 777-795.
Light, M., J. Reddoch, E. Hagsater y M Soto Arenas. (2003). Edward W. Greenwood. In memorial. Bol. Soc.
Bot. MAx. 71: 85-84.
Soto Arenas, MVA. 2003. Vanilla. Pp. 321-334. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase y F.N. Rasmussen.


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(eds.). Genera Orchidacearum vol. 3. Orchidoideae (Part two), Vanilloideae. Oxford University Press.
Oxford.
Soto Arenas, M.A. & R.L. Dressier. 2004. Vanilla. Pp. 383-387. In B.E. Hammel, M.H. Grayum, C. Herrera y
N. Zamora (eds.) Manual de Plantas de Costa Rica, vol. 3 Monocotiled6nea (Orchidaceae-Zingiberaceae).
Missouri Botanical Garden,-INBio-Museo Nacional de Costa Rica.
Soto Arenas, M.A. & J.D. Ackerman. 2005. Domingoa. Pp. 228-232. figs. 299.1, 2. In: Pridgeon, A.M., PJ.
Cribb, M.W. Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one).
Oxford University Press. Oxford.
Soto Arenas, M.A. & G.A. Salazar Chavez. 2004. Orquideas. Pp. 271-295 in A.J. Garcia-Mendoza y M.J.
Ordofiez y M. Briones-Salas (eds.) Biodiversidad de Oaxaca. Institute de Biologia, UNAM-Fondo
Oaxaquefio para la Conservaci6n de la Naturaleza-World Wildlife Fund, Mexico D.F.
Hagsater, E. & M.A. Soto Arenas. 2005. Epidendrum. Pp. 236-251, figs. 301.1, 2, 3. In: Pridgeon, A.M., PJ.
Cribb, M.W. Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one).
Oxford University Press. Oxford.
Hagsater, E. & M.A. Soto Arenas. 2005. Transfers to Epidendrum from Oerstedella Rchb.f. Lankesteriana 5(1):
73-75.
Soto Arenas, M.A. 2005. Alamania. Pp. 190-193, figs. 287.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Arpophyllum. Pp. 193-196, figs. 288.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase
& F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University
Press. Oxford.
Soto Arenas, M.A. 2005. Artorima. Pp. 196-199, figs. 289.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Barkeria. Pp. 199-205, figs. 290.1, 2, 3. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Subtribe Chysinae. Chysis. Pp. 173-177, figs. 283.1, 2. In: Pridgeon, A.M., P.J. Cribb,
M.W. Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford
University Press. Oxford.
Soto Arenas, M.A. 2005. Hagsatera. Pp. 254-257, figs. 303.1,2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Helleriella. Pp. 424-426, figs. 362.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, 1VA. 2005. Homalopetalum. Pp. 257-260, figs. 304.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W.
Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford
University Press. Oxford.
Soto Arenas, 1VA. 2005. Laelia. Pp. 265-271, figs. 307.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase & F.N.
Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Meiracyllium. Pp. 276-279, figs. 310.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase
& F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University
Press. Oxford.
Soto Arenas, MA. 2005. Microepidendrum. Pp. 279-282, figs. 311.1, 311.2. In: Pridgeon, A.M., PJ. Cribb,


LANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








Miguel Angel Soto Arenas: publications and conferences (1983-2009)


M.W. Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford
University Press. Oxford.
Soto Arenas, M.A. 2005. Nemaconia. Pp. 429-432, figs. 364.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase&
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Ponera. Pp. 432-434, figs. 365.1, 2. In: Pridgeon, A.M., PJ. Cribb, M.W. Chase &
F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Subtribe Ponerinae. Pp. 422-424. In: Pridgeon, A.M., PJ. Cribb, M.W Chase & F.N.
Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press.
Oxford.
Soto Arenas, M.A. 2005. Arpophyllum (pp. 22-25), Barkeria (pp. 46-51; en colaboraci6n colaboraci6n con E.
Hagsater), Isochilus (pp. 382-385) y Jacquiniella (386-389) In: Pupulin, F. (ed.) y colaboradores. Vanishing
Beauty: Native Costa Rican Orchids. vol. 1. Editorial de la Universidad de Costa Rica. San Jose.
van den Berg, C. & M.A. Soto Arenas. 2005. Artificial key to genera of subtribe Laeliinae. Pp. 184-186.
In: Pridgeon, A.M., PJ. Cribb, M.W Chase & F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4.
Epidendroideae (Part one). Oxford University Press. Oxford.
Soto Arenas, M.A. 2006. La vanilla: Retos y perspectives de su cultivo. Biodiversitas 66: 1-9.
Soto Arenas, M.A., G.A. Salazar & C. van den Berg. 2007. New combinations in Domingoa, Homalopetalum
(Orchidaceae: Laeliinae) and Nemaconia (Orchidaceae: Ponerinae). Neodiversity 2: 7-9.
Schlfiter, PM., M.A. Soto Arenas & S.A. Harris. 2007. Genetic variation in Vanilla planifolia (Orchidaceae).
Economic Botany 61(4): 328-336.
Soto Arenas, M.A., R. Solano Gomez & E. Hagsater. 2007. Risk of extinction and patters of diversity loss in
Mexican orchids. Lankesteriana 6: 123-132.
van den Berg, C., WE. Higgins, R.L. Dressier, W Mark Whitten, M.A. Soto-Arenas & M.W Chase. 2009. A
phylogenetic study of Laeliinae (Orchidaceae) based on combined nuclear and plastid DNA sequences.
Annals of Botany, 104 (3): 417-430.

BOOKS

Soto Arenas, M.A. 1990. (30/100 texts of plates, second author) in G. Salazar & E. Hagsater (eds.) Orquideas de
Mexico, part 1. Icones Orchidacearum. Asociacion Mexicana de Orquideologia. Mexico D.F.
Salazar Chavez, G.A. & M.A. Soto Arenas. 1996. El g6nero Lepanthes en Mexico. Asociacion Mexicana de
Orquideologia, A.C. Mexico D.F. 231 pp.
Halbinger, F. & M.A. Soto Arenas. 1997. Laelias of Mexico. Herbario AMO. Mexico D.F. 160 pp.
Soto Arenas, M.A. 2003. (137/200 texts of plates, first author) In: E. Hagsater & M.A. Soto Arenas (eds.).
Orchids of Mexico, parts 2 and 3. Icones Orchidacearum fasc. 5-6. Herbario AMO. Mexico D.F.
Hagsater, E., M.A. Soto Arenas, G.A. Salazar Chavez, R. Jimenez Machorro, M.A. Lopez Rosas & R.L. Dressier.
2005. Las Orquideas de Mexico. Institute Chinoin, Mexico D.F. (simultaneous edition in English, Orchids
of Mexico).
Soto Arenas, M.A. 2008. (87/100 texts of plates, first autor) In: E. Hagsater & Soto Arenas, M.A. (eds.) Orchids
of Mexico, part 4. Icones Orchidacearum fasc. 10. Herbario AMO. Mexico D.F.

OTHER PUBLICATIONS

Preparation and edition of the "Timbre de la Tuberculosis". Comit6 Nacional de la Lucha contra la Tuberculosis
y Enfermedades del Aparato Respiratorio". (1997).
Soto Arenas, M.A., E. Hagsater, R. Jimenez Machorro, G.A. Salazar Chavez, R. Solano Gomez, R. Flores


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Gonzalez & I. Ruiz Contreras. 2007. Las orquideas de Mexico. Catalogo Digital. Interactive Multimedi CD,
Win-Mac. Herbario AMO. Institute Chinoin, A.C. Mexico.
Soto Arenas, M.A., R. Solano Gomez, (with the collaboration by R. Jimenez Machorro, L. Sanchez Saldafia &
G.A. Salazar). 2007. Fichas tecnicas de las species de orquideas incluidas en la NOM-059-ECOL-2007
(SEMARNAT).

SPECIAL INVITED AND KEYNOTE CONFERENCES

"Influencia del Dosel Forestal en la Vegetaci6n Epifita". Sociedad Botanica de Mexico, March 1988.
"La Flora y la Vegetaci6n de Mexico". Museo Nacional de Historia, Castillo de Chapultepec, Cd. de Mexico.
January 1993.
"Population Studies in Mexican Orchids". 14th World Orchid Conference, Glasgow, Scotland, April 1993.
"Orquideas Mexicanas en Peligro de Extinci6n". V Congreso Nacional de Horticultura Ornamental, 1995, FES
Cuautitlan, UNAM- Asociaci6n Mexicana de Horticultura Ornamental. Keynote Conference.
"Laelias, Barkerias and Rhynchosteles. Jewels of the Mexican Orchids. Iowa Orchid Society. Des Moines, IA.,
1997.
"Systematics of Vanilla (Orchidaceae)". Royal Botanic Gardens, Kew, Jodrell Laboratory, U.K. April 1999.
"Genetic Resources of Vanilla", 16th World Orchid Conference, Vancouver, May 2000,
"Orquideas Mesoamericanas: Evoluci6n, experiencias y perspectives de conservation". Chairman conference. II
Seminario Latinoamericano de Orquideologia, San Jose, Costa Rica (in collaboration with G. Salazar and
E. Hagsater), May 2001.
"La Conservaci6n de las Orquideas de Mexico". Simposium de Conservaci6n, XII Congreso Mexicano de
Botanica, Queretaro (in collaboration with G. Salazar and E. Hagsater), October 2001.
"CITES, conservation and orchid taxonomy in the Neotropics; personal notes and a proposal of criteria for non-
commercial collections" (in collaboration with E. Hagsater), 17th World Orchid Conference, Shah Alam,
Malaysia, April 2002.
"Phylogeny of Epidendrum" (in collaboration with E. Hagsater), 17th World Orchid Conference, Shah Alam,
Malaysia, April 2002.
"Evoluci6n y recursos gen6ticos en Vanilla (Orchidaceae)". Seminarios de Botanica, Instituto de Biologia,
UNAM. May 2002.
"Evoluci6n de Epidendrum" y "Biogeografia de las orquideas del Neotr6pico" (Keynote Conferences), VI
Taller Intemacional de Orquideas, Jardin Botanico Orquideario Soroa, Universidad de Pinar del Rio, Cuba.
November 2002.
"Evoluci6n en Vanilla (Orchidaceae)". Seminario, Colegio de Posgraduados, Montecillos, Edo. de Mexico.
"Las Orquideas de Oaxaca" Jardin Hist6rico Etnobotanico. Centro Cultural Santo Domingo, Oaxaca, Oax.
"Vanilla The challenges of a crop based on an endangered species with a complex life history". Vanilla 2005.
Third International Congress, Boca del Rio, Ver., November 2005.
"La Vainilla Los retos de un cultivo basado en una especie amenazada con una historic de vida compleja".
Congress intemacional de productores de Vainilla. Papantla, Ver. May 2006.
"Notes for a proposal of an infrageneric phylogenetic classification of Epidendrum, the largest Neotropical orchid
genus" (in collaboration with E. Hagsater), 18th World Orchid Conference, Miami, Florida. U.S.A. January
2008.


IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA9(3) 281-284 2010


A NEW SPECIES OF VANILLA FROM SOUTH AMERICA'


MIGUEL A. SOTO ARENASt


Herbario AMO, Montafias Calizas 490, Mexico D.F. 11000, Mexico



ABSTRACT. A new species from the basin of the Rio Magdalena in northern Colombia is proposed, Vanilla
espondae.

KEY WORDS: Orchidaceae, Vanilla, V espondae, Rio Magdalena, Colombia


During my work with the phylogeny of Vanilla
it became evident that some specimens represent
undescribed species in the genus. The species here
described is among the most showy in the genus,
suggesting that additional, undescribed vanillas may
be remain to be found*.
The taxon is native to South America, where
Vanilla reaches its highest diversity, and from where
much more material is needed. Its relationships are
discussed.

Vanilla espondae Soto Arenas, sp. nov.

HOLOTYPE: [N.] Colombia: Tributary of R.
Magdalena, cult. F Perez-Vera 563, K-L! (illustration
voucher).

A Vanillae trigonocarpae Hoehne disco labelli papillis
verrucisque ornato recedit.

Hemi-epiphytic herb. Stems apparently thin,
terete, smooth, olive green, ca. 6.3 mm thick, the
intemodes as long as the leaves. Leaf shortly petiolate,
the petiole ca. 8 mm long, blade oblanceolate, abruptly
acuminate, the apex incurved, base obtuse, the basal
margin revolute, coriaceous-fleshy, green, 14.5 x
3.6 cm. Inflorescence axillary, a shortly pedunculate

1 Editor's note: Miguel Angel Soto Arenas passed away
August 27h, 2009. We wish to thank Dr. Phillip Cribb
for preparing the camera lucida drawing of the lip of the
holotype, and Rolando Jimenez Machorro for preparing the
attached illustration, prepared from a digital image of the
holotype and the rendering of the lip by Dr. Cribb.
* Editor's note: Additional new species from Mexico and
Central America are being described elsewhere in this issue:
see Soto Arenas & Dressier, pp. 285-354.


raceme, the rachis congested, with ca. 11 flowers,
bracts unknown. Ovary sub-trigonous conspicuously
white. Flowers: buds whitish at base, apex green,
with the midrib of petals protruding between the
sepals; flowers very showy, large, perhaps 12-15 cm
diameter when spread out; tepals ivory white, outer,
basal surface of the lip ivory-white, mid lobe and
throat ivory-white lined with yellow-ochre, papillae
of the midlobe yellow; the segments spreading.
Dorsal sepal recurved, apparently long oblanceolate,
acute, perhaps as long as the lateral ones. Lateral
sepals directed downwards, somewhat falcate,
obliquely oblanceolate, obtuse-subacute, base long
attenuate, with a prominent axial vein on the abaxial
surface and corresponding to the axial groove on the
adaxial surface; very smooth, 74-78 x 20 mm. Petals
spreading, somewhat arcuate, slightly recurved at
the apex, narrowly elliptic-oblanceolate, acute-
acuminate, convex, apically conduplicate, broader
than the sepals, longitudinally keeled on the outer
surface, the keel broad and conspicuous, at least 76
long, more than 16 mm wide (not well preserved).
Lip very showy, forming a long tube, marginally
fused to the column at least 2.7-2.9 cm; when spread
out trilobed, the lateral lobes scarcely defined with
rounded shoulders, oblong-triangular, tapering at
apex, ca. 28 x 12 mm; midlobe oblong, rounded, the
base somewhat narrowed and forming an isthmus,
ca. 16 x 15 mm; the disc covered by 7 showy rows
of papillae, longer towards the apex, the papillae
digitiform, up to 3.5 mm high, continuous with warty
veins at the apex of the midlobe. Column unknown.

ETYMOLOGY: This species is dedicated to my dear
friend Mrs. Nora Esponda, administrative assistant at









LANKESTERTANA


-'IP LLA























































HERB.HORT.tOT. TB. KW.

A' 6,,,,".L,. ..^ C -.,/./A...






FIGURE 1. Holotype of Vanilla espondae Soto Arenas, by permission of the Keeper, Herbarium, Royal Botanic Gardens,
Kew.

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS -A new Vanilla from South America


3 K-) 7^.wI '

i< 3 cm I'

LIt i*''



: ''^'


10 cm


sun-


FIGURE 2. Vanilla espondae Soto, prepared from the holotype by Rolando Jimenez M.; the lip based on a camera lucida
drawing by P. Cribb.

the Herbario AMO. Nora's work is evident in every DISTRIBUTION: Known only from the type; from
product of our team and she has been of indispensable the basin of the Rio Magdalena in northern
assistance during our research work. Colombia.

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








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This is one of the most beautiful Vanilla
species. Although it is not closely related to any
other Vanilla, it could be allied to V trigonocarpa,
with which it shares the huge, whitish flowers,
and bright yellow disc; V trigonocarpa lacks the
papillae and warts on the disc. Other species with
similarly adorned lips, like V helleri A.D.Hawkes


or V insignis Ames, have either smaller flowers or
green tepals.
The type specimen consists only of a single leaf,
a picture of the inflorescence, the 2 lateral sepals, a
petal, and the lip, yet the species is so different from
any other described species of Vanilla, that I have little
hesitation in proposing it.


LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA9(3) 285-354 2010


A REVISION OF THE MEXICAN AND CENTRAL AMERICAN SPECIES

OF VANILLA PLUMIER EX MILLER WITH A CHARACTERIZATION OF

THEIR ITS REGION OF THE NUCLEAR RIBOSOMAL DNA 1

MIGUEL A. SOTO ARENAS t23 & ROBERT L. DRESSLER4,5


2 Institute de Ecologia, UNAM, Circuito Exterior s.n. Ciudad Universitaria,
Coyoacan 04510 Mexico D.F., MEXICO
3Herbario AMO, Montafias Calizas 490, Mexico, D.F. 11000, MEXICO
4 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Costa Rica
5 Corresponding author: kerry@bio-photo.com

ABSTRACT. We present a revision of the Mexican and CentralAmerican species of Vanilla. There are 15 different
species in the area; Vanilla costaricensis, V cribbiana, V dressleri, V martinezii and V sarapiquensis are here
proposed as new taxa, and V pompona subsp. pittieri and V pompona subsp. grinrditoin are recognized at
subspecific rank. Vanilla calyculata, V hartii, V helleri, V inodora, V insignis, V odorata, V phaeantha, V
planifolia, Vpompona and V trigonocarpa are also described, illustrated, and their nomenclature, typification,
distribution, and other aspects of interest are discussed. Additionally, we include a key to the species. Several
Mexican and Central American species of Vanilla are closely related to V planifolia, V x tahitensis, and V
pompona, the cultivated species of the genus, and these are thus important in plant breeding. We also include
a cladistic analysis of nucleotidic sequences of the internal transcribed spacer region of the nuclear ribosomal
DNA, showing that this popular molecular marker is of sufficient variation to allow for species discrimination,
permitting, with some exceptions that are discussed, the recognition of sterile samples and indicating that it
is a good molecular marker to infer the phylogeny of this group. The similarity and relationship between V x
tahitensis* and V odorata is discussed.

Se present una revision de las species mexicanas y centroamericanas de Vanilla. Existen 15 species
reconocidas en el area. Vanilla costaricensis, V cribbiana, V dressleri, V martinezii y V sarapiquensis se
proponen aqui como nuevas species, Vpompona subsp. pittieri y Vpompona subsp. gitrndiloifar tambi6n se
reconocen con status subespecifico. Vanilla calyculata, V hartii, V helleri, V inodora, V insignis, V odorata,
Vphaeantha, Vplanifolia, Vpompona y V trigonocarpa tambi6n se described, ilustran y se discuten aspects
de su nomenclatura, tipificaci6n, distribuci6n, relaciones y otros puntos de interest. Adicionalmente, se incluye
una clave de identificaci6n de las species. Varias species mexicanas y centroamericanas de Vanilla estan
cercanamente relacionadas a V planifolia y V pompona, las dos species mas frecuentemente cultivadas, y
por lo tanto representan un germoplasma important para este cultivo. Adicionalmente se present un andlisis

1Editor 's note: Miguel Angel Soto Arenas passed away August 27th, 2009, before this manuscript was submitted for review.
The working manuscript presented here was still unfinished at the moment of his death, and was recovered among Soto
Arenas' electronic files. Despite its economic importance, Vanilla remains one of the most poorly studied of all large orchid
genera, and both the editors and reviewers considered that this publication is an important addition to the orchid and general
botanical literature. Ken Cameron, Phillip Cribb, Eric Hagsater, Gerardo Salazar, and Rodolfo Solano did their best to offer
corrections and comments to the original draft manuscript. In some instances, however, Soto Arenas quoted preliminary
analyses and refers to new taxonomic concepts we were unable to recover. We have not changed the letter of the original
manuscript, and such cases are indicated by editor's notes to warn researchers in the future who will find anomalies or have
concerns about some of the data.
* Editor's note: A paper on Vanilla tahitiensis has been published by Lubinsky et al., Neotropical roots of a Polynesian
Spice: the hybrid origin of Tahitian vanilla, Vanilla tahitensis (Orchidaceae). Amer. J. Bot. 95(8): 1040-1047. 2008. In the
article, the senior author acknowledges Soto Arenas for suggesting the topic of his PhD thesis.








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cladistico de secuencias nucleotidicas de la region de los espaciadores internos transcritos (ITS) de los
genes nucleares ribosomales, donde se muestra que las secuencias de este marcador molecular ampliamente
utilizado son especificas, lo que permit, salvo algunas excepciones que se discuten, el reconocimiento de
muestras est6riles y ademas lo hacen un buen marcador molecular para studios filogen6ticos en este g6nero.
Se comenta la similitud y relaci6n entire V tahitensis* y V odorata, y el origen hibrido de la primera.

KEY WORDS: Orchidaceae, Vanilla, Vanilla calyculata, V hartii, V helleri, V inodora, V insignis, V odorata,
V phaeantha, V planifolia, Vpompona, V trigonocarpa, Mexico, central Americas, new species


The Pantropical genus Vanilla Plumier ex Miller is
a group of orchid vines with about 107 species (Soto
Arenas, 2003; Soto Arenas & Cribb, 2010). Vanilla
is an ancient genus (Chase, 2001; Cameron, 2000,
2003) and the largest in the subfamily Vanilloideae
(Soto Arenas, 2003). Some Vanilla species are
grown as the source of an extract for flavoring and
perfumery, especially Vanilla planifolia Jacks., with
V x tahitensis J.W.Moore and V pompona Schiede
being much less cultivated (Correll, 1944; Purseglove,
1975). Vanilla exports generate US $60-80 million in
foreign exchange for producing countries (Smith et
al., 1992), perhaps more at present, and it is the most
profitable crop of the warm tropics.
The culture of V planifolia and V pompona
originated in Mexico, probably in northern Veracruz
(Bruman, 1948).Although frequently cited as native of
Mexico and elsewhere, there were until recently very
few records of V planifolia from wild populations,
and its precise original distribution is uncertain (Soto
Arenas, 1999). Many Vanilla species are similar
to V planifolia in floral and vegetative traits, and
abundant material in herbaria, both flowering and
sterile, is usually misidentified as V planifolia. Even
some Mexican plantations planted with wild Oaxacan
Vanilla vines include a mixture of V cribbiana, V
insignis, V odorata, and V pompona, besides the
true V planifolia. Similar "mixed" plantations occur
in Ecuador (P Lubinsky, pers. com.) and Guatemala
(Beza, pers. com.).
It is well-known that Vanilla has peculiarities that
make it a taxonomically difficult genus (Wood, 2003),
mostly due to the scarcity of flowering material (a
result of the gregarious, ephemeral flowering, usually
at the top of the forest canopy), the huge vegetative
variation and phenotypic plasticity characteristic of

* V6ase la Nota del Editor en la pagina anterior.

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


the hemiepiphytic growth habit (e.g. leafy and leafless
shoots and differently shaped leaves on the same
individual plant as in many hemiepiphytic aroids; Putz
& Holbrook, 1986; Ray, 1990), the poor preservation
of the membranaceous, strongly three-dimensional
flowers make them difficult to study, the notable
floral similarities between many species, in part due
to the pollination by Euglossine bees (which permits
the maintenance of species with little morphological
differentiation but with different floral fragrances (in
other genera of orchids); Williams, 1982), and the
problems derived from the less than perfect quality
of type specimens. Additionally, some species are
very rare, with sparse populations, and the vines are
long-lived perennials that only flower when they have
attained considerable size. Small pieces of these plants
may be transplanted to greenhouses or gardens, but
few of them ever attain the size and strength to flower.
There are abundant data that indicate that V planifolia
is in danger of extinction in Mexico (Soto Arenas et
al., 2004). Scarcity of material can be illustrated by the
flowering behaviour of the only Mexican population
of V hartii, which was found in anthesis only after 7
years of observations. Vanilla helleri and V martinezii
are each known only from two flowering collections,
while V costaricensis and V sarapiquensis are each
known from a single pressed specimen. Vanilla
phaeantha and V helleri are here reported for the first
time from Mexico, based on sterile material, whose
ITS sequences match with properly identified material
of these taxa. Most cultivated specimens of Vanilla in
botanic gardens and living collections never produce
flowers and therefore cannot be identified.
Vanilla plantations face several agricultural
problems, the most important being the root rot
disease caused by Fusarium batatis f. vanilla
(Childers et al., 1959). It has been suggested that
susceptibility to root rot is perhaps due to a narrow








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


genetic variation, and that is to be expected in this
crop due to its vegetative propagation (Purseglove,
1975; Purseglove et al., 1981; Smith, et al., 1992;
Soto Arenas, 1999; Cibrian, 1999). A study of the
circumscription of the cultivated species, their
distribution, variation, and information on their
related taxa, or those with which they may be likely
confused is a necessary step toward the establishment
of a germplasm bank and a breeding program in this
orchid in order to enlarge the genetic foundation of
the crop. In the absence of a thorough taxonomic
revision, and phylogenetic framework, the breeding
programs with Vanilla have used very distantly
related species that are unlikely to produce fruits with
commercially interesting aromatic properties (e.g.
with V aphylla, or V barbellata).
The taxonomy of the Central American Vanilla
species has been previously recognized as confusing
(Dressler, 1993), and the available regional treatments
are not accurate. Bouriquet (1954) revised the entire
genus Vanilla, including the Central American species,
but his study is out of date and he worked only with
herbarium material. In "The Orchidaceae of Mexico",
Williams (1951) listed four species as native of the
country, one of them, V pfaviana Rchb.f., is actually
a synonym of V inodora Schiede. In recent years,
Soto Arenas (1989, 1994) reported additionally V
mexicana Miller and V odorata. Later, Castillo and
Engleman (1993) have cited significant differences in
morphology, phenology, and compatibility behavior
in the cultivars of V planifolia (which suggested
more than one taxon or a strong genetic structure
within the species), and additional species have been
recognized in Mexico in recent years (Soto Arenas,
2003; Hagsater et al. 2005). On the other hand, V
insignis Ames (1934) was described from Honduras,
V helleri A.D.Hawkes (Heller & Hawkes, 1966) from
Nicaragua, and V. ,p c ititora Dressler (Dressler, 1979)
from Panama. Vanilla hartii Rolfe was reported from
several countries of the area (Correll, 1965), and V
phaeantha Lindl. cited from El Salvador (Hamer,
1974). McLeish et al. (1995) listed three species from
Belize; Hamer (1974) listed two taxa from El Salvador,
and four for Nicaragua (Hamer, 1984), while Heller &
Hawkes (1966) mentioned five from this same country.
Dressler (1993) included three species in his 'Field
Guide to the Orchids of Costa Rica and Panama'. Dix


and Dix (2000) reported four species for Guatemala.
The brief treatment recently published for Costa Rica
(Soto Arenas & Dressier, 2003) lists ten species for
this country, three of then unnamed, and it is largely
based on the present revision.
Special mention is necessary for the enigmatic
V x tahitensis J.W.Moore, described from Tahiti,
but apparently introduced to the botanic garden at
Papeete from Manila, Philippines in 1848, together
with the true Vplanifolia (Petard 1986: 123; Hermann
et al. 1989: 20). Vanilla tahitensis is very similar in
morphology to some Central American taxa, and very
different from the Old World vanillas. No confirmed
records of it, either wild or cultivated exist from
Central America, or in any other American country,
but evidence has been presented of its hybrid origin
from V planifolia and V odorata.
A study of the historical records, literature, and
field work, both in wild populations and plantations,
has proven that some of the reports for Mesoamerica
are based on misidentifications, some species must
be reduced to the synonymy of previously described
taxa, and that some other species of the area remain
undescribed.
This revision of the taxonomy of Mexican and
Central American vanillas, together with a molecular
tool may permit the correct identification of material
of this area, even if it is sterile. This is an important
issue, in view of the necessity of identifying
vegetative material of rare, economically important
plants that seldom produce flowers.
For most Central American taxa, sequences of the
Internal Transcribed Spacers of the nuclear ribosomal
DNA (ITS) were obtained and proved to be species-
specific with a high level of confidence, therefore,
they represent a tool for the identification of sterile
or juvenile living material. The ITS analysis is
compared with sequences gathered from two larger,
more difficult to sequence chloroplast genes (rbcL
and matK) in order to corroborate its utility.


Material and methods

Herbarium and spirit preserved material was
studied in most institutions that house Central
American orchids (AMO, AMES, BM, BR, CHAPA,
CR, ENCB, F, G, GOEL, HEPF, K, INB, LL, MA,
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


MEXU, MO, NY, P, RJ, SCZ, SEL, UCR, US, UV,
W, WU, and XAL.). Efforts were made to study
living material and wild specimens were observed
in Mexico, Guatemala, Costa Rica, and Panama.
Pollination observations and analysis of fragrances
by gas chromatography (conducted by Dr. Neil
da Acosta, Bush Boake Allen, London) were an
additional element in circumscribing some taxa and
are presented in detail elsewhere*.
Due to the phenotypic variation and plasticity seen
in the vanillas and mentioned above, the descriptions
of vegetative features were prepared only from
flowering shoots, often under well-illuminated
conditions. Stems and leaves of descending (shaded,
ground-oriented shoots) are etiolated, often thinner,
or even leafless. On the other hand, Vanilla flowers
frequently show strong shrinkage inflower dimensions
after being pressed; for this reason the measurements
of V cribbiana, V hartii, V inodora, V insignis, V
martinezii, V odorata, V planifolia, V pompona,
and V sarapiquensis were taken from living and/or
spirit preserved flowers. For the rest of the species
the dimensions were taken from pressed specimens,
and those of V calyculata and V trigonocarpa also
include measurements of living specimens published
elsewhere.
In those species in which more than five different
specimens (perhaps not necessarily different clones
in the case of V planifolia) were available, either
in fresh condition or in spirit, the dimensions of
flowering features are also given in mean and
standard deviation, in order to determine the possible
range of variation more accurately. The flowers of
all herbarium material examined were boiled or
reconstituted in a solution of ammonium hydroxide.
Internal Transcribed Spacer (ITS) sequences of
nuclear ribosomal DNA were collected in order to
determine if this widely used molecular marker could
be a useful tool to identify material. The sampling
(see Table 1) does not include V costaricensis nor V
sarapiquensis from which adequate tissue samples
were not available. A special effort was made to
confirm the specific identity of the specimens, even


of some sterile ones. Due to difficulties in preparing
vouchers, some samples remain unvouchered;
however, several are housed at AMO or MEXU;
pictures of other plants are kept in M. Soto's files
(which eventually will be deposited in AMO), but
no voucher specimens are known to exist for most
plants in living collections (e.g. Royal Botanic
Gardens Kew, Nancy Botanic Garden, Jardin
Nacional de Cuba, Marie Selby Botanical Gardens,
Missouri Botanical Garden, Finca La Gavilana),
although they were confidently identified from
the living material, and locality data may help to
eventually confirm the identity of the populations.
Sequences of South American specimens of the
V pompona complex were also included. Also
a sequence of V claviculata, from Puerto Rico,
another of V barbellata, from Florida, and another
from V mexicana Miller, species reported for our
area from unconfirmed records, were also included.
The sequences of Epistephium parritlloruii and
Lecanorchis multillora were defined as outgroups.
It is unknown which is the sister genus of Vanilla,
although morphological data suggest that it is likely
Dictiophyllaria, a poorly known genus which is
known only from the type specimen (Soto Arenas,
2003). Epistephium, Clematepistephium, Eriaxis
and Vanilla form a polytomy in a cladistic analysis
of sequences of the plastid gene rbcL. On the other
hand, Lecanorchis (achlorophyllous, and in which
amplification of rbcL gene has been unsuccessful)
was found to be sister to Vanilla from analysis of
the nrDNA gene 18S, although a more complete
sampling of this region places Epistephium sister to
Vanilla (Soto Arenas, unpublished data)**.
DNA was extracted mostly from fresh leaves
using a method based on Murray & Thompson
(1980), or from leaves preserved in silica gel (Chase
& Hills, 1991). Better quality DNA was obtained
using liquid nitrogen and adding 2% of PVP to the
extraction buffer. Vanilla tissues oxidize badly when
they are pressed, especially in the Membranaceous
species; all attempts to extract DNA from herbarium
specimens failed, although bad quality DNA has


* Editor 's note: The manuscript "Notes on the floral biology of mexican Vanilla (Orchidaceae) and the evolution of
pollination systems in the genus" has been found among Miguel Angel's unpublished manuscripts.
** Editor 's note: It is unclear to which data the senior author refers here.


LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.










SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


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SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


been extracted from a couple of South American
and Asian herbarium specimens of other species.
The ITS region, including the 5.8S gene was then
amplified with the primers ITS2, ITS3, ITS4, and
ITS5 (White et al. 1990; Baldwin, 1992) or 17SE
and 26SE of Sun et al. (1994). PCR reactions
were better when 3% of DMSO was added to the
cocktail. PCR products were run in low melting
point agarose gel and the band cut to be extracted
with QIAGEN Gel Extraction Kit (QIAGEN,
Ltd.) or QIAquick silica columns (QIAGEN, Ltd.)
adding guanidin chloride (35%) to remove primer
dimers. Bi-directional sequencing was performed
using cycle-sequencing (ABI Prism dye terminator
cycle sequencing ready reaction kit, PE Applied
Biosystems, Inc.) with the same primers with which
they were amplified using different automated
sequencers following manufacturer's protocols.
Electropherograms were edited using EditView, and
the resulted sequences were initially aligned using
Clustal x (Thompson, 1995) and adjusted by eye.
Although alignment of Vanilla species is easy, the
alignment with the outgroups is largely ambiguous
and alignment is based mostly on the lengths of
the regions. Phylogenetic analysis was performed
with PAUP* 4.0 (Swofford, 1998) with the
following specifications: Epistephium pairitlloriu
and Lecanorchis multitilor, were defined as the
outgroups. Anheuristic search with equally weighted
parsimony analysis. The analysis consisted of 1,000
replicates with SPR swapping and 20 trees saved
per replicate, to save time swapping on islands with
large numbers of trees and allowing the detection
of multiple islands of equally parsimonious trees
(Madison 1991). Then another equally weighted
parsimony analysis was performed, swapping
to completion of all the trees from the previous
analysis. This was followed by a bootstrap analysis,
holding 10 trees per replicate for 1000 replicates,
using SPR swapping and MULPARS. Bootstrap
values were obtained from 100 replicates.


Results

We recognize 15 Vanilla species in Mexico and
Central America. A key and a taxonomic treatment
are presented in the following pages. Five species,


Vanilla costaricensis, V cribbiana, V dressleri, V
martinezii, and V sarapiquensis are proposed as new
species and three subspecies are recognized within V
pompona (the typical subspecies, subspp. pittieri and
gria, Ilitlor 1n.
Amplification of PCR products was better with
primers 17SE and 26SE of Sun et al. (1994) and with
them only a single band was obtained. Two bands,
one probably from and endophytic fungus were
sometimes amplified using primers ITS4 and ITS5.
The analysis of nucleotidic sequences resulted in
669 characters, from which 246 were constant, and
271 sites were parsimony-informative. The heuristic
search yield a total of 39,090 most parsimonious trees
(tree length = 1070). The strict consensus of this tree
is presented in Fig. 1.
This tree is largely congruent with phylogenetic
analysis based on morphological and molecular
characters, either of plastid (rbcL gene, Soto 2003;
matK gene, Soto & Alvarez-Buylla, unpublished),
of nuclear sequences (18S and ITS sequences from
species from all the world). In all the analysis two
main groups of Vanilla are recognized. One of them
includes the Membranaceous vanillas (V inodora
and V martinezii in the tree; V costaricensis
and V sarapiquensis, from which DNA was not
available belong to this clade). The other lineage
of Vanilla includes the leafless plus the leafy, non-
membranaceous species.
Two very dissimilar sequences of V planifolia
were obtained from two different PCR reactions of the
specimen Dressler s.n. (cultivated at FLAS). One of
these sequences is practically identical to other eight
sequences of the same species included in the sample.
The anomalous V planifolia sequence goes sister
to a V mexicana sequence, forming a clade which
occupies a basal position to the rest of Vanilla. When
Clematepistephium smilacifolium and Eriaxis rigida
are included into the analysis, this clade occupies an
intermediate position between Clematepistephium-
Eriaxis-Epistephium and Lecanorchis (data not
presented). Vanilla mexicana is the type species of
the genus, and closely related to, even difficult to
separate from, other membranaceous vanillas (e.g.
V martinezii and V inodora). These anomalous
sequences make both Vanilla and the defined
outgroup non-monophyletic. The rbcL (Soto Arenas,
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.









LANKESTERIANA


VerjW odror. 4
Venge odweor 5
Venfle ~odrta S
VwMI~ .11 odrwr~
VwWO. WiNW&f 1
Venl~e iehi~nas 2
Vw~Ee of. hurfg

77 Vane hefrul
TT I IVenlle odoJ~tai
100 OYARO 0001WO 1
71 Von oidOYNaO 2
Vanie MaruMag 2
92 V8aw inki~mw 3

Vannia pwknffo"a 2
v~nW pfta" 3

Vanif paniOW 1a
Vsnlwp wafti I6a
VJw ApW9r0P 14
VaniW. paW CI1A22
Vnme phowifh. 1
Vnwe phooflI& 2
V6n5e phMarWfi' 3

Venle dmsWof# I


van08 M53kni 3



100 veapo~mpone B
INVaW mpbo.oe i po n I
VWIA POeRAOn2S ioehe 2
00 VW08 crgt~cloonff

Ve i paompanoon Prn 4
VSMP pompnopor n&mp 2

vese~sPorlonsootekmfilJ S

1I VsnAJ2a npinOs pi~ti 1
V9nnV p paomni pompw 2
92venla POM1PO2 -,ad 4


Van We pomponm pmt~ 4
VaanI. pomponspided 5
93 Van~ia pfOMPns gmandrfa 1
= Van ponpan P WnumIrvu 2
100I I vsnwe M wukults I
VWS xr 2
100 =EEW4~

VWWO~ Ovonocarm I
100 VWWO O"Wamerg 2
100 V~nrW* Vnawa" 3
Varew haffw I
VwW& harfff 3
97 96 V&W& h~o arlff 4
VenW&e dowtua 1
Vexie borowt 3
100 Yonit knodQw I
100 --- VlWW in0dj 4
Ven~e mena~ena
10 VA& kdb
Vwni~e pmw5oie Ie
EpLafph2um paa~n
LeF3no0a iiT malDNA







FIGURE 1. Strict consensus from 30,090 minimal trees obtained from ITS sequences of the nuclear ribosomal DNA.


LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


2003), matK, and 18S topologies (Soto Arenas et al.,
in prep) are incongruent with the position of these two
particular sequences of Vanilla and we regard them as
paralogous ITS copies. Cloning in 10 PGEM a PCR
product of V. cf. kaniensis Schltr. (a species from
New Guinea) yield three different ITS clones which
differ only in 1 and 2 DNA bases and are clustered
together in the ITS phylogeny, therefore ITS is a very
useful identifying tool.
Other paralogous ITS sequences of Vanilla (from
species of other regions) have occasionally resulted,
especially when DNA has been extracted from
herbarium specimens or less often from tissues dried
in silica gel. No more paralogous copies of American
vanillas have been detected, and both included
represent the two main lineages in the genus. If they
are excluded, both the defined outgroup and Vanilla
become monophyletic. The paralogous sequences
are largely divergent, and therefore easily identified.
This is probably because the duplication event that
produced them seems to have been previous to the
separation of Vanilla from their related groups.
The leafless group is not well-documented from
Mesoamerica (see discussion in 'Excluded taxa');
a Puerto Rican specimen of V calyculata and
another of V barbellata were included. There are
no confirmed reports of these species from Mexico
or Costa Rica, respectively; in any case, it is rather
surprising that leafless taxa are absent in Yucatan,
very close, and with similar habitats to places in
Cuba where leafless vanillas are present. The rest
of the Mesoamerican vanillas belong to the group
recognized by Soto (2003) as "American fragrant
Vanilla spp." Phylogeny of Vanilla is discussed in
more detail in Soto (2003).
What is important for identification purposes is
that ITS sequences are largely species-specific, since
the species form monophyletic groups with very high
bootstrap support.
The only exceptions are V odorata which proved
to be paraphyletic, since V helleri and V tahitensis
are nested in it. Two samples from east of the
Tehuantepec Isthmus share a different ITS sequence.
Vanilla tahitensis sequences are not different from
most of V odorata, a result which is not incongruent
with a the hypothesis of a recent hybrid origin of
this taxon (between V planifolia and V odorata),


as its morphology suggests. The Vanilla pompona
complex has a bootstrap value of 99%. All the
Mexican samples of V pompona plus two specimens
(SEL 1985-0211A and a sequence in the Gen Bank
U66819) form a clade with 99% of bootstrap support.
Several South American specimens of V. giranitlorai
form a weakly supported (63% bootstrap value)
branch, and the rest of the South American specimens
of V pompona and the Costa Rican, Honduran, and
Panamanian specimens of the V pompona complex
form a polytomy at the base. This structure in the V.
pompona complex is congruent with the distributional
disjunctions and suggests that an infraspecific
classification of the species is required.
In conclusion, ITS sequences are a very useful tool
to separate sympatric, vegetatively indistinguishable
vines like V planifolia, V helleri, and V cribbiana,
or to identify specimens with juvenile or etiolated
vegetative characters. We suspect that one of the
reasons for the lack of an important breeding effort in
Vanilla has been due to the difficulties of identifying
material.
The origin of V tahitensis has been much argued.
These data indicate clearly that V tahitensis is very
closely related to V odorata. Vanilla tahitensis is
clearly a member of the American fragrant clade of
vanilla, characterized by leafy plants and penicillate
calluses, and which is absent in the Old World Tropics
(as in New Guinea) where the vanillas geographically
nearest to Tahiti occur), therefore the hypothesis
that it was introduced recently to Tahiti is strongly
supported. Since vegetatively it has much broader
leaves than V odorata, it is almost certain that it is
not conspecific with this species. The flower is larger
than in Vplanifolia, and it is rather intermediate in all
traits between V planifolia and V odorata.
These molecular data, and other unpublished
data including many more species from other areas,
suggest that V phaeantha, V insignis, V helleri, V
odorata, and V tahitensis are the closest relatives of
the vanilla of the commerce, V planifolia, even much
more closely related than V pompona with which
it has formed hybrids that are at present cultivated.
The close relatives all have fragrant fruits and have
different habitat preferences; thus they represent
a pool of interesting traits with potential to be
incorporated into this crop.
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


KEY TO THE SPECIES AND SUBSPECIES

1. Leaves thin, membranaceous when dry, sometimes slightly chartaceous, rachis of the inflorescence lax with
remote bracts; lip united only basally to the column (< 4 mm), column smooth or basally keeled, but without
substigmatic hairs; lip without a penicillate callus formed by a tuft of laciniate scales, instead, the callus may
be form ed by longitudinal keels, or very fleshy cushions .............................................................................. ............................. 2
2. Bracts of the inflorescence foliaceous, very large, similar to small leaves ................................. ............................ 3
3. Lip distinctly trilobed, midlobe emarginate-bilobed, callus massive, cushion-like ....................... V inodora
3. Lip subentire, truncate at apex, callus made up of 2 prominent axial keels and several warty lateral
k e e ls ..................................................................................................................................................................................................... V. c o s ta r ic e n s is
2. B racts of the inflorescence scale-like, sm all (< 1 cm long) ........................................................................................................... 4
4. Lip ca. 42 mm long, with low callus made up of several axial, longitudinal, warty keels........... V martinezii
4. Lip ca. 32 mm long, with a massive callus made up of 2 broad, fleshy, sulcate, apically confluent
k e e ls .................................................................................. ............................. ........................................... ............................. s a r a p iq u e n s is
1. Leaves thick, coriaceous to fleshy, chartaceous when dry; rachis of the inflorescence dense with approximate
bracts; lip attached to the column up to the stigmatic region, usually > 2.5 cm, column with trichomes on the
ventral surface, lip with a penicillate callus formed by a tuft of laciniate, retrorse scales............................................. 5
5. Lip without a clear claw, rather cuneate, strongly trilobed, the midlobe short, transversely oblong and
covered by conspicuous, complanate papillae along nerves...................................................................................V helleri
5. Lip conspicuously long-clawed, claw usually very distinct from the blade, the latter entire to trilobed, the
midlobe variously shaped, smooth or variously covered with papillae, trichomes or warts ...................... 6
6. Flowers very large, lip > 10.5 cm long, inflated, margins crenulate-plicate, racemes few-flowered
(2-3) ........................................ trigonocarpa............................................................................................................................................................. V trigonocarpa
6. Flowers not so large, lip < 9 cm long, inflated or not, margins not crenulate-plicate, entire to denticulate
or only plicate; raceme several-flowered, usually with > 6 flowers, often much more................................. 7
7. Leaves conspicuously shorter than the intemodes, lip almost smooth, without conspicuous
appendages, except by minute warts at the apical part of the axial veins 8
8. Leaves acuminate; lip < 41 mm long, subentire ................................................................................... ... V hartii
8. Leaves obtuse to subacute, sometimes mucronate; lip > 60 mm long, trilobate at apex.......................
......................................................................................................................................................................... V p h a e a n th a
7. Leaves usually as long or longer than the intemodes, lip apex covered with papillae, retrorse
trichomes or the veins thickened, wavy and complanate-warty......................................................... 9
9. Lip fringed or denticulate at margins, with an apical thickening with warts, papillae, or retrorse
appendages, flowers mostly whitish-green, weakly scented...................................... .................. ..... 10
10. Stems sulcate, minutely papillose, flowers large, the lip more than 65 mm long, margin
long laciniate-fimbriate, midlobe covered by retrorse appendages up to 4 mm high .................
insignis................................................................................................................................................................................................... insignis
10. Stems non-sulcate, smooth, flowers smaller, the lip less than 65 mm long, margin laciniate,
erose or denticulate, midlobe with few (less than 5) retrorse papillae or warty .......................... 11
11. Leaves ensiform, long acuminate; lip margin fimbriate .............................................. V odorata
11. Leaves oblong to elliptic, acute or abruptly acuminate; lip margin undulate-denticulate,
not fimbriate ............................................................... planifolia
9. Lip entire or undulate at margins, without apical thickenings or if present without warts, papillae,
or retrorse appendages, flowers mostly cream-yellow, with strong and spicy scent .......................... 12
12. Midlobe of the lip longer than wide, emarginate, flower bell-shaped, with the apex of
the tepals reflexed ....................... .............. ...................... ......... ......................................... V calyculata
12. Midlobe of the lip broader than long, truncate to somewhat emarginate, flower
trumpet-shaped, the apices of the tepals not reflexed ............................. ................................ 13


LANKESTERIANA9(3), January 2010. 0 Unversidad de Costa Rca, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


13. Lip rhombic, with thickened, flat veins, leaves cuspidate .............................................. ................................. V. dressleri
13. Lip flabellate, obscurely trilobed, with a longitudinal, cushion-like thickening, leaves obtuse to acuminate 14
14. Stems 3-5 mm thick, leaves 10-22 x 2.3-7.5 cm; lip apex truncate, entire, ovary and sepals fairly
papillose-verrucose ..... ......................................................... V. cribbiana
14. Stems 10-24 mm thick, leaves 22-29 x 8-14 cm; lip margin undulate, ovary and sepals smooth ...............
............................................................................................................................................................................................................................... V... pompona

TAXONOMIC TREATMENT


1. Vanilla calyculata Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 7: 42-43. 1920.

TYPE: Colombia, Cauca: 1000 m M. Madero,
not located; neotype (here designated): VALLE:
Municipio de Tulua, Corregimiento Mateguadua,
Jardin Botanico, laderas en via de repoblacion
natural. Altura 1100 m. Enredadera; sepalos verde
claro, petalos amarillo claro, labelo amarillo intense,
frutos maduros color marr6n; muy fragantes. 29
sept. 1984. W Devia 815 holo. MO (3245054)!

COMMON NAMES: "Vanilla"

Hemiepiphytic vine, branching, leafy, up to ca. 4
m high. Stems flexuose, terete, green, 6-12 mm thick;
intemodes 7-15.5 cm long. Aerial, free roots apparently
subterete, pale brownish, up to 8.2 cm long, 2-3 mm
thick. Leaves sessile to subpetiolate, the petiole up
to 7-10 mm long; the blade oblong-lanceolate, rather
narrow, the base rounded, the apex acute, coriaceous-
fleshy, apparently stiff and xerophytic, the margins
slightly revolute, 7.5-21 x 1.7-4 cm. Inflorescence
a 12-14-flowered receme, 4.2-6 cm long, rachis
terete, green, ca. 4-6 mm thick. Bracts widely ovate,
obtuse, green, concave, progressively smaller, up to
10 x 6 mm. Flowers opening successively, 2-3 open
at once, pendant, bell-shaped, very showy, sepals
pale green to yellowish green, petals pale yellow, lip
deep yellow, ca. 9 cm long; strongly fragrant. Ovary
terete, smooth, sulcate, the grroves twisted, ca. 4-4.5
mm long, 4 mm thick. Dorsal sepal oblanceolate to
narrowly elliptic, apex subacute-obtuse, somewhat
thickened, subcalyptrate, recurved, base attenuate-
subunguiculate, basally canaliculate, convex above,
smooth, ca. 12-13-veined, 66-80 x 8-17 mm. Lateral
sepals oblong to narrowly elliptic, oblique, the lower
margin more curved, apex subacute-obtuse, somewhat
thickened, subcalyptrate, recurved, basally attenuate-
subunguiculate, smooth, ca. 12-13-veined, 67-80


x 12-17 mm. Petals obliquely oblanceolate, lower
margin more arcuate, apex attenuate, rounded, very
attenuate at base, acute, with an axial, flat keel on the
abaxial surface, 1 mm wide, ending in a free, conic,
blunt, short, 1 mm long process; ca. 11-12-veined,
65-80 x 9-16 mm. Lip attached to the column along
the margins of the basal half (ca. 41-46 mm), tubular,
cymbiform, deeper near the middle; axially grooved
on the abaxial surface, the groove well-defined
and deep, when spread out 78-90 x 36-45 mm; long
unguiculate, the claw apically rugose-papillose on the
inner surface, 26 x 3-3.8 mm; the blade trilobed, ca.
36-veined, veins branched, the lateral lobes forming an
inflated tube around the column, when flattened widely
triangular-semiovate, margins entire, undulate near
the midlobe, ca. 37-39 x 14 mm; midlobe dilated and
recurved, approximately transversely oblong, deeply
emarginate-bilobed, margins crenulate-undulate,
veins conspicuously thickened, 14-16 x 21-26 mm;
penicillate crest at ca. 41 mm from the base, 4.5 x
3.8 mm, made up by 8-10 flabellate, shortly lacerate-
laciniate, retrorse, scales, some of them united to each
other along the lateral margins; disc with progressively
more thickened veins from the center to the apex of the
lip, forming an apical, swollen, low cushion, obclavate
in outline, rugose-papillose at the apex, 25-27 x 6 mm,
2-3 mm high; with a group of transversal, yellow-
orange hairs at the basal part. Column very elongate
and slender, subtrigonous-subclavate, slightly sigmoid,
51-55 mm long, ca. 4 mm wide; ventral surface flat
and lanuginose at the distal half; apex dilated (6 mm
wide) with vertical wings, narrow, triangular-flabellate,
undulate, ca. 4 x 1 mm. Stigma trilobed, the lobes
emergent; rostellum trapezoid-flabellate, 2 x 4.5 mm;
lateral lobes transversely oblong-flabellate, 1.2 x 1.8
mm. Anther versatile, attached to the wide clinandrium
margin by a broad filament, triangular-ovate, truncate-
emarginate, 5 x 5 mm. Fruit fragrant, 8-15 cm long,
apparently thick and trigonous. Fig. 2, 17A.

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.









LANKESTERIANA


5mm


CM

"* I





































S~. 5mm



FIGURE 2. Vanilla calyculata Schltr. Based on published pictures (Hamer, 1981) and Devia 815. Drawing by R.
Jimenez.

LANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.


ai,
--








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


DISTRIBUTION: Along the Pacific slope of El Salvador,
Honduras, Costa Rica, and Colombia; a fruiting
specimen from Michoacan, Mexico, may belong here.
ECOLOGY: Hemiepiphyte at relatively high altitudes
(800-1300 m), often in rather dry places. Flowers
recorded in April (El Salvador and Honduras) and
September-February (Colombia).
The identity of V calyculata has been obscure
and no original specimens are known to us. However,
Schlechter's description can be applied with
confidence to the taxon above described and illustrated
in Fig. 1. No other species known from Colombia (V
columbiana, V dressleri, V espondae, V hostmanii, V
methonica, V odorata, V pompona, V ribeiroi, and V
sprucei) agrees with the description of V calyculata.
The specific epithet makes reference to the calyculus,
a minute, cup-like structure sometimes distinguished
in some vanillas and more developed in the species of
the related genera Epistephium and Lecanorchis. The
calyculus in V calyculata is rather conspicuous when
compared with other vanillas, and is more evident
in pollinated ovaries that start to swell. Schlechter's
description differs from our material of V calyculata in
the column described as glabrous instead of pubescent
below the stigma (a trait found in all American
penicillate Vanillas) and its smaller measurements of
the perianth segments. We believe that Schlechter's
description could had been based on a bud. However,
the calyculus, the sessile, oblong, rather small leaves,
the deeply excised midlobe of lip, the pubescent lip claw,
the 3-5 thickened midveins on the disc, the unusually
high altitude for a Vanilla and its strong resemblance
with the Brazilan V chamissonis (its sister species, Soto
unpublished) suggest that the name V calyculata may
be applied to this taxon with confidence.
Vanilla calyculata is distinguished from V
chamissonis by its larger, bell-shaped, pendant,
yellowish flowers with revolute, flaring tepals, and the
longer midlobe of the lip.
Despite its unique characters, specimens of V
calyculata have been regarded by Reichenbach and
Rolfe (1896; e.g. Lehmann 2263) as V pompona.
Later it was reported from El Salvador (Hamer, 1974)
as V phaeantha Rchb.f Jose Linares, from Escuela
Agricola Panamericana has told us that this species is
abundant, well-known, and usually confused with V
pompona in the dry valleys of central Honduras.


The specimen O. Pank sub F Hamer 203 bears
peloric flowers where the column has two anthers and
two very odd stigmas.

OTHER RECORDS: EL SALVADOR: La Palma-Finca El
Refugio, rio Nanuapa, 1000 m alt., terr. climbing oaks or
manzana rosa, collected 3.4.1969. April 11 1969, 0. Pank
y F Hamer 203 AMES(*112862; *113837)! HONDURAS:
COMAYAGUA: Fruit triangular, up to 6 in. long. Climbing
in shrubs, thicket along river, plain near Sihuatepeque. 1050
m altitude. 7/23/36, TG. Yuncker, R.F Dawson & H.R.
Youse 6045 *AMES(46667)! G! K! *MO(1115382)! NY!
MORAZAN: flores cremas, bejuco sobre arbustos. Aguas
abajo de la Quebrada de Sta. Clara. Alt. 850 m, Abril 28,
1948, A. Molina 808 *F(1676153)! Bejuco, creciendo
sobre rocas y arbustos, flores amarillentas algo fragantes
(por la tarde). Quebrada Santa Clara, ca. 2 km al norte del
Zamorano, alt. 800 m. Mpio de San Antonio de Oriente. 19
de junio de 1996. J. L. Linares 3386 MEXU! El PARAISO:
Mpio. Moroceli, Quebrada El Cajocote conocida tambi6n
como El Terrero, 8.7 km al N de Moroceli, por el camino
a Mata de Platano. Veg. riparia, veg. circundante selva
baja caducifolia y bosque de pino-encino, 14 10'10" N,
8651'06"W, 680 m. Bejuco creciendo sobre Guettarda
macrosperma, flores amarillo palido con el apice amarillo
oro, olor suave y dulce, 29 abril 2004, J. L. Linares 7313
MEXU! [cf., sterile] "Vainilla", trepadora sobre arboles y
matorrales, de Barranco de las Mesas, Alt. 900 m, Agosto
1, 1957, L.O. Williams & A. Molina 8575 F(1590341)!
Open savannah, Las Mesas region near Yuscaran. August,
1960, H.W. Pfeifer 1454 *US(2563382)! COLOMBIA:
TOLIMA: Fingerdick, bis 5 m lang. Bl. dickfleischig &
dunkelgrin. Bit. grinlich meist mit heller Lippe. Frachte
bis 8 cm lang. Auf Baumen in Zavannen -Waldern bei La
Plata. 800-1500 m. 3.12.1882. EC. Lehmann 2263 BM! G!
Vanilla, white and red inside, cultivated E. Dryander 2379
BM! "Columbien" Lehmann W(59043, in part, the flower
and probably the right shoot)!
The following sterile specimens may belong here:
MEXICO: MICHOACAN: Hacienda de Coahuayula,
Feb. 1901, G.M. Emrick 16 F(95476)! COLOMBIA: [cf.,
leaves atypically broad] TOLIMA: La Plata, 1000-1300,
*Lehmann 6278 AMES(14875)!

2. Vanilla costaricensis Soto Arenas, sp. nov.

TYPE: COSTA RICA: ALAJUELA: Llanura de
San Carlos. Alt. 200 m., Feb. 20, 1966. Lip white,
tepals same green as leaves; vine. Lowland rain
forest between Los Chiles and Venecia. A. Molina
R., L.O. Williams, WC. Burger and B. Wallenta
17565, holo. CR(062310)! iso. MO(2367689)!

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


Species Vanillae oroanae similis floribus minoribus,
labello integro breviore differt.

Hemiepiphytic vine, leafy. Stems keeled, 8 mm
thick (in dried condition); intemodes ca. 12.5 cm
long. Aerial, free roots dorsiventraly compressed,
pale brownish, ca. 2.5 mm wide. Leaves petiolate, the
petiole canaliculate, ca. 17 mm long, 9 mm wide; blade
elliptic, abruptly mucronate, base acute to attenuate,
green, membranaceous (in dried condition), 21-22 x
8.7-9.5 cm. Inflorescence similar to the vegetative
shoots, but smaller, elongate, a 3-4-flowered raceme
(probably longer and more floriferous when completely
developed), 28-32 cm long, intemodes up to 6 cm long;
peduncle 15-23 cm, rachis ca. 3 mm thick. Bracts
foliaceous, subpetiolate, the petiole ca. 3 mm long;
blade elliptic, acute-acuminate, base obtuse-rounded,
membranaceous, 2.9-7.3 x 1.6-3.4 cm. Flowers
successive, 2 open at once, with spreading segments,
ca. 3.5 cm in diameter; tepals green, lip white. Ovary
rather sigmoid, 34-40 x 3.5 mm. Dorsal sepal strongly
twisted and the margins contorted-widely undulate,
elliptic, apex acute, rounded, base cuneate, ca. 10
veined, smooth, 41 x 11 mm. Lateral sepals narrowly
elliptic, acute-subacute, base obtuse, margins undulate,
reflexed, smooth, ca. 10 veined, 41 x 10.5 mm. Petals
narrowly elliptic, apex truncate, notched, base widely
cuneate, strongly twisted, widely undulate-contorted,
with the lateral margins reflexed, difficult to spread out
without distortion, ca. 10 veined, 35.5 x 9 mm. Lip
attached to the column less than 2 mm, slightly arcuate,
almost straight, with the lateral margins erect forming
a throat around the column, quadrate-flabellate, entire,
apex truncate, obcurely trilobed, the lateral lobes as
long as the midlobe, or slightly longer, base rounded-
truncate, the apical margin entire to dentate, ca. 26
veined; 24-27 x 20-21 mm; callus made up of a pair of
flat, broad keels along the axial line, from the base to
the beginning of the apical third, then separated in 3 low,
erect, congested keels ending at the apex; additional
lateral keels with complanate warts adorn the basal
half. Column relatively short, straight to very slightly
arcuate, semiterete, with a dilated apex, 16 mm long;
smooth, except by the callus of verrucose keels at base,
ca. 5 mm long; vertical wings flabellate, ca. 1.5 x 3 mm.
Stigma lobed, the midlobe, convex, very prominent,
almost perpendicular to the column axis, ca. 2 x 1.5
mm; lateral lobes much smaller, fused in an ovate-

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


subquadrate, emergent blade, ca. 0.5 mm long. Anther
galeate, protruding at apex, laterally compressed, 4 mm
long, with a long, thick filament 1.2 mm long. Fig. 3.

DISTRIBUTION: Known only from northern Costa Rica.
ECOLOGY: In lowland rain forest at 200 m altitude;
flowering in February.
This species is known only from the type locality
and it has been misidentified as V mexicana Miller, a
different, closely related species from the West Indies
and northeastern South America. Vanilla mexicana
has a distinctly trilobed lip, with the apex of the
lateral lobes slightly above the middle of the lip; in V
costaricensis the lip is entire, flabellate, with the apex
truncate and just obscurely trilobed, but the lateral
lobes of the apex are subequal or slightly longer than
the midlobe; additionally, the keels at the apex of the
lip in V mexicana are much more conspicuous. Vanilla
costaricensis has large bracts similar to those of V
inodora Schiede, easily distinguished by the cushion-
like, axial callus on the lip. Also closely related are
V guianensis and V martinezii with much larger
flowers, straight sepals, more rugose keels, and shorter
inflorescence, with reduced bracts. The species was
illustrated by Dr. Rafael Lucas Rodriguez (Rodriguez
C. et al., 1986) based on a specimen that has not been
located.
The closest relative of Vanilla costaricensis is
V oroana Dodson of Ecuador. They have similar
inflorescences and flower morphology, with the
column almost identical. However, V oroana has a
longer, relatively narrower, trilobed non-truncate lip,
and the surface of the lateral, basal part is minute, but
conspicuously more sculptured than in V costaricensis.
Vanilla oroana flowers are in general much larger.
In the examined material of V costaricensis (and
also V oroana) the lateral lobes of the stigma are fused
to each other to form an ovate-subquadrate, emergent
blade. This is notoriously different from the concave
stigma found in other membranaceous (e.g. V inodora).
However, observation of this trait is difficult because
of the poor preservation of the column, and the large
amount of sticky substance in the stigmatic area, visible
in fresh material but absent in pressed specimens.

OTHER RECORDS: COSTA RICA: ALAJUELA: Llanura de
San Carlos. Alt. 200 m., Feb. 20, 1966. Lip white, tepals
same green as leaves; vine. Lowland rain forest between
Los Chiles and Venecia. A. Molina R., L.O. Williams,








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


I 3cm














-
3cm







2 mm

4mm







.-.








10 cm








FIGURE 3. Vanilla costaricensis Soto Arenas. Based onA. Molina et al. 17565, except floral dissection and column
based on A. Molina et al. 17567 Drawing by M. Lopez.


LANKESTERIANA 9(3), January 2010. C Universidad de Costa Rica, 2010.








LANKESTERTANA


W.C. Burger and B. Wallenta 17567 *SEL(016793)!
*F(1771938)!

3. Vanilla cribbiana Soto Arenas, sp. nov.

TYPE: MEXICO: CHIAPAS: Mpio. Ocosingo:
Estaci6n de Biologia Chajul, en el borde del Rio
Lacatum; a 680 m del Puente Hamaca, por la
vereda a Arroyo Miranda, ca. 180 m s.n.m., selva
alta perennifolia en piano, ca. 8 m de alto, flores
con tepalos blanco-cremosos, sepalos verdosos en
la superficie extema, labelo atrompetado, amarillo
intense con rayas anaranjadas en las venas, labelo
extendido y truncado, sin papilas, no deflexo en el
apice, flores bien abiertas. Fragancia muy intense
y conspicua, a mentol y citrico. Ovario verde-
blanquecino con papilas. 20 junio 1996, M. Soto
7945 yR. Solano, holo. *AMO!; iso. K !, AMO (in
spirit)!

Vanillae hostmanii primo adspectu similis, sed foliis
minoribus, inflorescentiis minoribus paucifloribus,
labello obscure 3-lobato, lobo medio distinct,
subquadrato, quinque nervis papillosis incrassatis in
centro apice omato versus circa decem nervous in lobos
laterales extensos

COMMON NAMES: "Vainilla".

Hemiepiphytic vine, somewhat branching, leafy,
up to 12 m high. Stems terete, smooth, green, 3-5 mm
thick; intemodes 3-9 cm long. Aerial, free roots pale
subterete, brownish, ca. 2 mm thick; ,ill.i. clii, aerial
roots conspicuously dorsiventrally compressed, 2-5
mm wide. Leaves subpetiolate, petiole 1-1.5 cm long;
blade obliquely elliptic to elliptic-oblong, abruptly
acuminate-apiculate, stiff, brittle, 10-22 x 2.3-7.5 cm.
Inflorescence a ca. 10-flowered raceme (rarely up to
30 flowers), 30-37(-110) mm long, rachis 18-22(95)
mm long, 4 mm thick. Bracts ovate-triangular, obtuse,
very concave, progressively smaller towards the apex,
up to 8 x 4 mm. Flowers successive, 2-3 open at
once, with spreading segments, apparently ephemeral,
very showy, white-cream tepals, sepals externally
cream-greenish, lip deep yellow with orange stripes,
ca. 8 cm diameter; fragrance strong, mentholate and
citric, similar to that of V pompona. Ovary arcuate,
dorsiventrally slightly compressed, thickened at the
very base, whitish-green, conspicuously papillose,

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


more densely towards the perianth, sulcate, the grooves
twisted, 44-46 mm long, 4-5.5 mm thick. Dorsal
sepal narrowly elliptic, apex subacute-rounded,
subcalyptrate, minutely apiculate, base attenuate, flat,
apically concave, 9-veined, minutely papillose on the
abaxial surface, the papillae in longitudinal rows and
forming transverse, undulate rows, very fleshy and
stiff; 63-65 x 12.5-13.5 mm. Lateral sepals obliquely
oblong-elliptic (upper margin curved, lower margin
more straight), apex subacute-rounded, subcalyptrate,
base attenuate, canaliculate basally, slightly convex,
apically concave, apex somewhat recurved, margins
conspicuously involute, ca. 12-14-veined, minutely
papillose in the abaxial surface, the papillae in
longitudinal rows and forming transverse, undulate
rows, very fleshy and stiff; 60-65 x 12.5-14 mm.
Petals oblanceolate, very slightly oblique (the lower
margin more straight), slightly arcuate, apex obtuse,
widely rounded, somewhat thickened at apex, base
attenuate, basally canaliculate, slightly concave
towards the apex, with an elevated, axial, flat keel on
the dorsal surface, ending in a long triangular, flat,
free (ca. 2 mm) process; dorsal surface conspicuously
canaliculate apically, surface colliculate, the cells in
longitudinal rows, 14-veined; with granular, thread-
like, somewhat branched inclusions; 64 x 12.5-13.5
mm. Lip attached to the column along the margins
of the basal half (ca. 28 mm), tubular, trumped-
shaped, cymbiform, deepest near the middle; axially
grooved on the abaxial surface; when spread out 50-
52 x 38 mm; unguiculate, claw with 4 obscure bands
of unicelluar, minute, elongate, yellow-brownish,
trichomes; the blade obscurely flabellate, trilobed,
margin conspicuously entire, subtruncate, lip with
inclusions similar to those found in the petals; lateral
lobes widely and obliquely triangular, overlapping
above the column, 33 x 16 mm; midlobe small,
subquadrate-ovate, slightly deflexed, smooth, the
apex flat, ca. 8.5 x 14 mm; penicillate callus made
up by ca. 10 congested, retrorse, trapezoidal, laciniate
scales, the scales sometimes united to each other along
the lateral margins, ca. 5 x 4 mm; the region just after
the penicillate callus, smooth, with denser inclusions;
with 5 low, rugose, rounded, densely papillose keels
confluent in an apical cushion-like thickening, 4-6
secondary, more inconspicuous keels, shorter, not
reaching the apex. Column elongate, 34 mm long,









SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


1cm


1 mm


2cm



FIGURE 4. Vanilla cribbiana Soto Arenas. Based onM. Soto 7945. Drawing by M. Lopez.

IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








LANKESTERIANA


3 mm wide; ventral surface pubescent-lanuginose at
the middle, hairs yellowish; apex dilated, 4 mm wide;
vertical wings trapezoidal, the lower margin projected
and acute, ca. 2 x 4.3 mm. Stigma trilobed, the lobes
emergent, with a membranaceous, convex rostellum,
4 mm wide; lateral lobes subquadrate, rounded, ca.
1.2 x 1.2 mm. Anther versatile, articulate to the wide,
convex, clinandrium, ovoid-subquadrate, 3.5 x 3 mm.
Fruit short, thick, trigonous in cross section, 10-16 cm
long, 1-1.4 cm thick; fragrant when ripe, aroma similar
to common vanilla, but weaker. Fig. 4, 16A.

DISTRIBUTION: Known from Mexico (Oaxaca and
Chiapas), Guatemala, Belize and Honduras.
ECOLOGY: Hemiepiphyte in rain forest on soils with
variable drainage in areas with 2500-4000 mm of
rainfall, at 150-350 m altitude. It is the most common
Vanilla in many areas with tropical rainforest of the
Selva Lacandona (Chiapas) and the Peten (Guatemala).
It is the only penicillate Vanilla in N Central America
blooming during the rainy season, in July and August.

Vanilla cribbiana is a member of the V hostmanii
group. Like the other members of this complex, it has a
slightly trilobed to subentire lip with a disc adorned by
some thickened veins. It is different from V hostmanii
Lindl. from Amazonia, probably its closest relative,
by its smaller leaves, shorter inflorescence with fewer
flowers, more defined and subquadrate midlobe of
the lip, and only ca. 5 thickened, papillose veins in
the center of lip apex, not ca. 10 and spreading to
the lateral lobes. From Vanilla dressleri Soto Arenas
from Costa Rica to Colombia, it is distinguished by
its denser raceme, more defined apical lobe of the
lip, less conspicuous thickened veins on the disc,
papillose-granulose outer surface of the sepals and
broader, thicker leaves. Vanilla ruiziana Klotzch
(synonym V weberbaueriana Kraenzl.) from Peru and
Bolivia has a larger, ovate midlobe of the lip which
bears 1-3 elevated, axial keels near the apex and the
tepals are acute and the sepals neatly calyptrate. The
flowering period is July-August, at the beginning of
the rainy season, anf it is also distinct. Vegetatively
it is easily confused with V planifolia but the leaves
of V cribbiana are usually more elliptic, basally more
attenuate, and acuminate at the apex. However, the
vegetative differences are difficult to appreciate, and
both species are easily confused if flowers are not

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


available, and especially in sterile shoots grown in
shade. Therefore, Vanilla cribbiana has been confused
with V planifolia, and also with V pompona and even
V inodora, in the herbaria and in the plantations.
The fruits of this species are fragrant, and probably
of some potential commercial interest, but the vine is
not a strong grower, nor does it flower freely, at least
in the plantations of northern Oaxaca, where it was
introduced when confused with V planifolia (Perez
Mesa, pers. com.). It is pollinated by an unidentified
Eulaema bee. Beza (com. pers.) reports that the fruits
of this species have been cured in Guatemala.

OTHER RECORDS: MEXICO: OAXACA: cultivated by
J. Perez Mesa at Instituto Tecnol6gico Agropecuario #
3, San Bartolo Tuxtepec, from plantation in Jaltepec de
Condoyoc, near Ma. Lombardo, 24-IV-1997, J. Perez sub
M. Soto 8504AMO(in spirit)! CHIAPAS: Mpio. Ocosingo,
Estaci6n de Biologia Chajul, ca. 200 m s.n.m., selva alta
perennifolia despu6s de la sabana II, 19 junio 1996, M
Soto 7941 & R.Solano *AMO(sterile)! Chajul, Camino a
Arroyo Miranda. Fruto trigono, ca. 16 cm de largo, 14
mm de grosor, amarillo, rugoso, inflorescencia con ca. 21
flores, 22 junio 1996, S. Sinaca sub M. Soto 7953A MEXU!
Mpio.Ocosingo, Estaci6n de Biologia de Chajul, camino
a la Sabana I, selva alta perennifolia en loma, ca. 230 m
s.n.m. 13-IV-1997, M. Soto 8387 *AMO(sterile)! Mpio
Ocosingo, 1.5-2 km al SW de la Colonia Benito Juarez
Miramar, sobre la desviaci6n a Tierra y Libertad. Acahual
de 18 afios derivado de selva alta perennifolia, con Bursera
simaruba, Vochysia hondurensis, 360 m s.n.m., 20N,
9112'W, Hierba epifita; fr. inmaduro, 20 agosto 1993, A.
Reyes-Garcia y M. Sousa 2029 MEXU(584270, 584313)!
Mpio. Ocosingo, entire Bonampak y el Rio Lacanji, selva
alta perennifolia con Dialium guianense, Brosimun spp.,
Ficus glabrata y Terminalia amazonia, 300-350 m s.n.m.,
abril 2000, M. Soto 9617, S. Maldonado, P. Schliftter, L.
L6pez AMO(sterile)! [cf, fruit apparently sulcate] a 2 km
del Crucero Corozal, camino Palenque-Boca de Lacantuim,
Mpio. Ocosingo, 180 m s.n.m., hierba epifita, fruto verde,
selva alta subperennifolia, 13 feb 1985, E. Martinez 10299
MEXU! Mpio de Ocosingo, carretera Palenque-Marquez
de Comillas, Crucero San Javier; selva alta-mediana
subperennifolia, inundable, perturbada con ( /,, ..! ,'(1/ .
Vochysia y muchas epifitas, 16-IV-1997, M. Soto 8438-8440
AMO(sterile)! GUATEMALA: PETEN: Fleshy epiphytic
vine. Canchacan, in high rain forest of southeastern
Pet6n. July 14 1959. C.L. Lundell 16457 *LL(x2, buds)!
MO(3832548)! "Vianilla", vine, fruits black, fragrant,
Dolores, in low forest of pinal about 800 m south of
the village on the Machaquila Road, May 18, 1961, E.
Contreras 2333 MEXU(511605)! NY! LL(fruit)! LL(buds)!








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


"Vainilla" Fleshy vine, fruit green; Dolores, on Rio Mopan
trail, in high forest, October 17, 1961, E. Contreras 3063
LL(fruit)! BELIZE: STANN CREEK: vanillala. Vine; fls.,
yellow. In high ridge on hill top. Middlesex, 2 July 1939.
P.H. Gentle 2894 *AMES(58082)! K! LL! NY! TOLEDO:
Jimmy cut, Alt. 40 m, vive, hanging from tree, no flowering
or fruiting, stiff, thick leaves, 1973, C. Whitefoord 1816
BM! HONDURAS: ATLANTIDA: Near Tela. Guaymas.
Clambering over tree. March 17, 1923 0. Ames II 211
AMES(36945, fruit)!

4. Vanilla dressleri Soto Arenas, sp. nov.

TYPE: PANAMA. COLON: End of Pipeline road
onRioAgua Salud, 9-10 miN of Gamboa. Tropical
wet forest. Elev. 20-50 m. Vine. Flowers greenish-
yellow, lower lip white. Column yellow. Sweet
smelling, 15 April 1982, S. Knapp 4621 & J. Mallet
(holo. MO 3032052!).
Species Vanillae hostmanii similis sed floribus
longioribus, labello longiore subintegro anguste
obtuso differt.

Hemiepiphytic vine, leafy. Stems flexuoue,
apparently terete, grooved, 3-4 mm thick (in dried
specimens); intemodes 7.5-11.5 cm long. Aerial, free
roots terete, 2 mm thick; ,ll. i i!i, aerial roots strongly
flattened, up to 4 mm wide. Leaves petiolate, the
petiole canaliculate, 11-18 mm long, ca. 5 mm wide;
the blade oblanceolate, obovate to narrowly elliptic,
much larger than the intemodes, sometimes slightly
oblique, basally rounded to attenuate, apex acuminate-
cuspidate, fleshy, chartaceous (in dried specimens),
mesophytic, conspicuously pendant in living condition,
10.5-21.5 x 3.0-6.5 cm. Inflorescence a 3-10 flowered
raceme, rather lax, contiguous bracts distant up to 11
mm; peduncle 15-35 mm long, rachis 15-43 mm long,
at least 4 mm thick. Floral bracts patent, spreading,
concave, widely ovate to elliptic, obtuse, up to 14 x
8 mm. Flowers successive, 1 open at a time, big and
showy, tepals white to greenish-yellow, lip orange-
yellow in the inner surface, fading to white, with
brownish veins, column yellow; fragrance clove-like.
Ovary subterete, straight, thickened towards the apex,
bisulcate, the grooves twisted, very inconspicuously
and minutely papillose, 42 mm long, 4-5 mm thick;
sometimes slightly calyculate. Dorsal sepal linear to
narrowly elliptic, apex obtuse, rounded, subcalyptrate,
thickened, with 2-3 warts at the apex of the outer


surface; base narrowed, canaliculate; conspicuously
keeled on the outer surface, blade apparently
somewhat concave, ca. 13-veined, stiff, fleshy, 72 x
11.5 mm. Lateral sepals obliquely linear to narrowly
elliptic, apex obtuse, rounded, calyptrate, thickened,
inconspicuously warty at the apex on the outer surface,
base narrowed, concave, blade ca. 11-veined; obscurely
keeled on the outer surface, stiff, fleshy, 67 x 12 mm.
Petals linear to narrowly elliptic, slightly oblique,
apex widely rounded, base narrowed, basally concave,
with an elevated, axial, flat keel on the outer surface,
the keel ending in an acuminate, laterally compressed
process, ca. 2 mm long, apex strongly grooved, ca.
10 veined; 68 x 13.5 mm. Lip attached to the column
along the margins of the basal half(ca. 28 mm), tubular,
apparently concave, axially grooved on the abaxial
surface; when spread out 65 x 39 mm; unguiculate,
the claw canaliculate, lanuginose becoming pubescent
towards the blade, 15 mm long, 5 mm wide near the
blade; blade subrhombic, margin undulate, pleated,
apex obtuse-rounded, very slightly notched, ca.
24-veined; the ca. 11 veins of the apical part thickened
and rather warty, with the warts flattened, the central
ones rather elevated; the region corresponding to the
lateral lobes ca. 36 x 15 mm; the wanting midlobe 10 x
15 mm; penicillate callus at 28 mm from the lip base,
made up by ca. 12, obtriangular, basally long attenuate,
fimbriate scales, 5 x 2.5 mm. Column elongate, rather
straight, ca. 34 mm long, 3 mm wide; densely pilose on
apical 2/3 of the ventral surface, below the stigma; apex
dilated, vertical wings oblong, lower apex acute, ca. 1.5
x 3 mm. Stigma trilobed, the lobes emergent, rostellum
strongly convex, lateral ones apparently very narrow.
Fig. 5, 16B.

DISTRIBUTION: Known from Costa Rica, Panama and
Colombia.
ECOLOGY: In lowland to submontane (20-1000 m),
wet forest. Flowering from March to early June. The
species seem to be fairly common, at least in the Rio
Savagre area of W Costa Rica. The clove-scented
flowers suggest a male euglossine bee as pollinator.
ETYMOLOGY: Dedicated to Dr. Robert L. Dressier,
authority in orchid evolutionary biology and in orchid
floristics of Central America.

This large-flowered species is a member of the V
hostmanii group, characterized by its large flowers with
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.









LANKESTERIANA


10cm


4cm


S-i






















2 cm


FIGURE 5. Vanilla dressleri Soto Arenas. Based on G. McPherson 9196. Drawing by M. Lopez.

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


scarcely trilobed lips, granulose ovaries and sepals, and
the distal veins of the disc thickened. Among the group,
it is distinguished by the rather lax inflorescence, with
the flowers distant (ca. 1 cm), patent bracts, subentire
lip, with the blade subrhombic in outline, and very
scarcely pappilose sepals; the other members of the
group, V cribbiana, V hostmanii and V ruiziana have
conspicuously granulose sepals. From V cribbiana it is
additionally distinguished by the narrower, thinner leaves
which are more long acuminate-cuspidate, more lax
inflorescence, less defined apical lobe, veins of the disc
more conspicuously thickened, and flowering time mostly
in March-May, not in July-August. Vanilla hostmanii
from Amazonia has very long inflorescences that bear
many more flowers (e.g. 40-50) and the leaves are larger,
thicker, but not as strongly acuminate-cuspidate. Vanilla
ruiziana, from Peru and Bolivia has larger, broader, less
acuminate-cuspidate, and thicker leaves, but is similar to
V dressleri in having rather elevated axial keels of the lip.
Vanilla gardneri from Brazil has a subacute lip apex and
oblong leaves, not cuspidate.
Central American specimens of V dressleri have
been previously identified as V planifolia and V
insignis, distantly related species, easily separated by
the distinctly trilobed lips with well developed retrorse
warts.

OTHER RECORDS: PANAMA: Flowers yellow green;
lip yellow with green patch at apex. Collector, N.H.
Williams, Rio Iguanita, fl in cult, 14 April 1978, SEL
56-76-11[accession num.], *A. Pridgeon s.n. SEL(019244;
the leaves; flowers may belong to V phaeantha)! COLON:
Santa Rita Ridge, southeast of Colon, c. 920'N, 7945'W.
Along ridge road, 10-12 miles from Transisthmian
Highway, c. 550 m. Vine; perianth white, the lip orange-
yellow within basally; flower clove-scented. 21 May 1986.
G. McPherson 9196, holo. MO(3432756)! COCLE: Huge
plant purchased in the El Valle market. Flowers pale yellow-
green. 6 March 1976. R.L. Dressier, C. Luer J. Luer & P.
Taylor 768 *SEL(009187)! COSTA RICA: ALAJUELA:
Reserva Biol6gica Monteverde, Rio Pefias Blancas,
1019'N, 8443'W, 900 m. Epifita. Flores blancas con
amarillo. 7 June 1988. W. Haber 8471 & E. Cruz CR INB!
MO(3714711)! Reserva Monteverde, Rio Pefias Blancas,
1020'N 8443'W, 820 m. Bejuco. Flores blancas. 10 June
1987. W. Haber 7423 & E. Cruz INB! Upala, San Jose, Villa
Nueva, 11 km al noreste de San Jose. 1059'N, 8507'W,
40 m. Terrestre, trepadora, escandente. Flores en bot6n
verde amarillento. 18 April 1988, *G. Herrera 1840 CR
MO(3864177)! CARTAGO: 12 km S of Turrialba by air, 4


km SE ofPejibaye along Rio Gato. Disturbed primary forest
along river. Vine. 700 m alt. 9048'N, 8342'W. Vine.16-17
April 1983, *R. Liesner 14435 MO(3102177)! SAN JOSE:
Sepals and petals greenish yellow. Lip cream-colored with
greenish-brown raised lines. Callus in middle of lip raised
and with rows of attached hairs or brushes. Vine. Parque
Nacional Braulio Carrillo; Carrillo station, 300-600 m, 31
May-5 June 1980, *C. Todzia 1291 LL! HEREDIA: Sep
& pet interiorly creamy, faintly greenish, lip embraces
column loosly, interior of lip and base yellow, apical 3rd
fading to white, leaves suculent, stems single, not branched.
Sarapiqui: (April) .... The lip has been pressed .., do not
reveal the actual appearance of the flowers at complete
anthesis, and they have no perceptible odour at 10:30 a.m.
Flowers in axillary, stubby branches of from 5 to 8 flowers.
Leaves drooping apparently from a rather slender stem,
April 1959, *C.H. Lankester 1746 SEL (011365)!. LIMON:
Reserva Biol6gica Hitoy Cerere, Valle de la Estrella. Bosque
primario, bosque secundario. 700 m Epifita, flores amarillas
con blanco, 4 December 1990, G. Carballo 325 INB!
PUNTARENAS: Canton de Golfito. P.N. Corcovado. Valle
de Coto Colorado. Estaci6n Esquinas: Secci6n Esquinas.
0846'00"N 8315'00"W, 100 m. Bejuco, frutos verdes.
17 July 1993, M. Segura 121, F Quesada & R. Aguilar INB!
COLOMBIA: VALLE: Rio Calima (region del Choc6);
La Trojita, 5-50 m alt., Bejuco trepador; flores amarillo
blanquecinas, 19 febr.-10 mar. 1944, *J. Cuatrecasas 16550
AMES(71363)! Cordillera Occidental; vertiente occidental:
Hoya del rio Anchicaya, lado derecho, bajando a La plant,
bosques, 200-350 m alt.. Bejuco herbaceo, crasiuisculo;
tepalos ocraceo blanquecinos; labelo amarillo claro. 27
sept. 1943, *J. Cuatrecasas 15220 AMES(71364,71365)!
CHOCO: Parque Nacional de Utria: Colecciones realizadas
en la serrania ubicada al este de la ensenada de Utria, en un
recorrido oeste-este, entrando por la casa de la Seiora Ana
Elida (Mesica) siguiendo la trocha que conduce al acueducto
cruzando por la quebrada Aguada. 620'N, 7720'W, 0-100
m. Epifita; flores amarillas, 30 Mayo 1990, *F Garcia C.
& E.D. Agualimpia 325 MO(3878341)! CHOCO(not seen).
WITHOUT LOCALITY: No collection data: Climbing
vine; sepals and petals pale yellowish-green; petals with flat
ridge along center; lip white with crenulate, entire margins,
inner throat with yellow and brown lines; column yellow;
sweet, spicy fragrance. 6 May 1992, *S. W Ingram 1421
SEL(066924)! same data *S. W. Ingram 1422 SEL(066925)!

5. Vanilla hartii Rolfe, Bull. Misc. Inform. Kew: 133.
1899.

TYPE: Trinidad, Cabasterre Arima, Hart 6355,
holo. K!, iso. AMES(67785)!
V leprieurii R. Port., Bull. Soc. Bot. France, 98:
94. 1951.

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LANKESTERIANA


Type: "Guyane Frangaise: Cayenne, dans les
forts humides, Leprieur (s.n.), en 1846" (not
seen, drawing!)

COMMON NAMES: "Vanilla", vanillala.

Hemiepiphytic vine, branching, leafy, relatively
weak, up to 3 m high; perhaps up to 8-10 m long. Stems
terete and smooth, green, 3-6 mm thick; intemodes 7.0-
12.4 cm long. Aerial, free roots terete, pale brownish,
8-12 mm long, 1-2 mm thick; attaching roots up
to 6 cm long, 2 mm wide, conspicuously flattened.
Leaves usually slightly shorter than the intemodes;
conspicuously petiolate, the petiole straight or twisted,
canaliculate, with a clear junction line with the blade, ca.
10-14 mm long, 1.5-3 mm broad (not flattened); blades
elliptic, acuminate, rounded at base, coriaceous, green,
9-11 nerved (and a similar number of smaller veins
intercalated); 6.0-11.5 x 1.4-3.5 cm. Inflorescence a
short raceme with about 5-8 flowers, 13-30 mm long,
3-3 mm thick; peduncle of 4-7 intemodes; rachis 5-15
mm long. Bracts sessile, variable, triangular-ovate,
acuminate to obtuse (the upper ones), concave (the upper
ones) to cymbiform, not very stiff, progressively smaller,
3.5-9 x 2.5-5 mm. Flowers successive, 1-2 open at a
time, segments spreading, ephemeral, but at least some
of them remaining for more than one day, relatively
small, tepals white, the sepals tinged with green, lip white
with grayish or brown faint lines on the throat and an
inconspicuous green cushion at lip apex; ca. 3.8 cm high;
fragrance imperceptible. Ovary terete, green, smooth,
2.7 cm long, 1.2 mm thick. Sepals fused at base ca. 2
mm, subunguiculate and canaliculate at base, smooth.
Dorsal sepal oblanceolate, apex acute, rounded, slightly
thickened, flat to slightly concave, the apex slightly
incurved, 12-veined, 39-49 x 6-8 mm. Lateral sepals
obliquely oblanceolate, the lower margin more arcuate,
apex obliquely subacute, rounded, thickened, slightly
subcalyptrate, and obscurely warty, 12-veined, 39-46
x 7-8.6 mm. Petals obliquely oblanceolate, somewhat
arcuate, the lower margin more straight, long attenuate
at base, apex subacute to widely rounded, ca. 12-veined,
smooth, slightly concave; with an axial, flattened keel on
the abaxial surface, ending in a cylindric, acute, attached
process, 39-50 x 6-8 mm. Lip attached to the column
margins ca. 24-26 mm, long tubular, trumpet-shaped,
basally gibbous, with an abaxial longitudinal groove;
when spread out 37-43 x 14-25 mm; the claw slightly
LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


sigmoid, ca. 16 mm long, minutely papillose-pubescent;
blade cymbiform, ca. 7 mm depth, subentire or entire,
constricted near the apex and appearing slightly trilobed,
the apex deflexed, obovate-flabellate, ca. 21-veined,
basal half with inconspicuous rows of papillae up to
the penicillate callus, distal part with thickened veins,
inconspicuously papillose, the axial ones more prominent
and forming an apical inconspicuous cushion, margins
denticulate-undulate, plicate, apex truncate; penicillate
callus at 26-30 mm from the lip base; ca. 4 x 3 mm,
built up by ca. 7 flabellate-praemorse, retrorse, lacerate-
laciniate scales. Column very elongate and slender,
semicylindric, 32-33 x 2-2.5 mm, apex dilated ca. 3-4
mm wide; slightly pubescent on the flat ventral surface,
below the stigma, vertical wings flabelate-bilobed, with
granulouse inclusions, ca. 3 x 1.5 mm, filament broad,
thick, ca. 1 mm long, 2 mm wide. Anther versatile,
obovoid, ca. 3 x 2 mm. Stigma trilobed, the lobes
emergent; with a flabellate rostellum, with the lateral
margins reflexed, the margin denticulate, ca. 3.5 mm
broad; lateral lobes emergent, ovate-quadrate, tongue-
shaped, ca. 1 x 1 mm. Fruit linear-fusiform, elongate,
cylindric, not fleshy, slightly calculated, dehiscent
along 2 lines; fragrant, characteristic vanilla aroma, very
sweet; 92-140 x 4-5 mm. Fig. 6, 16C.

DISTRIBUTION: Mexico (Chiapas, perhaps reaching
eastern Oaxaca), Belize, Guatemala, Honduras,
Nicaragua, Costa Rica, Panama, Trinidad, Guyana,
French Guiana, and probably Brazil.
ECOLOGY: Rare in wet forests at low altitudes, usually
growing on understory treelets, in deep shade. Flowering
time January to April. The Mexican population of V
hartii is sympatric with V cribbiana, V insignis, V
odorata, V planifolia, and V inodora, but V hartii is
confined to the top of small hills with shorter forests and
open understory with tree ferns and large sedges; the
other vanillas are absent from this habitat although they
may be found a few meters away. Female Euglossa bees
have been observed visiting the flowers, but we do not
know if they are effective pollinators. Vanilla hartii is a
shy flower-producer; we have visited over many years
the Mexican population during the flowering season,
and we have found open flowers only twice, in the
spring of 2000, and then in spring of 2002.

This species was first cited from Central America
by Correll (1965). The Central American material of V









SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


2cm


2 cm


4mm


1cm

5mm






I I














4 cm


FIGURE 6. Vanilla hartii Rolfe, based on M. Soto 9729-9731. Drawing by M. Lopez.


10 cm


IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








LANKESTERIANA


hartii has slightly larger flowers (e.g., lip is 37-43 vs.
32 mm long) and the lip apex was described originally
as acute to subobtuse, not obtuse to widely rounded.
However, examination of flower of the type and
pictures from Trinidad and Mexico show that they are
indistinguishable, and that the floral differences may
due to the herborization process. Three collections
of V hartii from the Pacific slope of Costa Rica are
vegetatively stouter than Mexican and Trinidadian
plants, and the flowers are also the largest examined.
We do not know if these differences may indicate that
they should be regarded as a distinct taxon; although
molecular data suggest that they are closely related
(see ITS analysis).
Vanilla leprieurii is here considered as a synonym
of V hartii. Porteres (1954) stressed that his V leprieurii
was distinct because the nervation in the center of the
lip was denser, with the zone before the nervation
thicker, more imbricate scales in the penicillate callus,
and the apex of the lip more emarginate, apiculate and
pleated. Furthermore, the lip looks constricted, almost
trilobed in the published drawing. All these features
are variable in the material examined and again, they
seem to be the result of the way in which the flowers
were pressed.
Vanilla hartii is a relatively small vine similar
to the V planifolia group. It is somewhat similar to
V odorata but has shorter racemes, much smaller
flowers, a gibbous lip base, subentire lip, denticulate-
undulate (not lacerate-fimbriate) lip margin, a cushion-
like thickening at the lip apex without retrorse papillae
(vs. the 3-4 longitudinal rows of apical, retrorse, big
papillae) and broader and shorter elliptic leaves (vs.
long triangular-ensiform). The elliptic leaves, shorter
than the intemodes, and the slender stems make it
vegetatively similar to V bicolor from the Caribbean
and Guyanas. However, V bicolor has acute to
subacuminate lip, larger, tan-colored flowers with
yellow lip, and stouter, more elongate inflorescence.
Most Nicaraguan and several of the Central
American specimens previously assigned to V
planifolia are actually V hartii. Also the reports of wild
V planifolia from Rio Palenque Center in Ecuador are
based apparently in V hartii (P Lubinsky, com. pers.).
Vanilla hartii is morphologically similar to
V planifolia, and even confused with it in many
treatments. However, all the molecular data place it in

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


a rather basal position among the American penicillate
vanillas. Vanilla hartii tolerates damper, more shady
conditions, but the fragrant fruits are much smaller,
and it is not a free-flowering plant.
OTHER RECORDS: MEXICO: CHIAPAS: Estaci6n de
Biologia de Chajul, selva alta perennifolia, selva de loma
con Calophyllum y sotobosque de cyperaceas altas y
helechos arborescentes, sobre el camino a Arroyo Miranda,
16 07'35"N, 90 54'35"W; 200 m s.n.m., 12-IV-1997,
M. Soto 8347 *AMO(sterile)! 8350 *AMO(fruit)! same
data 14 abril 2000, M. Soto *9727(x2), *9729(x2), S.
Maldonado, L. L6pez y P Schlfitter AMO!; same data:
flor blanca, t6palos algo mas verdosos, especialmente en
el engrosamiento apical, con lines inconspicuas, caf6s,
tenues en la garganta, sin fragancia aparente. Rondada por
Euglossa hembra, pero no capturada ni vista polinizando la
flor. Fruto maduro con olor caracteristico a vanilla, muy
dulce M. Soto 9730, S. Maldonado, L. L6pez y P Schlfitter
*AMO! Sobre la Vereda La Granja, Estaci6n de Biologia
de Chajul, selva alta perennifolia en zona de loma con
muchos arroyos, algo perturbada, ca. 16 07'N, 90 54'W,
200 m s.n.m. escasa, en floraci6n, 15 de abril de 2000,
M. Soto 9731 y P Schlfitter *AMO! BELIZE: TOLEDO:
"Vianilla" Vine, flowers white, in broken Cohune Ridge,
between Orange Point and Moho River, April 28 1952, P.H.
Gentle 7673 MEXU(511492)! Southern Maya Mountains,
Bladen Nature Reserve, mountain, 1.7 airline north of
Ex Xux archeological site, 16 31'05"N, 88 54'11"W,
500-600 m, vine, flower white, in tree fall gap, 24 May
1996, G. Davidse 36251 BM! COROZAL: [cf.] "vanilla",
vine, P.H. Gentle 328 F(713628)! GUATEMALA:
IZABAL: Leaves subcoriaceous, dark dull green above,
slightly paler dull green below. Stem terete, dull green.
Petals and sepals pale greenish-white. Lip white. Leaves
somewhat narrower than in typical V fragrans. Swamps
of Salomon Creek, 1/2-1 mi. south of Bananera, alt. 50 m.
April 6, 1940. J.A. \i, ....... 38944 *F(1043051)! [cf.]
Quebradas, 19-22, May 1919, H. Pittier 8589A NY(sterile)!
*US(1013493; sterile)! HONDURAS: ATLANTIDA: [cf],
sterile Lancetilla Valley, near Tela, altitude 20 to 600 m;
vanillala, creeping on tree in wet forest; frequent, Dec. 6,
1927-Mar. 20, 1928, PC. Standley 52824 *AMES(36946,
fruit)! F(582560)! *US(1407340; sterile)! NICARAGUA:
ZELAYA: Monkey Point; ca. 1135'N, 83 39'W, elev 0-20
m; beach and bluff near village; vine on understory tree,
flower pale green, 7 Apr 1981, W.D. Stevens, B.A. Krukoff
20021 *SEL(047700)! [cf] Ibo, drainage of Cano Sung
Sung, N of road between Puerto Cabezas and Rio Wawa;
approximately 14 9-11'N, 83 29-31'W, elev. less than 10
m; gallery forest and adjacent savanna. Vine on tree trunk,
sterile. WD. Stevens & B.A. Krukoff 10667 SEL(036668)!
[cf] Caho Zamora on Rio Rama; ca. 11 57'N, 84 16'W,








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


elev. ca. 10 m; gallery forest along canio, pasture land on
plain. Epiphytic vine, sterile. W.D. Stevens, B.A. Krukoff
8835 *SEL(054718)! without data [the attached flower;
leaf, comments and analytical drawing on envelope perhaps
belong to V planifolia] A.H. Heller s.n. *SEL(03849)!
COSTA RICA: PUNTARENAS: Hilly slopes west of Villa
Nueva and Rio Naranjo. Evergreen rainforest formations on
the seasonally dry Pacific slope. Elev. ca. 200 m; 928'N
8428' W. Vine growing in partial shade 1 m up on stump.
Greenish-white flowers. 10-12 Feb 1988, W. Burger, K.
Swagel & J. G6mez-Laurito 12251 F(2009056)! Canton de
Osa, Aguabuena. Cuenca superior de Quebrada Aguabuena.
842'40"N 8331'40"W, 200-400 m. Bejuco trepador
colgante. Inflorescencia de bracteas verdes. Flor blanco
verde, labelo blanco con mancha puntual, apical, verde,
column blanca, polinios amarillo blanco. 18 January
1991, G. Herrera 4846 INB! MO SAN JOSE: Rio Savegre,
aproximadamente 1 km antes de la entrada a la Finca La
Gavilana, cerca de los limits del Parque Nacional Tapanti,
selva lluviosa con Anacardium, ca. 200 n.s.n.m., trepadora,
con capsulas, relativamente muy robusta, E. Salas 1 &
M. Soto INB! HEREDIA: Sarapiqui, Chilamate. Finca El
Bejuco, S end of Cerros Sardinal (N of Rio Sarapiqui).
1027'N, 8404'W, 60 m. Vine climbing in understory tree
at edge of forest to 3-4 m (transplanted here from swampy
area of primary forest nearby). Sepals and wing petals
cream-white. Labellum pure white, gibbous at very base,
constricted 1/4-1/3 towards apex, gibbous again (more
prominently) in apical 3/4, slightly narrowed before rotate
margin. Labellum with grayish lines within. 24 January
1987. M. Grayanum 7998 & T Ray *MO(3593651)!
PANAMA: PANAMA:cf] Rio Tecumen. Moist forest;
herbaceous vine; scarce, January 3, 1924, P.C. '..*i.. .
29353 *AMES(31441; sterile)!

6. Vanilla helleri A.D.Hawkes, Phytologia 14(1): 34,
19-20. t. 13.1966.

TYPE: NICARAGUA: Dept. Chontales: 2 miles
south of La Libertad on the road to Sto. Tomas,
growing as a vine on a wild avocado (Persea sp.)
tree, alt. 1900 feet, April 1962, A.H. Heller 7946,
holo. AMES? (not located), iso., SEL(fragments
and h., iri,''

Hemiepiphytic vine, branching, leafy. Stems
flexuous, terete to slightly subquadrate, sulcate, dark
olive green, minutely and inconspicuously papillose-
rugose inliving condition; 5-10 mm thick; intemodesup
to 15 cm long, the apical stems conspicuously covered
with glaucous wax. Attaching, aerial roots flattened,
1-3 mm wide. Leaves subpetiolate, the petiole 9-12


mm long, the blade oblong-elliptic to oblanceolate
(perhaps also elliptic), fleshy, abruptly constricted
towards the acuminate apex (acumen 10-15 mm), 12-
15 x 2.5-4.5(-6) cm. Inflorescence a 12-20-flowered
raceme, candelabrum-shaped, 4-10 cm long; peduncle
1-2 cm long. Bracts triangular-ovate, acute to obtuse,
dark green, up to 9 x 8 mm. Flowers successive, 1
open at once, with partially spreading segments,
ephemeral, showy, medium-size, sepals whitish,
greenish at the apical half on the abaxial surface, petals
whitish, lip bright yellow with orange appendages on
the midlobe, column white; ca. 6 cm diameter. Ovary
slender, subterete, white, apex green, 39-50 mm
long, 3 mm thick. Dorsal sepal oblanceolate-elliptic,
subacute-subobtuse, slightly concave, fleshy, slightly
subcalyptrate at apex, 11-veined, 40-42 x 11-14 mm.
Lateral sepals obliquely elliptic, obtuse, slightly
concave, fleshy, subcalyptrate, 11-veined, 40-45 x
13-14 mm. Petals oblong-elliptic, oblique, subacute
to obtuse, rounded, 11-13-veined, dorsally keeled, 40-
47 x 11-13 mm. Lip attached to the column margins
up to the stigmatic region, forming an inflated throat,
cuneate, without a claw, the blade clearly trilobed,
when spread out 35-40 x 30-33 mm; lateral lobes
flabellate-obovate, rounded, margins entire to slightly
repand with branched veins, ca. 5 mm wide at apex;
midlobe subquadrate to transversely oblong, truncate,
covered with retrorse, complanate, appendages up to
3 mm long, along the veins; disc with inconspicuous
axial rows of minute warts, with abundant trichomes
on the sides of the basal half; penicillate callus at
20-23 mm from the base, almost continuous with the
axial warts and the apical appendages, a tuft of long
laciniate, scales, much divided, ca. 5 x 5 mm. Column
subclavate, rather short and thick, semicylindric, 28
x 4 mm, apex dilated ca. 5 mm wide; ventral surface
with fine glandular hairs below the stigma; vertical
wings subtriangular, subacute, filament broad, thick,
ca. 1 mm long, 2 mm wide. Stigma trilobed, with a
convex, flabellate midlobe, the margin entire, ca. 3.5
mm broad; lateral lobes emergent, quadrate, tongue-
shaped, ca. 1 x 1 mm. Anther versatile, transversely
ellipsoid, ca. 3 x 3 mm. Fig. 7.

DISTRIBUTION: Costa Rica and Nicaragua, and also
apparently in Oaxaca, Mexico. Our knowledge of
the distribution of V helleri is very fragmentary. It
is known from the type locality on the slopes of the

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2 cm
iI


2 cm


FIGURE 7. Vanilla helleri Hawkes. Based on G. Davidse & R. WK Pohl 1503. Drawing by M. Lopez and M. Soto.

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


10cm








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Cordillera Chontalefia, and on the Pacific slopes of S
Costa Rica, especially in the Osa Peninsula.
ECOLOGY: In rainforest, with a short, marked dry
season, at 200-570 m altitude; it flowers in March-
April.

This species has a lip with elongate, complanate
appendages, similar to that found in V insignis. The
outline of the lip, with a cuneate base lacking a claw is
very anomalous among American penicillate vanillas
and is reminiscent of some Old World species. In the
original description Hawkes stressed that this species
has no close allies in Central America. However,
morphological and molecular data suggest that it is
closely related to V insignis and V odorata, but the
flowers are much smaller, and both these species
have conspicuously long-clawed lip. Vanilla helleri,
described 30 years ago is one of the rarest vanilla in
the area; it is known only from a couple of flowering
herbarium specimens. Despite its rarity in the herbaria,
V helleri could be more common than usually thought.
Several specimens of it are growing at Finca La
Gavilana, Rio Savegre, Costa Rica. Additionally,
several non-flowering plants, that may be this species,
have been located around Sta. Maria Chimalapa, in
Oaxaca, Mexico. Most Mexican specimens have rather
elliptic, broader leaves than Costa Rican material, but
they share the sulcate stems, papillose stem surface,
abundant wax on stem apex, and the nucleotidic
sequences are very similar to those of Costa Rica.
Originally, the Mexican specimens were thought to be
a hybrid between V planifolia and V insignis, since V
helleri has vegetative traits intermediate between these
species. We hope to see their flowers some day.
OTHER RECORDS: COSTA RICA: PUNTARENAS: 2 km
SW of the intersection of the Rio Ceibo and Carretera
Interamericana; elev. 200 m. Large vine climbing in a tree.
18 March 1969, G. Davidse 1503 & R. W Pohl F(1731785)!
MEXICO: OAXACA: [cf.] Planta j6ven, silvestre, en
selva alta perennifolia con Sebastiania, Quercus oleides,
Terminalia, Acacia, Ampelocera, ca. km 42 del camino
Mezquite- Sta. Maria Chimalapa, 390 m s.n.m., 20 Marzo
1997, A. Cibridn 11 & M. Soto (not preserved). [cf.]
Tallos asperos al tacto y subpapilosos, longitudinalmente
sulcados; al sureste de Sta. Maria Chimalapa, vereda hacia
el Rio Milagro, paso Quetz Tug, 15/XI/98 a 6/XII/98, H.
Herndndez V7 sub M. Soto 8806 AMO(sterile, photo)! [cf.]
Hojas subcuneadas en la base, algo oblanceoladas y algo
oblicuas; 3 km al este de Sta. Maria Chimalapa, cerca de la


cascada Arroyo Sangre. 15/XI/98 a 6/XII/98 H. Herndndez
V3 sub M. Soto 8811 AMO(sterile, photo)! [cf.] Tallos muy
delgados, sulcados, asperos, algo glaucos, hojas elipticas,
flexibles y delgadas. Aproximadamente a 12 km al suroeste
de Sta. Maria Chimalapa, a la orilla del camino Sta. Maria-
El Mezquite, en la bajada de Zacatal, 15/XI/98 a 6/XII/98,
H. Herndndez V8 sub M. Soto 8817 AMO(sterile, photo)!
Tallos muy delgados, sulcados, asperos, hojas elipticas,
flexibles y delgadas. Aproximadamente 1 km al noreste de
Sta Maria Chimalapa, a 5 m de la vereda de Paso Lagarto.
H. Herndndez V2 sub M. Soto 8818 AMO (sterile, photo,
ITS sequence)!
REFERENCES: Hamer, Orch. Nicaragua, Ic. P1. Trop. pl.
1192. 1984; Hamer, Selbyana 11. Orch. Center. A. p.
847. 1990.

7. Vanilla inodora Schiede, Linnaea 4(4): 574-575.
1829.

TYPE (?): "Baynilla de puerco Misantlensium";
Mexico, Misantla. Schiede 1044, K; "fruto inodori,
Vanilla inodora nob. interin Vaynilla de puerco,
Misantlensis .... Misantla, Mart 29 Schiede & BM!
W(s.n.; x2)!
Vanillapfaviana Rchb.f., Gard. Chron. 2, 20. 230.
1883.
Holotype: "Mexico" actually from Chiriqui,
Costa Rica, R. Pfau [269] W(19347)!
Vanilla preussii Kmzl., Notizbl. Bot. Gart. Berlin-
Dahlem. 7: 320. 1919.
Type: Bei der Planzung El Baul im Buschwald,
Heisst bei den Eingeborenen "Vainilla
silvestre" (Preuss n. 1445), not seen.

COMMON NAMES: "Vainilla de puerco" (Ver.); vanillala,
vanillaa cerro amarillo" (Oaxaca).

Hemiepiphytic herb, vigorous,up to 12 m high.
Stems terete, ca. 4-5 mm thick; intemodes ca. 7.5
long. Leaves usually hanging and with bending
blades; petiolate, the petiole ca. 1-2 cm long, twisted;
blades elliptic to broadly elliptic, abruptly acuminate,
thin, membranaceous when dry; 12-21 x 4.5-7 cm.
Inflorescence similar to the vegetative shoots; ca.
30 cm long, bearing 3-5 flowers; the raceme bears
progressively smaller foliaceous bracts. Bracts
similar to the leaves, although smaller or much
smaller, 3.3-12 x 1.0-5.2 cm; the intemodes 3.5-7
cm long, ca. 2 mm thick. Flowers resupinate, with
spreading segments, very showy, tepals apple-green or
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312 LANKESTERIANA


1cm


FIGURE 8. Vanilla inodora Schiede, based on M. Hernandez s.n. sub M. Soto 7864. Drawing by R. Jimenez.

LANKESTERIANA9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


yellowish green, shiny, lip white with yellow-orange
tinge in the throat; ca. 4.5-5 cm diameter; fragrance
sweet strong or faint medicinal, to weak solanaceous
alcaloids; flowers lasting 2-3 days. Ovary straight to
slightly arcuate, terete, smooth, 3-sulcate, the grooves
twisted, 47-55 (48.671.25) mm long, 3-4 (3.330.47)
mm thick. Tepals contorted, with very undulate,
somewhat revolute margins, and the apices recurved
or rolled backwards. Dorsal sepal erect, elliptic-
lanceolate, broadly obtuse to acute, ca. 11-veined,
44-50 (472.19) x 12-18 (13.42.8) mm. Lateral
sepals descending, elliptic-lanceolate, broadly obtuse
to acute, ca. 11-veined, 41-44 (42.41.02) x 13-17
(15_1.79) mm. Petals spreading, elliptic-lanceolate,
attenuate towards the obtuse apex, axially canaliculate
on the abaxial surface, ca. 9-veined, 43-47 (451.67) x
9-11 (101.26) mm. Lip attached to the column ca. 2.5
mm at base; arcuate, trilobed, the lateral lobes covering
the column, the midlobe spreading and deflexed; 27-36
(333.16) mm long, 29-34 (31.61.62) mm wide when
spread out; lateral lobes erect, semiobcordate, broadly
rounded, reduplicated and covering the distal half of
the column; 20-24.5 (22.72.23) x 9-13.5 (12.21.69)
mm; midlobe subcuadrate, spreading, deeply
emarginate, the margins undulate, 12-17 (14.81.94)
x 13-19.5 (15.82.38) mm; the midlobe and part of
the disc with a massive, fleshy, elevated, cushion-like
callus, 16-21 (181.79) mm long, 5-8 (6.81.16) mm
broad, ca. 3-5 mm high. Column strongly arcuate,
the apical part resting on the callus surface, 21-24
(22.61.16) mm long, 5-6.3 (5.760.56) mm wide at
the apex. Stigma a well-defined, transversely oblong-
panduriform cavity with thickened, yellow borders;
without a rostellum. Anther hinge-like, broad,
attached to the clinandrium, and forming together with
it, a couple of lateral auricles. Pollen soft, somewhat
sticky. Fruit cylindric with attenuate base and apex,
not aromatic, 18 cm long, ca. 8 mm thick. Fig. 8.
DISTRIBUTION: Mexico (Puebla, Veracruz, Jalisco,
Guerrero, Oaxaca, Chiapas), Belize, Guatemala,
Nicaragua (Heller & Hawkes, 1966), Costa Rica, and
Panama.
ECOLOGY: Vanilla inodora grows in wet forests from
150 to 1600 m altitude; it is the only Vanilla that
inhabits the cloud forests of the area, and is found in
the lowlands only in sites with more than 2500 mm
of rainfall. Flowering apparently without a defined


period. It is sometimes found in savanas with slow
drainage.
Vanilla growers believe that this species can be
self-pollinated; the large fruit set in some populations
supports this idea; however, other populations have a
fruit set as low as 2.5 %. The flowers remain in good
condition for 2-3 days, and are therefore long-lived
compared with other vanillas; often older and younger
flowers are at anthesis on the same raceme.
The flowers of this vanilla resemble those of some
species of Schomburgkia group of Laelia in the flower
structure; the tepals are contorted, strongly twisted,
with a varnished appearance and the column lies on the
lip surface; pollination must be carried out by large,
strong bees that try to enter to the throat separating
the lip from the column, as in Schomburgkia or
Barkeria; in the latter this work is done by carpenter
bees (Xylocopa), and they might be good candidates
to pollinate this species. We have observed carpenter
bees approaching V inodora in Chiapas, but they were
not been seen landing on the flower.
V inodora was collected by Schiede near Misantla;
the original collections are housed at K, BM and
W. None of the herbarium specimens bear flowers,
although probably some specimens have had fruits
(see Rolfe 1896); so its identity has been somewhat
obscure. The type specimen is evidently a member
of Portere's subsect. Membranacea, and as only a
single species of this group is found in Mexico, and it
is common in Veracruz, we discard the possibility of
other species, from elsewhere, to which the name V
inodora has been applied. There is no evidence that the
Mexican species is conspecific with the Haitian Vanilla
mexicana, as suggested by many authors, despite the
specific name of the latter. The large foliaceous bracts
of V inodora are larger than in other Membranaceous
species and are clearly visible in the type.
Vanilla inodora has been known in recent years
in the region as V pfaviana Rchb.f. In their Orchids
of Guatemala, Ames and Correll (1952) mentioned
that they had not examined Guatemalan material of
V inodora and that the species could be conspecific
with V pfaviana. It seems, however, that the sterile
specimens (or those very badly preserved) of this
species were always identified as V mexicana or V
inodora, while the material with flowers was thought
to be V pfaviana. Vanilla inodora and V mexicana

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


are similar, but the massive, elevated, fleshy callus of
V inodora is very different from the 3-keeled callus
found in V mexicana; furthermore, V mexicana can
have much larger leaves. It should be noticed that the
callus of V inodora may appear to be keeled in old,
withering flowers or in young buds.
The type of V preussii Kraenzl. was destroyed in
the Berlin bombing, but the description matches V
inodora very well. Furthermore, Schlechter, who must
have examined the type specimen identified Tiirckheim
II 1764 as V. preussii.
The allied species in Central America are V
martinezii, from Guatemala, that is easily distinguished
by the congested inflorescences, V costaricensis
with entire lip lacking a massive callus, and V
sarapiquensis, with short midlobe and a callus formed
by two high, massive confluent ridges at the distal part
of the lip. However, its closest relative may be the
Andean V methonica Rchb.f. & Warsz., with a similar,
emarginate lip, but having 3 thickened and somewhat
rugose keels instead the fleshy massive callus, and
much smaller inflorescence bracts.

OTHER RECORDS: MEXICO: JALISCO: Steep mountainsides
3-10 km generally east on the road to Mina Cuale, from the
junction 5 km northwest of El Tuito, Mpio. Cabo Corrientes;
pine-oak forest on decomposed granitic soils, with
Podocarpus, oaks, and other deciduous trees in rocky
stream valleys, elev. 850-1150 m. Seen once, in rocky
stream valley. Sterile; fleshy herbaceous vine climbing 10 m
or more in trees, 16-19 February 1975, R. McVaugh 26397
MEXU! Mpio. Cabo Corrientes, km 2.6 de la brecha hacia
a la izquierda que sale del km 9.8 del camino El Tuito-
Cuale, bosque de galeria de Hedyosmum, Inga, Podocarpus,
Magnolia, Clusia, en medio del bosque sabanoide de pinos
y encinos, sobre granitos intemperizados, 900 m s.n.m.,
20 22'29.4" y 105 15'21.9". Banco con suelo profundo
con vegetaci6n densa, similar a acahual de selva montana
lluviosa con Heliconias, Philodendrum tripartitum,
Lindenia y trepadoras. Hemiepifita, cerca 10 m de alto, con
muchos frutos, cerca 400 m de extension. Muy rara, s61o un
esp6cimen visto. 27/junio/1998 M. Soto 8626 y E. Huerta
*AMO(x6)! IBUG! GUERRERO: System of Teotepec,
near San Vicente, N.E. ofAtoyac, in mixed oak-pine forest
on tree trunks. 100 16'W, 17 17'N, 850 m, January 10,
1933. 0. Nagel sub E. Oestlund 1984 AMES (with sketch
by B. Ames; 41396)! *AMES(51815)! *MO(11411617;
sterile)!; San Vicente, north-east of Atoyac, towards Mt.
Peineta. In mixed forest on trees & shrubs, 100 16'W,
17 17'N, ca. 100-1000 m, 10 Jan 1933. 0. Nagel sub E.
Oestlund 1985 *AMES(41476, sterile)! BM(sterile)!

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


*MO(1145095; sterile)! PUEBLA: Vicinity of Puebla,
Venant des sierras (De la decoration florale a l'archevech6)
Dec. 8. 1907. B.G. Arsene 1682 *AMES(71359, sterile)!
*MO(843101, sterile)! NY! *US(1032010; fruit)!
VERACRUZ: Miradores, 4/42 Liebmann 297 K! Mirador,
Liebmann W(11762)! Hacienda de Java, 5/1841, Liebmann
295/296 W(13537, 13538, 11761)! Zacuapan, June 1919,
C.A. Purpus 8481 *AMES(71356)! NY! Region of
Zacuapan, near Rancho Viejo, climbing on tree trunks, ca.
700 m, 10 Jul 1935, C.A. Purpus sub E. Oestlund 4876
*AMES(sterile, 51843)! *US(1809800)! Near Zacuapan.
On trees and shrubs, 12 Feb 1932, 96 52' W, 19 12' N, ca.
900 m, O. Nagel sub E. Oestlund 2683 *AMES(sterile,
41475)! *US(1809491)! Region of Zacuapan, on tree
trunks, 8 Oct 1935, ca. 800 m, C.A. Purpus sub Oestlund
5045 *AMES(fruit, 51838)! Falda del Volcan de San Martin
Tuxtla, San Andres Tuxtla, 18 35', 95 09'W, alt. 800 m
s.n.m., selva alta perennifolia primaria, suelo negro arenoso,
buen drenaje, aluvial, calido himedo, lluvioso, escasa, floor
blanca, 29-10-1973, J.I. Calzada 01031 BM! CHAPA!
MEXU! El Mirador, Mpio. Totutla, bosque de encino en
canada, 1000 m, herbacea trepadora, flor verde, centro
blanco, fruto verde, escasa, 18-VI-1973, F Ventura 8417
*AMO(436)! CHAPA! MEXU! Lote 67, Estaci6n de
Biologia Tropical Los Tuxtlas, 95 04' y 95 09' O, 18 34' y
1836' N. Mpio. San Andres Tuxtla, borde selva alta
perennifolia, 300 m s.n.m, bejuco herbaceo, fruto verde-
grisaceo, vanillala, julio 14 de 1986, S. Sinaca C. 835
*AMO(7066)! OAXACA: Cerro Martin, cerca de Usila,
400 m selva alta perennifolia, predio de Sebastian Arista, 22
mayo 1993, M. Herndndez s.n. AMO(in spirit)! Usila, M.
Herndndez s.n. AMO(in spirit)! Usila, 2 mayo 1994, M.
Herndndez s.n. AMO(in spirit)! Cerro Martin, ca. Usila,
1992, M. Herndndez s.n. AMO(in spirit)! La Escalera,
Chinantla. "Vainilla Cerro Amarillo". abril-1990. M.
Herndndez Apolinar sub lM Soto 10707 AMO! 1844,
Mexico, Karwinski M W(19347, sketch)! Mpio. Sta. Ma.
Chimalapa: Arroyo Sangre ca. 2 km E de Sta. Ma.
Chimalapa, selva perturbada con Calophyllum, Tapirira,
Brosimum, etc., suelo cafe parduzco con much hojarasca,
250 m, 16 54'30" 94 40', 20 abril 1985, bejuco, flor blanca,
s6palos verde, aromatica, en canada, comfin, usos se pone
en aceite para el cabello, H. Herndndez 1125 CHAPA!
MEXU(564872)! Orquidea epifita, acahual derivado de
selva alta perennifolia, loc.: Cuaje, Mpio. Ixtlan de Juarez,
Dto. Ixtlan, Sierra Norte, 17/05/94, E. Torres 367
AMO(16696; young fruit)! TABASCO: Hierba epifita
como bejuco, flor con 5 t6palos semienriscados y una quilla
blanca, inflorescencia con bracteas foliares, asociada a
pukte en vegetaci6n riparia, Teapa, 31-05-1990, V Ramdn
& A Sol 309 MEXU [sterile]! CHIAPAS: "Local name:
vanillala, in wet forest, Libertad Acacoyagua, June 1 1948,
E. Matuda 17912 AMES(66534; young fruit)! F(1616949)!








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


MEXU(85052)! En Crucero Corozal, camino Palenque-
Boca Lacantuim, Mpio. Ocosingo, 180 m s.n.m., bejuco con
fruto, selva alta subperennifolia, 8 enero 1986, E. Martinez
15730 *LL! MEXU(436979)! MO(4272282)! *XAL!
Triunfo, Dec. 1936, E. Matuda 360 *US(1689405)! Triunfo-
Juarez, Escuintla, Dic. 1936,E. Matuda 369MEXU(85064)!
Arroyo Miranda, entire los 6 y 20 km del arroyo partiendo
del Rio Chajul, Mpio. Ocosingo, 150 m s.n.m., primaria,
orilla de arroyo, bejuco perenne, 8 m, flor blanca, nom. vul.
vanillala, trepador, 20-02-1985, G. Castillo et al. 4011
*XAL! 2 km al NW de Lacanja-Chansayab; Mpio. de
Ocosingo, 400 m, enredaderas; vainas verdes, lisas, negras
cuando maduras; numerosas semillas negras cubiertas con
una secreci6n pegajosa de las paredes internal de las vainas;
abundante en remanente de selva alta perennifoia
relativamente madura, abril 16, 1991, M Gonzdlez Espinosa
et al. 1413 CHIP! MEXU(563979)! Bonampak, Mun.
Ocosingo, 350 m s.n.m. selva alta perennifolia, suelo
arcilloso. "Vainilla de montafia". Enredadera muy escasa,
hasta de 7 m de largo. abril 1982. M. Soto 1020 AMO!
Estaci6n de Biologia Chajul, Mpio. Ocosingo, ca. 200 m
s.n.m., 1608' N, 90'53' O. Cerca del Puente Hamaca en la
vereda a la Sabana I. Loma con selva mediana-alta
perennifolia, en la base de la loma. Muy escasa. 23 junio
1996. M. Soto 7954-A yR. Solano *AMO! km 223.6 de la
carretera Palenque-Marqu6z de Comillas, ca. de Benem6rito
de las Americas, selva median subperennifolia con
Cocoloba y Roupala, 220 m s.n.m., 11-IV-1997, M. Soto
8342 et al. AMO(also in spirit)! Mpio de Ocosingo, ca. de
las ruinas Los S6tanos-El Zapote, Estaci6n de Biologia de
Chajul, selva alta perennifolia inundable con Bactris, 230
m, 1608'N, 90'53' O. Sobre Guarea en sombra densa, con
frutos. 12-IV-1997. M. Soto 8343 *AMO! Estaci6n de
Biologia de Chajul, junto al Rio Lacantun, vereda La
Granja, en la intersecci6n con la vereda a La Sabana, selva
alta perennifolia con Ficus glabrata en suelos profundos,
arenosos, probablemente inundables estacionalmente, floor
verde con labelo blanco, fragancia dulce, intense, 15 de
abril de 2000, M. Soto 9726 & P. Schliitter *AMO!
BELIZE: TOLEDO. Gracie Rock, Sibun River, 1 May
1935, P.H. Gentle 1672 *AMES(42338)! *LL! "Vainilla".
Climbing plant, on cohune tree, white flowers, in cohune
ridge, near river beyond Columbia. January 31, 1947, P.H.
Gentle 6152 *F(1599314)! G! *LL(x3)! NY! *US(2572753)!
"Vainilla". Climbing vine, white flowers, in acahual, near
Ocotal, Pine Ridge, 3 miles in trail from 7 Miles, Punta
Gorda-San Antonio Road, January 20, 1950, P. Gentle 6957
*LL(x3)! MEXU(511462)! *MO(3832518)! "Vianilla",
vine, in acahual, Feeders Road leading to Big Fall, April 13,
1950, P.H. Gentle 7023 *LL(fruit, x2)! "Vianilla", vine, in
cohune ridge, one mile from 7 Miles, San Antonio-Punta
Gorda Road, July 1, 1950, P.H. Gentle 7078 *LL(sterile
inflorescence, x 2)! "Vianilla", vine, in cohune ridge, near


Columbia, August 15, 1950, P.H. Gentle 7108 *LL(fruit)!
GUATEMALA: IZABAL: [cf, sterile] wet forest,.
"Vainilla". Creeping on tree trunk. Near Entre Rios, alt.
about 18m., April 30,1939,P.C. \i. ,. ., 72709F(991636)!
ESCUINTLA: [cf, sterile] El Zapote, in jungle, on tree of
Ficus, April 9, 1937, W.C. Muescher 12480 F(905455)!
SAN MARCOS: [sterile] vanillala, climbing, leaves fleshy
coriaceous, rich green above, practically same color but
slightly paler beneath, above Finca El Porvenir on "Todos
Santos Chiquitos", lower south facing slopes of Volcin
Tajumulco, alt. 1300-1500 m, March 7, 1940, J.A.
so, l..,....?l 37076 F( 1041850)! HUEHUETENANGO:
[cf] Epiphyte, alt., At 3000 ft. alt. Cerro Chiblac, between
San Rafael and Ixcan, Sierra de los Cuchumatanes, July 22,
1942, JA. Steyermark 49171 *AMES(63277, sterile)!
F(1495682)! SUCHITEPEQUEZ: [cf, sterile] Epiphyte on
tree on bark, leaves fleshy subcoriaceous, dull dark green
above, dull green beneath, stem pale green, in cafetal on
opposite side of Finca, southern lower slopes of Volcin
Zunil, vicinity of Finca Las Nubes, along Quebrada Chita,
east of Puebo Nuevo, alt. 500-800 m, Feb 2, 1940, J.A.
.\ i .... ,' 35412 F(1041244)! BAJA VERAPAZ: Wald in
Paujal, 1000 met April 1907, Bl. grin, Lippe weiss, H. von
Tilrckheim I 1764 *US(825825)! HONDURAS: COLON:
[cf] Vine, flower white, Guaranta, Wispernini Camp, 75-
100 ft., tropical rain forest, March 1938, C. vonHagen & W.
vonHagen 1352 F(942976)! NY! ATLANTIDA: [cf] banks
of the Salado River, above the village of Salado ... on the
mountain slopes and coastal plains, vicinity of La Ceiba,
July 10, 1938, TG. Yuncker 8335 *AMES(fruit, 50661)!
NY! COSTA RICA: ALAJUELA: Reserva Biol6gica
Monteverde. Rio Aguas Gatas, Laguna de Arenal, El
Castillo. 1026'N 8444'W, 600-1000 m. Epifita semi-liana
dentro del bosque. Flor con caliz verde, corola blanca en
forma de tubo. 11 August 1989, E. Bello 1146 INB! [cf,
without flowers] Reserva Forestal de San Ramon; camino
entire el Rio San Lorenzo y la estaci6n. 1012'53"N,
8436'28"W. Epifita trepadora, frutos inmaduros verdes. G.
Herrera Ch., I. Chac6n, D. Herndndez, A. Solis yH. G6mez
386 SEL(062313)! CARTAGO: vanilla, "Chiteria"
Chitaria, alt. 750 m, 15/4/36, F Solis F(833816)! [cf,
sterile] LIMON: Epiphyte, 7 km al SW de Bribris, 100-250
m, May 4 1983 [sterile], L.D. G6mez, R. Liesner, E.
Judziewicz 20437 MEXU! MO [cf, sterile] San Clemente,
Apr. 1920. Lankester (k331) K! SAN JOSE: Herbaceous
vine, attached by roots. Fls green with white labellum, in
forest. Vicinity of El General, alt. 1130 m, Feb. 1936, A.
Skutch 2592 AMES(*44231, *44232, *44233)! K!
*MO(1105371)! NY![ Cataratas de San Ramon, marzo de
1931, A.M. Brenes 13679 F(906350)! PUNTARENAS:
Canton de Osa. Fila costefia. Rio Piedras Blancas, cerca de
la casa. Cerro Anguciana. Fila Cruces, 0849'02"N,
8311'23" W, 900 m. Bejuco trepador. Caliz verde, labelo
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LANKESTERTANA


blanco. 10 December 1993. R. Aguilar 2736 INB!
PANAMA: BOCAS DEL TORO: [cf.] Vanilla, epiphyte;
flower pale green, Big Bight, Vicinity of Chiriqui Lagoon,
Oct., 27, 1940, H. von Wedel 2880 *AMES(61513, sterile)!
REFERENCES: Ames, Bot. Mus. Leafl. Harvard Univ.
4(3): 26-29.fig. p. 29. 1936.

8. Vanilla insignis Ames, Bot. Mus. Leafl. 2(8): 101-
102.fig.p. 103. 1934.

TYPE: REPUBLIC OF HONDURAS, Dpt.
Comayagua, Esquias, El Rio Funes. Epiphyte in
river-valley forest at 2,500 feet altitude. Sepals and
petals green, lip white at base. April 27, 1933.J.B.
Edwards 407, holo. AMES(40085)!; isotype
AMES(40086)!

COMMON NAMES: "Sisbic" (Maya), vanillala, vanillaa
cimarrona" (Veracruz).

Hemiepiphytic, branching, leafy vine, usually very
vigorous, up to 30 m high; up to a hundred meters
long. Stems flexuouse, subterete, with a conspicuous
groove along the entire internode, surface fairly
rugose-papillose, dark to olive green, ca. 6-9 mm
thick; intemodes, 10-17 cm long. Terrestrial roots
pubescent, brown-whitish, ca. 2-3 mm thick; aerial
free roots, terete, greenish gray, ca. 1 mm thick; aerial
attaching roots semicylindric, flattened in the surface
in contact with the substrate, ca. 3-4 mm wide. Leaves
subpetiolate, petiole canaliculate, up to 11 mm long,
4 mm wide;blade oblong-elliptic, abruptly acuminate
at the apex, rounded at base, coriaceous-fleshy,
rather stiff, 4.2-19.5 x 1.5-3.4 cm. Inflorescence a
6-12(21)-flowered raceme, candelabrum-shaped,
4.4-11 cm long; peduncle fleshy, subterete, slightly
compressed, 15-25 mm long, 6.5-8 mm thick; rachis
28-90 mm long, progressively slender towards the
apex. Bracts sessile, small, broadly ovate, obtuse,
concave, fleshy, progressively smaller towards the
apex, commonly less than 9 x 6 mm, rarely up to 10 x 10
mm. Flowers successive, 1-3 open at once, ephemeral
(from 8:00 to 14:00 hrs; starting to close at noon),
very showy, segments completely spreading, tepals
pale apple-green, shiny, lip cream-white with orange
to dull yellow appendages on the midlobe, extreme
apex green, penicillate callus cream, column white;
ca. 11-12.4 cm wide, 9-11 cm high; fragrance weak,
spicy. Ovary subterete, very slightly dorsiventrally

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


compressed, smooth, arcuate, 2-sulcate, the grooves
almost straight, 45-54 (48.753.27) mm long, 4.5-5.5
(50.41) mm thick. Dorsal sepal long oblanceolate,
apex acute, disatally rounded, slightly thickened,
subcalyptrate, base attenuate-subunguiculate, claw ca.
22 x 11 mm, basally canaliculate, concave at middle,
fairly incurved at apex, ca. 12-13-veined; 69-76.5
(73.16 2.56) x 10.5-14 mm (11.61.24). Lateral
sepals obliquely oblanceolate, the lower margin
more arcuate, apex subacute, thickened, apiculate,
subcalyptrate, smooth, the abaxial surface minutely
warty, especially at the minute apicule, base attenuate-
subunguiculate, basally canaliculate; slightly concave
towards the apex, margins somewhat involute, ca.
11-14 veined, 64-74 (69.673.19) x 12.5-13.5 (12.92
0.35) mm. Petals obliquely linear to oblanceolate,
somewhat arcuate, apex obtuse, distally rounded,
oblique, subcalyptrate, base long attenuate; concave,
with a very conspicuous, axial, elevated, flat keel at
the abaxial surface, ending in a subtrigonous, free,
adpressed process, ca. 2 mm long; ca. 12-14-veined,
68-74 (70.92.11) x 9-10 (9.760.39) mm. Lip
fused to the column along the margins of the basal
half (37-44 mm), long tubular, slightly concave,
fairly inflated near the base of the blade; the apex
conspicuously recurved-deflexed; axially grooved on
the abaxial surface, the groove deep; when spread out
65-73 (69.332.49) x 30-35 (32.6+1.62) mm; long
unguiculate, adaxial surface of claw hairy on the distal
half, the trichomes dense, more or less in rows; ca. 26
x 7 mm; the blade approximately obovate-flabellate
in outline, trilobed, ca. 30-veined, the veins branched
above the middle, thickened forming low, obscure, flat
keels near the base, disappearing near the basal third
of the blade; lateral lobes subelliptic, oblique, ca. 33
x 10 mm, margins long laciniate-fimbriate, especially
near the joint with the midlobe; cilia up to 7 mm long;
midlobe ovate-suborbicular to oblong, obtuse, the
margin undulate-crenulate to lacerate-dentate towards
the lateral lobes, 11-15 (131.41) x 12.5-14 (13.37
0.65) mm; penicillate callus at ca. 34 mm from
the base, 4 x 5 mm; made up by ca. 13 imbricated,
retrorse, flabellate, praemorse to densely lacerate-
laciniate scales, sometimes united to each other along
the lateral margins, scales almost without adpressed
processes on the surface, continuous towards the lip
apex with 7 rows of tubercles, near of the base of the









SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


3cm
I I


2cm[












































FIGURE 9. Vanilla insignisAmes. Based onM. Soto 7684. Drawing by M. Soto.

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


midlobe becoming enlarged to form conspicuous,
retrorse, subtriangular, thick, complanate, sometimes
conical, obtuse appendages, that cover almost entirely
the surface of the midlobe, distributed approximately
in 10 rows, the central appendages bigger, up to
4 mm long, 1 mm wide; extreme apex thickened,
mound-shaped. Column very elongate and slender,
semicylindric-trigonous, 49-52 (50.371.08) mm
long, 4 mm wide; ventral surface flat, with scarce,
minute trichomes below stigma; apex dilated, with
vertical flabellate, somewhat trilobed wings, ca. 1.5
x 2.3 mm; clinandrium galeate, ovate, convex, 2 x 2
mm. Stigma trilobed, the lobes emergent, midlobe
(rostellum) a transversely oblong, convex blade,
axially grooved, ca. 3 x 4 mm, covering and parallel
to the lateral lobes, these quadrate, rounded, convex,
slightly divergent from each other, ca. 1.5 x 1.5 mm.
Anther versatile, saddle-shaped, axially canaliculate,
3 mm wide, 1.8 mm thick, attached to the clinandrium
margin by a thick, approximately semiterete filament.
Pollen in monads, not forming a clear pollinarium
but an ill-defined, sticky mass. Fruit short, thick,
subclaviform-semifusiform, rounded, swollen towards
the apex, green turning yellow when ripe, dehiscent
along a single line, fragrant (less than V planifolia),
like a blend of common vanilla with coconut; ca. 7-14
cm long, 0.7-1.75 cm thick (n = 6). Fig. 9, 17C.

DISTRIBUTION: The Caribbean watershed of N Central
America, in Honduras, Belize, Guatemala, and Mexico
(Yucatan, Quintana Roo, Campeche, Chiapas, Tabasco,
Oaxaca, and Veracruz; perhaps also in Puebla).
Reported from Panama (Dressler, 1993) but the report
seems to be based on specimens of V dressleri.
ECOLOGY: From the level to ca. 900 m elevation.
Vanilla insignis is probably the most common vanilla
in Mexico (Soto Arenas, 2003), where it is widely
distributed and forms large populations. It grows
in dry and wet areas (1000 to 4000 mm of annual
rainfall), but in the latter, is confined to savannas with
especial edaphic conditions. It has been recorded only
from calcareous substrates. In the Yucatan Peninsula
it grows in the subdeciduous forests of Bucida
buceras, Brosimum alicastrum, and Manilkara sapota,
often with the understory dominated by the palm
Cryosophila argentea; these areas have slow drainage
during the rainy season, and are frequently associated
to the flooded areas ("tintales") with Haematoxylon
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


campechianum.In the much moister areas of Chiapas,
it is found in savannas with Coccoloba belizensis,
Quercus oleoides, and Roupala borealis.In Central
Veracruz V insignis grows in tropical deciduous forest
or in warm oak forest.
Some specimens in Campeche and Chiapas seem
to occupy areas up to 4,000 m2, and undoubtedly they
are the largest plants of any Vanilla of the area. Some
of these specimens may prove to be the most massive
orchidaceous specimens in the world, with weights of
many tons. It flowers in April and May; fruits become
ripe in March-April. The flowers are visited by male
bees of Eulaema polychroma.
This is another member of the Vplanifolia complex
and it has been considered conspecific with Vplanifolia
by some authors (e.g. Williams, 1956). However, it is
clearly a distinct species; the flowers, though larger,
are rather similar to those of V planifolia in pressed
specimens. The floral fragrance is also similar to that of
V planifolia (1-2-dimethil-ciclopentane, ethyl acetate,
and 1-8-cineol as principal constituents, although
ocimene-trans is notoriously absent.
It is easily recognized because the stems have
internodes conspicuously sulcate and their surface is
fairly rugose-papillose. Similar vegetative traits are
found in some very distantly related Asian species,
namely V yersiniana Guillaumin & de Sigaldi, V
moonii Thwaites, and it allies. The sulcate stems are
thought to be an adaptation that permits the stem to store
more water during the rainy period in seasonal areas,
since the groove expands and becomes inconspicuous
when the stem tissues are swollen. The characteristic
flowers are large, ca. 11-12 cm in diameter, with green
tepals, cream-white lip, the midlobe adorned with thick,
triangular, retrorse projections, up to 4 mm high. The
fruit (ca. 10-12 cm long) is thick and fragrant when ripe,
but the aroma is similar to a mixture of vanilla with
coconut.
The ITS tree (Fig. 1) and a survey of additional
genomic regions, including also non-Mesoamerican
Vanilla species show that the closest relatives of V
insignis are V odorata Presl, V helleri A.D.Hawkes,
V uncinata Huber ex Hoehne, and V tahitensis
J.W.Moore. Vanilla odorata, V uncinata, and V
tahitensis have much narrower leaves, non-sulcate
stems, smaller flowers, and less developed retrorse
appendages on the lip. From V helleri, from Oaxaca,








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Nicaragua and Costa Rica, with which it shares the
papillose stem surface, sulcate intemodes, and long,
retrorse, orange-coloured papillae on the distal part of
the lip, it can be easily distinguished by the long claw
of the lip found in V insignis vs. the cuneate lip blade
of V helleri.
Vanilla insignis was described in 1934, and it is
surprising that it has not been reported until recently
from Mexico (Camevali et al., 2001; Soto Arenas,
2003) although it was collected in Veracruz by C.A.
Purpus in 1919. Neither has it been reported so far from
Guatemala or Belize, although a picture of V insignis
appeared wrongly identified as V pfaviana in "Native
Orchids of Belize" (McLeish et al., 1995). However,
this species has been known and used by the Mayas for a
long time, who call it "sisbic". Almost all the specimens
previously identified as V planifolia or V fragrans
from the Yucatan Peninsula and Veracruz belong to V
insignis. The report of V odorata from Quintana Roo,
based on Cabrera 4611 (Soto Arenas, 1989) is based in
a specimen with buds that is actually V insignis.
Vanilla insignis, could add desirable features to
the commercial vanillas; it also has fragrant fruits, it
is a more xerophytic, stouter species, and apparently
tolerates clayey soils, seasonally flooded in summer.
We have cultivated this species and it is also the
Mesoamerican species most tolerant to low and high
temperatures, and much more resistant to the attack of
pathogens than Vplanifolia.

OTHER RECORDS: MEXICO: VERACRUZ: Zacuapan,
June 1919, C.A. Purpus 8482 *AMES(71362)! NY! Near
Zacuapan, in humid forest on shrubs and on trees, 13 Feb
1932, 9652' W, 1912' N, ca. 900 m, O. Nagel sub E.
Oestlund 2682 AMES(*41478, *51844, sterile)! Region
below Zacuapan, on shrubs, humid forest, 8 Jul 1936, ca.
800 m, C.A. Purpus sub E. Oestlund 5961 *AMES(51845)!
Near Zacuapan, in humid forest on shrubs, 15 Jun 1935, ca.
800 m, C.A. Purpus 4866 AMES(*51849 fls. in spirit not
seen, *51848, sterile)! Carretera Xalapa-Veracruz, km 16
SE of Xalapa, 1 km SE of main Jalapa-Huatusco highway,
5 km SW of bridge over Rio Los Pescados, 5 km (by air)
SE of Tuzamapan, Mpio. Coatepec, 19021'N, 96 50"W,
680 m alt., "selva baja caducifolia", thorn scrub along now
dry canyon, now very dry and most trees without leaves.
Vine to 3 m, fruits green, hanging, March 19, 1983, M. Nee
& K. Taylor 26045 F(1985066)! *XAL! Mpio. Emiliano
Zapata, Cerro de Chavarrillo, 1926'N, 96047'W, alt. 850
m s.n.m., selva baja, primaria, suelo arcilloso, pedregoso,
color negro, muy seco, calido, bejuco perenne, 6 m, escaso,


fruto verde; trepadora, 15-04-1979, G. Castillo & L. Tapia
531 F(1963631)! NY! *XAL! Mpio. Emiliano Zapata,
desviaci6n de la carretera Xalapa-Veracruz, 16 km al SE
de Xalapa, a 900 m de la carretera, enredadera, flor blanca
y amarillo, escasa, vanillala, 23/V/1976, C.H. Ramos 402
MEXU! [cf, fruits] Mpio. Soteapan, San Fernando, 18 17',
600 m, acahual, selva alta perennifolia, 2-3 m, vanillala,
19-IX-86, (usos) aromatizante de aceite, M.C. Gonzdlez R.
303 *XAL! OAXACA: [cf., sterile] Forests ca. 25 km east
of Mogofie, near Rio del Corte. On shrubs, rooting with
long aerial roots in leafmould. Isthmus of Tehuantepec, ca.
9457' W, 170' N, alt. ca. 100 m, 20 feb 1935, 0. Nagel
sub E. Oestlund 4584 AMES(51847)! *US(1805098)!
Plan Juan Martinez, camino Reforma-Ayozintepec, 80 m.
Comprada al Sr. Eugenio Hilario Justo, quien la colect6
expresamente para nosotros. 19-III-1997, M. Soto 8120
y A. Cibridn *AMO(sterile)! TABASCO: [cf., sterile]
Balancan, carretera no. 25, km 45 del entronque con la
carretera E.W.O. hacia la carretera W-10, 10 m s.n.m.
selva median subperennifolia primaria, asoc. Manilkara
sapota, calido himedo, epifita, perenne, 5 m, escasa, nom.
vul. vanillala, 06-12-1975, P.E. Valdivia 2063 XAL!
CHIAPAS: [cf, fruits] La Cueva, al NW del Rancho
Corocito, Reserva del Ocote, Mpio. Ocozocuautla. Alt.
770 m s.n.m., selva median perennifolia, primaria, suelo
negro delgado con rocas calizas, ruderal, hierba, perenne,
3 m, escasa, fruto verde, nom. vul. vanilla, 29-04-1983,
J.I. Calzada, P. G6mez & B. G6mez 9695 *XAL! Mpio.
Ocosingo, Estaci6n de Biologia de Chajul, Sabana I, a unos
3 km del Rio Lacantuim, bosque sabanoide con Roupala,
Byrsonima, Scleria, Pteridium, ca. 200 m s.n.m. 16008'
N, 90053' O. 13-IV-1997, M. Soto 8361 *AMO(buds)!
CAMPECHE: Tuxpefa, Dec. 1, 1931, C.L. Lundell 1070
F(700398)! Selvas medianas subperennifolias (Manilkara-
( I i.... !-I,,.,. y bajos inundables, entire el Ejido 20 de
Noviembre y las ruinas de Rio Bec, Reserva de Calakmul,
Campeche. Abundante, plants mas pequefias que en Nueva
Vida, s6lo una vista con botones. Los mayas del ejido no
la conocen. 11-IV-1995. M. Soto, E. Martinez, G. Tavera,
et al. 7656 *AMO! Reserva de Calakmul, cerca de Zoh
Laguna, selva median subperennifolia con Cryosophila
argentea, ca. 200 m altitude, 14 abril 1995, t6palos verdes,
labelo crema con ap6ndices anaranjados, frutos aromaticos,
fragancia a vanilla y coco, M. Soto 7667 AMO(x2, also in
spirit)! same data, [fruits], M. Soto 7670 *AMO! Reserva
de Calakmul, ca. de Nuevo Becar, en bajo, 13-IV-1995. M.
Soto 7681 *AMO! Ejido Nueva Vida, al N de Zoh Laguna,
selva median subcaducifolia-subperennifolia de Brosimum
alicastrum, Protium copal, Platymiscium sp., con much
Cryosophila argentea, 230 m s.n.m.; plant vigorosa a la
orilla del chilar; dos flores abiertas, 12.4 cm de didmetro,
cerraron cerca de las 12:30; 3 Eulaema se aproximaron a
las flores, no se posaron. Fragancia especiosa, d6bil, no
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


identificada, 16-IV-1995, M. Soto 7684 & E. Martinez
*AMO! AMO(in spirit)! Ejido El Refugio, selva inundable
de Bucida buceras a la orilla de la laguna, no la hay en
el bajo de Haematoxylon contiguo. 230 m s.n.m., 17-IV-
1995, M. Soto 7685 & E. Martinez *AMO(x2)! 200 m,
1835'N, 8924'W, 1 Feb 1076, P. Alvaro M & G. Bacao
173 MO. OUINTANA ROO: Dense forest near shore of
lagoon Chichankana, on shrubs. N part of lagoon, 8843'N,
1952' W, ca. 50 m, 16Aug 1935, 0. NagelsubE. Oestlund
4973 *AMES(51850, sterile)! *MO(1145555; sterile)
*US(1805110; sterile). Mpio. F. Carrillo Puerto, camino al
Ejido X'konha', 4 m s.n.m., lat. 19028'N, long. 8803'W,
selva median perennifolia primaria, calido himedo, suelo
pedregoso de color negro con much material organica, asoc.
con arboles, es epifita, abundancia regular, bejuco, 10 m,
perenne, tallo suculento, flor blanca, 8-V-1981, JS. Flores,
E. Uccn 8236 CICY *XAL! En C. Vallarta, a 17 km al oeste
de Puerto Morelos. Bejuco herbaceo con botones florales.
Selva median con Manilkara, Vitex y Thrinax, 17 de abril
de 1983, E. Cabrera 4611 yH. de Cabrera *AMO! MEXU!
[cf, sterile] A 16 km al S de la terminal del Ferry, cerca de
la entrada a Palancar, selva baja a median con abundante
Lonchocarpus, Dalbergia etc., suelos inundables, epifita
sobre tronco, 22/Nov/1982, E. Cabrera 9768, 0. TPllez, yE.
Linares MEXU(421555)! La Pantera, 1997, Carnevali s.n.
AMO(sterile)! YUCATAN: San Antonio, Rancho al sur de
Pixoy, lat. 20042'N, long. 8814'W, alt. 22 m s.n.m., selva
baja caducifolia, secundaria, en la orilla de una mensura,
suelo moreno, pedregoso; abundancia regular, hierba, 6 m,
perenne, frutoverde; obs. provocacomez6nenlapiel, 12-08-
1983, E. Ucan 2761 CICY *XAL(fruto)! Mpio. Valladolid,
Ebtun cabecera rumbo a Pixoy, lat. 20041'N, long. 8814'W,
alt. 22 m s.n.m., selva baja caducifolia, secundaria, en la
orilla del camino, suelo moreno, abundancia regular, hierba,
6 m, perenne, flor verde amarilla, 11-05-1983, E. Ucan
2463 *XAL! GUATEMALA: IZABAL: [sterile] climbing
on dry pine slope; leaves coriaceous, dull green above, paler
dull green beneath, stems dull olive-green, warty rugulose,
with a sulcation on each side, between Milla 42.5 and ridge,
6 miles from Izabal, Montafa del Mico, 65-800 m altitude,
April 1, 1949 JA. Steyermark 38539 F(1043863)! [cf., fruit]
A 8 km al NO de El Estor, 210 m s.n.m., hierba trepadora
con fruto, sabana,30 agosto 1988, E. Martinez 23348 & D.
Stevens MEXU(480867)!MO(3656561)!Punta Palma, Sto.
Tomas, 100 m de la entrada de la playa por el lado norte,
3 6 4 plants en la playa, 22 febrero 1998, M. Dix sub M.
Soto 8611 AMO! ALTA VERAPAZ: [cf, sterile] Climbing,
stems terete, deep green, savanna north of Concepcion, 3-5
miles southeast of Finca Yalpemech, near Alta Verapaz-
Pet6n boundary line, alt. 100-110 m, March 23, 1942, JA.
Steyermark 45233 *AMES(sterile, 63988)! F(1195510)!
BAJA VERAPAZ: [cf] Jocol6, climbing up trees, wild
species of Vanilla, fruits said to be short, used as flavoring,
IANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


100 ft, Jan 30 1921, H. Johnson 1178 AMES(22753, sterile,
perhaps V cribbiana)! PETEN: "Vainilla", fleshy vine,
Tikal National Park, Bajo de Santa F6, salida de Arroyo
Corriental, in tintal on Aguada T6rminos road, March-June,
1959, C.L. Lundell 15940 *LL(fruit)! same data C.L. Lundell
15818 *LL(fruit)! BELIZE: COROZAL: Maskall, Dec.
1933, P Gentle 1063 *AMES(40496, sterile)! NY(fruit)!
TOLEDO: "Vianilla", vine, in cohune ridge, Canada Hill-
Alta Vista Road, November 2, 1953, P.H. Gentle 8054
*LL(fruit)! NICARAGUA: ZELAYA: Cerro Waylawas,
10 km south of Siuna; elev. 250 m. Vine; scrambling over
rocks; leaves thick, fleshy (sterile). 5 June 1978. D. Neill
4219 SE(049333)!

9. Vanilla martinezii Soto Arenas, sp. nov.

TYPE: GUATEMALA: IZABAL: Mpio.
Livingston, El Golfete, a 20.4 km al NE de Rio
Dulce por lancha camino a Calix, bejuco herbaceo,
flor verde con amarillo y labelo blanco; selva
median perennifolia 'swampo', 15047'06"N,
8851'42"W; E. Martinez S. 36410 y D. Alvarez,
holo. MEXU!, iso. AMO! BIGUA! MO!

Vanillae inodorae similis sed foliis membranaceis-
chartaceis, inflorescentia brevis laxa quasi sub sessili,
bracteis viridis non foliaceis, floribus majoribus,
sepalis rectis, petalis undulatis, labello subintegro
vel quinque lateribus, fere long quam lato, carinis
humilibus inconspicuis ornato.

Hemiepiphytic vine, leafy, up to 10 m high.
Stems terete-subquadrate, somewhat keeled (in dried
condition), ca. 2-4.5 mm thick; intemodes 3.9-7 cm
long. Aerial, free roots pale brownish, dorsiventrally
compressed, 2-3.3 cm long, ca. 1 mm wide. Leaves
petiolate, the petiole up to 16 mm long, canaliculate;
blade elliptic, acuminate, base obtuse, somewhat
conduplicate; membranaceous-chartaceous (in dried
condition), 7.5-21 x 3.5-7 cm. Inflorescence strongly
disimilar to the vegetative shoots, 26-50 mm long, a
short, lax, 4-6-flowered raceme (rarely branched at
base), subsessile, the peduncle up to 13 mm long, the
rachis zigzag, at least ca. 3 mm thick, flowers separated
by 9-14 mm; with 1-2 peduncle bracts, clasping,
ovate, concave, up to 10 x 7 mm. Floral bracts ovate,
concave, obtuse to acute, apparently thin, 4-17 x 2-6
mm. Flowers successive, apparently 2 open at once,
very showy, with rather spreading segments, tepals
yellow green, lip white, ca. 5.5 cm high, 6.5 cm wide.







SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


I i
N )d


FIGURE 10. Vanilla martinezii Soto Arenas. Based on the type, E. Martinez 22790, andM. Soto 8602a. Drawing by M.
Lopez and M. Soto.
IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








LANKESTERTANA


Ovary terete, smooth, arcuate to straight, 2-sulcate,
the grooves almost straight, 52-53 mm long, 4 mm
thick. Dorsal sepal lanceolate elliptic, apex acute, base
obtuse, basally and apically grooved on the abaxial
surface, obscurely and broadly keeled at the middle,
margins thinner, somewhat reflexed and slightly
undulate along its entire length, fleshy, incurved at
apex, concave at middle, ca. 11-veined; 45 x 14.5 mm.
Lateral sepals very slightly oblique, elliptic, the upper
margin may be or not almost straight, acute to obtuse
at apex, almost subcalyptrate, slightly recurved and
curved upwards; basally obtuse, slightly canaliculate-
concave, broadly keeled on the abaxial surface, the keel
narrower and higher at apex, smooth, margins thinner,
minutely inflexed, almost entire and just slightly
undulate, little pleated to straight, ca. 11-veined, 46-
47 x 17-18 mm. Petals arcuate, obliquely elliptic to
lanceolate, somewhat arcuate upwards, apex long acute-
acuminate, recurved-rolled, base acute, subtruncate,
conspicually grooved all length of the abaxial surface,
ca. 10 veined, margins undulate and pleated, especially
near the apex, 43-47 x 11-13 mm. Lip fused basally
to the column ca. 7 mm, arcuate, funnel-shaped,
the lateral lobes-margins erect and forming a gullet
around the column, the apical lobe-margin deflexed;
impossible to flatten without some distortion, subentire-
obscurely trilobed, trapezoidal-subpentagonal to
suborbicular-subpentagonal, base truncate with widely
rounded 'shoulders', apex truncate-rounded, minutely
mucronate, appearing triangular in natural position,
axially grooved on the abaxial surface; 42-44 x 44-47
mm, ca. 27-34-veined, veins branched above the middle;
callus lip length, made up by a broad, 7 mm wide,
flat, fleshy plate constructed by 3 obscure keels from
the base to near the middle, progressively becoming
ovate-triangular in cross section and then divided in
ca. 11 low, sinuous, rugose-warty, inconspicuous keels
reaching the apex as a narrow high, almost smooth keel;
the lateral ca. 10 veins (on each side) slightly raised and
minutely rugose-warty; lateral lobes-margins almost
entire, and apparently somewhat reflexed, apical lobe
margins slightly undulate-pleated. Column relatively
short, strongly arcuate, the basal part forming a rather
abrupt 600 angle with the apical part, semiclaviform,
apex dilated, smooth, ca. 28 mm long (across de arch);
vertical wings oblong, inconspicuous, 4.5 x 1 mm.
Anther strongly attached to the clinandrium by a broad,

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


short filament, ovoid, ca. 4.5 long, 2 mm wide. Stigma
a cavity, the midlobe concave, oblong, perpendicular to
the column axis. Fruit clearly dehiscent, opening by 2
sutures, leaving unequal valves, 2-3 and 7-10 mm wide
respectively, non-aromatic except by a resinous faint
smell, dark brown, blackish inside, with ellipsoid seeds
with slightly warty surface; 9.5-15 cm long, 6-8 mm
thick before dehiscence. Fig. 10.

DISTRIBUTION: Known only from eastern Guatemala, but
also to be expected from adjacent wet areas of Belize
and Honduras. It may also be native to Mexico. DNA
from a sterile specimen of a membranaceous Vanilla
collected in Crucero Corozal, in the Selva Lacandona,
Chiapas, Mexico, A. Ibarra P. 2222?, was sequenced
(ITS, matK), and it shows a strong relationship to V
martinezii, yet it seems different from the sympatric
V inodora; we suppose that it belongs to V martinezii,
although its sequences are somewhat divergent from
the Guatemalan material. This locality is known to
have populations of very thermophilous plants, that
are very rare in other rain forest areas of Mexico (e.g.
Lacandonia schismatica, Chysis limminghei, Warrea
costaricensis, Ligeophila clavigera, Specklinia haberi,
Maxillaria alba); furthermore the substrates in this
area are partially flooded and with peat-like soils.
ECOLOGY: In lowland, wet, swampy areas of high
rainfall. Locally abundant. Flowering in February and
July. As far as we know, this vine grows only on islets
in the delta of the Polochic River into Lake Izabal and
similar habitats near the coast in the area known as
Golfete; its habitat could be very specific, since these
islets have an unusual peat-like soil. In the only flower
that we have examined, the midlobe of the stigma is
perpendicular to the column body, and the anther is
also protruding. The fruit set in Vanilla martinezii is
very high (up to 53% in a clone) which suggests that it
could be self-pollinated.

Vanilla martinezii is known only from two or
three nearby localities. Vanilla martinezii is a species
of the membranaceous group; which, together with
V costaricensis, V inodora, and V sarapiquensis,
are the only members of this clade in Mesoamerica.
It is different from other species by the following
combination of characters: membranaceous-
chartaceous leaves, short, lax, almost subsessile
inflorescences (strongly different from the vegetative








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


shoots); short, non-foliaceous bracts, large flowers
with straight sepals, undulate petals, and by the huge
subentire to subpentagonal, free lip, about as long as
wide, and with very low and inconspicuous axial keels.
Both V costaricensis and V inodora have elongate
inflorescences similar to their vegetative axes, and leaf-
like bracts. Vanilla inodora is also easily distinguished
by its cushion-like, axial callus and its emarginate
lip; V costaricensis is more similar to V martinezii,
but has smaller flowers, twisted-undulate sepals, and
the lip surface is adorned with a rugose sculpturing,
especially at the sides of the lip.
The most similar species is Vguianensis Splitgerber,
from Guyanas and Amazonia (better known by its
synonyms V acuta Rolfe and V latisegmenta Ames &
Schweinf.). Both have membranaceous-chartaceous
leaves, short, few-flowered inflorescences with small,
non-foliaceous bracts, subentire lip which is strongly
veined, and similar column morphology with basal
wings. However, in Vanilla martinezii the axial keels
of the callus are warty, numerous, and not well-defined
at the apex, some of them extending to the basal,
lateral sides of the lip and the latter is not as trilobed at
apex; in V guianensis the callus is formed by 5 basal
keels and 3 rather prominent ones, but it is not or only
scarcely verrucose, and the lip apex is trilobed.
Vanilla martinezii seems to be another endemism
of the wet lowlands around Lake Izabal, a site which
has been proposed as a primary refuge for a diverse
tropical rain forest biota (Toledo, 1982; Wendt, 1989,
1993). Its affinities with the Amazonian V guianensis
make its distribution even more interesting.

OTHER RECORDS: GUATEMALA: IZABAL: Creek Lagarto,
Ensenada de los Lagartos, El Estor, 2 m s.n.m., bejuco,
flor blanca, selva median perennifolia inundable, 16 julio
1988, E. Martinez 22790, P. Tenorio, H. Droege & M. Diaz
MEXU(480869)! Lago Izabal, desembocadura del Rio
Polochic, Creek Lagarto, al SW de El Estor, selva median
inundable con Pachira aquatica, sobre suelos con much
material organica (peat), con Epidendrum stamfordianum,
E. flexuousum, E. cardiochilum, E. raniferum, Oncidium
sphacelatum, 0. luridum, Pleurothallis marginata, P.
sertularioides, Maxillaria crassifolia, M. elatior, Gongora
aff. quinquinervis, Coryanthes picturata, Myrmecophylla
brysiana, Sobralia decora; cerca del nivel del mar, ca.
1528'N, 8923'W; comin, hasta de 10 m de alto; 24 febrero
1998, M. Soto 8601aAMO! mismos datos, 23 capsulas de
43 flores, M. Soto 8602a AMO!


The next specimen from a nearby locality is sterile, but
it matches V martinezii in vegetative aspect; however it
could be V inodora Schiede, usually with broader leaves:
IZABAL: Vicinity of Quirigua; altitude 75 to 225 m, May
15-31, 1922, P.C. \i.,-- .r24554

10. Vanilla odorata C.Presl, Reliq. Haenk. 1: 101.
1827 [1830].

TYPE: [ECUADOR:] Hab. in Guayaquil, Haenke,
holo. PR(305753)! iso. (x2; 305751,305752)
[all sterile and mounted with fragments of a
Dimerandra species].
V ensifolia Rolfe, Kew Bull. 1892, p. 141.
Syntypes: [COLOMBIA:] Cauca, Pefiol,
Aout, Goudot, K! [with a drawing of the other
syntype, "Leaves, flowers & seed of Vanilla
-- Patia, presented by Mr. J. Hanbury 1884"
Herb. Pharmaceut. Soc.]; P(flowers poorly
preserved)!
Epidendrum vermifugum Sess6 & Moc., Fl.
Mex. ed. 2: 201. 1894.
Lectotype: (Soto Arenas, 1994): "Epidendrum
vermifugum, Sesse, Mociho, Castillo &
Maldonado (4358) MA [sterile]!; isolectotype
"Epidendrum vermifugum de Mexico absque
foliis" [only floral buds, mounted with
"Epidendrum uniflorum"], BM! F(848611)!

COMMON NAMES: vanillaa Tlatepusco" (Usila, Oax.);
"Vainilla de Teutila", Humboldt.

Hemiepiphytic vine, branching, leafy, up to 6 m
high. Stems flexuose, terete, smooth, dark green, with
whitish dots, 4-6 mm thick; intemodes 7-10.5 cm long.
Terrestrial roots conspicuously pubescent; aerial, free
roots terete, green to pale green, 1 mm thick; ,lil., .niii
aerial roots strongly flattened. Leaves subsessile; the
blade lanceolate to ensiform, very narrow, sometimes
oblique, acuminate to long acuminate at apex, very dark
green on both surfaces, 8-13 x 1.5-2.7 cm; ca. 1.8 mm
thick in fresh. Inflorescence a 6-12-flowered raceme,
rachis 30 mm long, 6 mm thick. Bracts subsessile,
concave, membranaceous, subsessile, progressively
smaller, ca. 8 x 5 mm. Flowers successive, 1-2 open at
once, with spreading segments, ephemeral (from 7:00 to
16:00 hrs), showy, tepals whitish green, translucent, lip
greenish white, the throat striped with pale yellow lines,
callus white, column white; ca. 8 cm high, 7 cm wide;

IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.






LANKESTERIANA


Cim


3cm

10 cmiI




FIGURE 11. Vanilla odorata Presl. Based onM. Soto 6617. Drawing by R. Jimenez.
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


2 cm


5cm


LI^


-7


3cm


q








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


fragrance weak, green, fresh. Ovary terete, smooth,
ca. 3 mm thick. Dorsal sepal linear to oblanceolate,
apex acute, rounded, subcalyptrate, base attenuate-
subunguiculate, concave, 9-veined, 47-54 x 8.0-8.3 mm.
Lateral sepals narrowly elliptic to oblanceolate, oblique,
apex subacute, rounded, subcalyptrate, base attenuate-
subunguiculate, concave, ca. 12-veined, 44.5-52 x 9.5
mm. Petals obliquely linear to oblanceolate, somewhat
arcuate, apex obtuse, rounded, sometimes emarginate,
oblique, fleshy, slightly calyptrate, base long attenuate;
concave; with a very conspicuous, axial, elevated, flat,
keel at the abaxial surface, ending in a cylindric, free,
adpressed process, ca. 1 mm long; ca. 11-veined, 46-53
x 7 mm. Lip attached to the column along the margins
of the basal half (ca. 23-29 mm), tubular, cymbiform,
the apex abruptly deflexed; axially grooved on the
abaxial surface, the groove well-defined and deep; when
spread out 42-49 x 19-26 mm; long unguiculate, the
claw dense papillose at base and apex on the the inner
surface, the papillae digitiform, unicellular, 13-14 x 2.7-
3 mm; the blade obovate-flabellate in outline, obscurely
trilobed, ca. 20-veined, the veins branched in the distal
third and slightly thickened; the lateral lobes narrow
and obliquely obovate, margins long lacerate-fimbriate
(cilia 1-3.5 mm long), ca. 25 x 8-9 mm; midlobe
approximately subquadrate-semiorbicular, the margin
undulate-crispate, laciniate, obscurely emarginate, 7-9
x 7-11 mm; penicillate callus at 29 mm from the base,
3-4.5 x 3 mm; made up by ca. 9 imbricate, flabellate-
praemorse, retrorse, laciniate scales, sometimes united
each to other along the lateral margins, with some
adpressed processes on the surfaces; continuous towards
the lip apex with 3 inconspicuous keels, progressively
thickened, less defined, and more confluent at apex,
almost forming an apical, fleshy cushion, ca. 14-15 x
2 mm, with 3-4 conic, prominent, retrorse papillae and
many small, elongate warts; other warts on the veins of
the lateral lobes. Column very elongate and slender,
semicylindric, 33-38.3 mm long, 2 mm wide; ventral
surface flat and papillose, the papillae digitiform, septate,
bigger towards the apex, absent at the basal third; apex
dilated, with vertical wings, inconspicuously trilobed,
ca. 2 mm long, 3 mm wide; clinandrium prominent,
transversely oblong, concave, ca. 2 x 3 mm. Stigma
trilobed, the lobes emergent; rostellum a flabelliform,
somewhat convex blade, ca. 2 x 3 mm, covering and
parallel to the lateral lobes, quadrate, slightly divergent


each to other, ca. 1 x 1 mm. Anther versatile, attached
to the clinandrium margin by a laminar, broad filament;
body urceolate in outline, with 2 upper, ovoid, small,
divergent lobes; 3 x 2.5 mm. Fruit narrowly cylindrical,
slightly compressed, attenuate at apex, dark green, 17-
17.5 cm long, 8-10 mm thick; strongly fragrant, aroma
similar to that of Vplanifolia. Fig. 11.

DISTRIBUTION: Mexico (Veracruz, Oaxaca, Tabasco, and
Chiapas), Guatemala, Belize, Honduras, Nicaragua,
Costa Rica, Panama, Colombia, Ecuador, Peru,
Bolivia, and probably Brazil.
ECOLOGY: This species can be common in secondary
vegetation derived from tall, evergreen, tropical forest
and rainforest, in gaps or in slopes with cleared canopy
in primary forests. It is known from the sea level to
650 m altitude. Flowering time is mainly in April and
May, but the vanilla growers in the Oaxacan Chinantla
mention other sporadic flowerings in August and
November. The flowers remain open until 2:30 PM.

The narrow, lanceolate to ensiform, long acuminate
leaves, thin stems, translucent whitish-green tepals,
white-greenish lip with the throat striped with pale yellow
and the laciniate margins of the lip are characteristic.
Vanilla insignis is very similar, but it has thicker, sulcate,
rugose-papillose stems, broader, xeromorphic leaves,
and the retrorse appendages of the callus are much more
numerous, bigger and orange-yellow.
Mesoamerican specimens of V odorata are very
similar to that illustrated by Blanche Ames (Ames,
Sched. Orch. 9: 1-6. 1925) based on Ecuadorian
material and supposedly close to the type; perhaps
the segments are slightly broader and the wings of the
column have better-defined sinuses, and the laciniae
can be shorter in the Ecuadorian and Colombian (e.g.
Fonnegra et al. 1784) material. With these bases, the
small differences do not seen to warrant the recognition
of separate taxa.
Some specimens of Vanilla odorata from Belize
have been identified previously in herbaria as V hartii
Rolfe, and a picture of V odorata was labeled as V
hartii in "Native Orchids of Belize" (McLeish et al.,
1995). Soto (1989) reported V odorata from Mexico
based on a specimen with buds from Quintana Roo
(Cabrera 4611 & Cabrera), but that specimen actually
belongs to V insignis.
Humboldt cited a "vanilla de Usila". From his

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


description it seems that the involved species was V
odorata, which is well-known by the Chinantecan
Indians of the region, and grown on a small scale at the
present time.
At the beginning of the 19th century some vanillas
were used as vermicide in Mexico and Cuba, and that
is probably the origin of the Sesse and Mociflo name.
In the same way, the name "lombricera" was applied to
some vanillas in Cuba (Bold6 & Estevez, 1990).
Vanilla odorata produces aromatic fruits, similar to
those of Vplanifolia, with a strong, pleasant fragrance;
they are very much appreciated in the regions where
the species is wild. In Chiapas they are occasionally
used to flavor rums; in this area the species is much
more common than V planifolia, and actually it is
the only vanilla collected for its beans in the Selva
Lacandona, as in many other areas of Tropical America.
In northern Oaxaca the species is grown as a curiosity
in the plantations, intermingled with V. planifolia; the
growers mention different cultural requirements from
those given to V planifolia, since it needs stronger
sunlight. Although the fragrance is appreciated, the
beans are difficult to manage, since they are attacked
by fungi and because they dehisce if treated the same
way as the fruits of V planifolia; open fruits of vanilla
usually bring a lower price. Vanilla odorata is found in
dry to damp sites, usually in areas with higher rainfall
than those of V planifolia, and seems to be tolerant
to a wide range of light conditions. Vanilla odorata is
undoubtedly a species to be considered in any breeding
program with commercial vanillas (see comments
under Vanilla tahitensis, in "Excluded species").

OTHER RECORDS: MEXICO: VERACRUZ: [cf., sterile]
"Vainilla". Vine on "Jimba" shrubs and on trunks of trees.
Leaves leathery, glabrous. Common in some areas in dense
forest, absent in others, Fortuno, Coatzacoalcos River, alt.
30-60 m, February 1937, L. Williams 8919 F(897099)!
OAXACA: In sylvis umbrosis prope Lobani, Chinantla,
Oajaca, Liebmann 6142 W(11759)! Tuxtepec, Ejido
Chiltepec, vanillala, domina Lonchocarpus, M. Sousa
947 MEXU [sterile]! Dto. Tuxtepec, Usila, Arroyo Iguana,
vainillal de Silvano Bautista, ca. 250 m s.n.m., 24 abril
1992, M. Soto 6617 & M. Herndndez *AMO(x3; illustration
voucher)! Dto. Tuxtepec, Mpio. San Jose Independencia,
Cerro Clarin, en el extreme SW de la Presa Temascal, ca.
120 m s.n.m., 29 abril 1994, M. Soto 8829a & U. Sdnchez
AMO! AMO(in spirit)! same data, abril 1994, U. Sdnchez
s.n. y 6 AMO(in spirit)! Mpio.Valle Nacional, Arroyo de

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


Banco, vainillal del Sr.Lazaro Perez Justo, 250 m s.n.m.
con flores, muestreada para fragancia; menos atacada por
plagas y pat6genos que V planifolia, 23-IV-1997, M.Soto
8501 & M. Herndndez *AMO(flowers, photos)! San Felipe
Usila, en cafetal, vanillaa tlatepusco", 23, abril 1995, M.
Herndndez s.n. AMO(flowers in spirit)! km 49.8 del camino
Sochiapa-San Juan Lalana, 1.2 km antes de San Juan Lalana,
210 m s.n.m. Cafetal derivado de selva alta perennifolia, en
el fondo de canada rodeada de encinares calientes. Terrenos
de Galino T6llez, 18-III-1997, M. Soto 8115 & A. Cibridn
AMO(fruit)! Dto. Tuxtepec, Mpio. San Jose Independencia,
Cerro Clarin, en el extreme SW de la Presa Temascal, ca.
120 m s.n.m., 29 abril 1994, M. Soto 7631 & U. Sdnchez
AMO! AMO(in spirit)! CHIAPAS: alrededores del sitio
arqueol6gico de Bonampak, vegetaci6n secundaria derivada
de selva alta perennifolia, 350 m s.n.m., abril 1981, M. Soto
1001 AMO! Mpio. Ocosingo, km 5 del camino del Crucero
San Javier a Bonampak, vegetaci6n secundaria derivada
de selva alta perennifolia, sobre suelos rojos arcillosos,
ca. 350 m s.n.m. 26 junio 1996 M. Soto 7959 & R. Solano
AMO(fruit)! Mpio. Ocosingo: Estaci6n de Biologia Chajul,
en el borde del Rio Lacatuim; camino a Arroyo Miranda,
selva median subperennifolia inundable sobre terrenos
plans con Scheelea y Sabal, ca. 180 m s.n.m., muy escasa,
21 junio 1996, M. Soto 7950 y R. Solano AMO(sterile)!
BELIZE: Trail through light jungle, near Camp 2. Alt.
2000 ft. Wiss (vine) with glossy green succulent leaves and
stems. No flower or fruit seen. 20.8.1976. C. Whiteford 1316
MO(2584013)! GUATEMALA: IZABAL: [cf., sterile]
Twining vine; leaves deep green, succulent; fruit green,
thick and succulent, twisted, exuding clear thick liquid when
crushed; faint vanilla odor. Quebradas, 19-22 May 1919,
H. Pittier 8589A NY! US(1013492); ALTA VERAPAZ:
"Vainillita" Chirujija Oxec.; near the Finca Sepacuit6, April
23, 1902, O.F Cook & R.F Griggs 735 *US(408445)!
PETEN: "Vainilla". Fleshy vine, Tikal National Park, Tikal,
in botanal north of hotel, January 20, 1961 E. Contreras
1841 *LL(fruit)! La Libertad and vicinity, Aug.-Nov. 1933,
M. Aguilar H. 164 *AMES(40519; steril)! HONDURAS:
COLON: Capuchin site east, mangrove forest. 1.8 mi
strip on the north bank of rio Guaimoreto between old
bridge and opening of Laguna Guaimoreto 4.5 mi NE of
Trujillo on old road to Castilla. Lat. 15 57'30"N; Long.
85 54'30"W. 2 Feb 1981, J. Saunders 1008 *LL(sterile)!
*SEL(038496, fruit)! Lancetilla, 150 ft., Yuncker 4993
NY! NICARAGUA: SEGOVIA: E of Jalapa, elev. 1600
ft., May-June, A.H. Heller 6106 SEL(013289; 003851,
drawing, fragments)! "Segovia Prov.", A.H. Heller s.n.
F(1598348)! without data, A.H. Heller s.n. SEL(003851)!
ZELAYA: [sterile]" Vainilla", bejuco, sobre arboles,
Guamil de segunda clase. Area de Ocotal, Rio Grande,
Guamil o brefias sobre areas pantanosas, a lo largo del
Rio Grande, Alt. 0-15 m, Abril 23, 1949, A. Molina 2312








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


F(1364505)! Cerro Waylawas, ca. 1339'N, 8448-49'W,
elev. ca. 100-268 m; sheer dog tooth limestone peak and
plain on E side of peak. Pendant epiphyte, sterile. 16 March
1978. W.D. Stevens, B.A. Krukoff7385 SEL(049332; cf,
steril)! COSTA RICA: ALAJUELA: Upala, San Jose,
Alrededores de Laguna Las Camelias, 1028'N, 85"08'W
100 m, bejuco trepador, 19 Nov 1987, G. Herrera 1346
INB(sterile)! MO(3709189)! LIMON: Hamburg Finca, on
the Rio Reventaz6n below Cairo, altitude about 55 meters.
Large epiphytic vine; laeaves and fruit dark green. "Vanilla
nevermanii", "Vainilla". Febr. 19, 1926, P.C. i... .r. & J.
Valerio No. 48917 *AMES(32665; fruits)! *US(1309433)!
[cf] Hamburg Finca, on the Rio Reventaz6n, F Nevermann
s.n. *AMES(33060; poorly preserved flowers)! PANAMA:
PANAMA: [cf] Barro Colorado Island, 10 Jan. 1940, EW
Hunnewell 16433 *AMES(sterile, 87907)! COLOMBIA:
ANTIOQUIA: Hierba trepadora, t6palos externos verde
amarillosos, interos amarillosos. Carretera Mutata-
Pavarand6, entire Haciendas La Esperanza y Mocari,
150 m s.n.m. Marzo 6 1987. R. Fonnegra, FJ. Rolddn,
J. Betancourt, B. Echeverry, O. Escobar 1784 K!
*MO(3592235)! PERU: LORETO: Leticia on the Amazon
River, September 1929, L. Williams 3161 *AMES(43483;
sterile)! Herbaceous vine. Yurimaguas, lower Rio Huallaga;
alt. about 135 m; dense forest, August 23-September 7,
1929, E.P. Killip & A.C. Smith 29065 *AMES(43484; old
inflorescences)! BOLIVIA: IXIAMAS: Best kind here,
fruit fragrant, vine, common, climbing over small trees in
damp forest, 15-20 ft, 1000-1500 ft alt, Dec 13 1921, O.E.
White 1115 *AMES(28024; fruits)! BENI: Rurrenabaque.
Rank growing species; swamp woods, common. Fls. said to
be white, fruits make good vanilla. 900-1000 ft alt., Dec 3,
1921, O.E. White 1821 *AMES(28026; old inflorescence)!
REFERENCES: Ames, Sched. Orch. 9: 1-6, fig. 1. 1925;
Schweinfurth, Orch. Peru, Fieldiana 30(1): 43. 1958;
Hamer, Ic. P1. Trop. pl. 1193. 1984; Soto Arenas, Orquidea
(M6x.) 13(1-2): 295-300, figs. p. 296, 297. 1994.

11. Vanillaphaeantha Rchb.f., Flora 48: 274. 1865.

TYPE: Cuba, C. Wright 3351 W; AMES(71001,
iso.)! BM(another specimen is V. cf. poiteai)!
G(7889/109)! K(iso.)!

Vanilla planifolia var. macrantha Grisebach,
Vcat. P1. Cub. 267. 1866.

COMMON NAMES: "Tapia" (Panama).

Hemiepiphytic vine, branching, leafy. Stems
terete, smooth, green, 4-12 mm thick (in dried
condition); intemodes 8-15.5 cm long. Aerial, free
roots, terete, pale brownish, up to 10 cm long, ca. 2
mm thick; attaching roots strongly flattened up to ca.


5 mm wide. Leaves subsessile, the petiole twisted,
canaliculate, up to 8 mm long; blade narrowly oblong
to narrowly elliptic, base rounded-subcordate, apex
acute-apiculate, chartaceous (xeromorphic), green,
margins revolute, 10.3-19 x 2.2-5 cm. Inflorescence
a 4-6-flowered raceme, 25-45 mm long, ca. 4-6 mm
thick, peduncle 15-29 mm long, rachis, 11-15 mm.
Flowers successive, 1 open at once, ephemeral,
the segments spreading, at least 75 mm long; tepals
greenish-white to cream, lip white stained with
brownish at edges or orange-yellow. Ovary straight
to arcuate, subterete, smooth, 42 mm long, ca. 4 mm
thick. Dorsal sepal long oblanceolate acute-subacute,
rounded at apex, subcalyptrate; base long unguiculate,
attenuate, canaliculate, the claw ca. 24 mm long, ca.
3-4.5 mm wide, blade concave, ca. 10.5 mm wide,
ca. 12 veined, smooth, thick and fleshy, total length,
62-74 mm. Lateral sepals strongly oblique, long
oblanceolate, arcuate, apex acute, subcalyptrate and
minutely warty on the apex of the outer surface; base
long unguiculate, attenuate, canaliculate, claw ca. 22
x 5 mm; blade concave, 12-12.5 mm wide, ca. 14
veined, smooth, thick and fleshy, total length 58-72
mm. Petals long linear, arcuate, slightly sigmoid, apex
subacute-obtuse, base long attenuate, canaliculate-
conduplicate basally the rest concave, with an elevated
axial, flat keel on the outer surface, ending in a
triangular, acute, terminal process ca. 1 mm long, ca.
10-veined, thinner than the sepals, 61-73 x 8.5 mm.
Lip attached to the column along the margins of the
basal half (ca. 42-52 mm), tubular, trumpet shaped,
conspicuously cymbiform, deepest near the middle,
axially grooved on the lower surface, when spread out
62-72 mm; unguiculate, the claw canaliculate, with
two rows of trichomes on the apical half, 26-34 x ca.
3 mm; the blade flabellate, somewhat trilobed, margin
slightly undulate, ca. 36-veined, apex emarginate;
obliquely long obovate, rounded 30-33 x 12-14 mm;
midlobe distinct, transversely oblong, the margins
conspicuously reflexed, revolute, emarginate, 5 x 11
mm; penicillate callus at 40-45 mm from the base,
made up by ca. 12 congested, retrorse, approximately
trapezoidal, fimbriate scales, the scales regularly
united each to other along the lateral margins, ca. 9 x
5.5-6 mm, with an apical callus, low, inconspicuous,
thickened. Column elongate, conspicuously sigmoid,
51-60 mm long; ventral surface lanuginose towards the

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rca, 2010.









LANKESTERIANA


5 cm










3 cm


4 cm

FIGURE 12. VanillaphaeanthaRchb.f.. Based onR.S. W.,.' ... .. I. . .;, ... O.F Cooks.n. SEL! (inflorescence
with leaf) and C. WK Powell 412 (inflorescence with 2 open flowers); Dwyer et al. 4691 (column). Drawing by M.
Lopez.


LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


apex, below the stigma; apex dilated, vertical wings
somewhat bilobed, the lower lobe falcate, ca. 2 x 3.5.
Stigma trilobed, with a convex, midlobe, ca. 3 x 4.5
mm, lateral lobes transversely oblong. Fig. 12, 16D.

DISTRIBUTION: U.S.A. (Florida), Cuba, Lesser Antilles,
Venezuela, Trinidad, Mexico (Yucatan and Quintana
Roo), ?Costa Rica, and Panama. It is unknown if
similar plants found in Ecuador, Guyanas and Brazil
belong here or to V bahiana.
ECOLOGY: In Mexico this species grows in tropical
deciduous or semideciduous forest near swampy areas,
at about 100 m above sea level. Mexican specimens
are juveniles and no flowering or fruiting specimens
have been recorded. In Panama the species grows in
coastal vegetation and in savannas with Curatella,
Clethra, and Roupala.

A previous report of V phaeantha from Central
America was based on misidentified specimens of V
calyculata. Vanilla phaeantha is very common in the
West Indies, but it has been scarcely recorded from the
mainland, except from Florida. However, it reaches
the coast of Venezuela (G. Camevali, pers. com.),
and also the Yucatan Penninsula and central Panama,
in our study area. Mr. Neal Byrd, owner of Finca La
Gavilana, Province of San Jose, Costa Rica, cultivates
a Vanilla phaeantha that was apparently collected from
the wild somewhere in Costa Rica. The only flowering
specimens of this species that have been available to us
are from Panama, where it seems to be common on the
Pacific slope.
The oblong-elliptic, xerophytic, usually green-
yellowish leaves are usually shorter than the intemodes.
The flowers are very large, with greenish tepals, and
the lip is white with yellow stripes. It is related to V
bahiana Hoehne from Brazil. From the V planifolia
and V odorata complexes it can be distinguished by its
smooth disc.

OTHER RECORDS: MEXICO: YUCATAN: Cenote Mucul,
Cerca de Sotuta, selva baja caducifolia, ca. 100 m s.n.m.,
G. Camevali 4885 (cultivated specimen, not preserved,
ITS sequence)! OUINTANA ROO: Mpio. Oth6n P Blanco,
Ejido Caobas, alrededores de la Sabana del Jaguactal, unos
21 km al sur de la carretera principal Xpujil-Chetumal, A.
Cibridn 21, 23, R. Jiminez, G. Carevali AMO(sterile)!
COSTA RICA: LIMON: Guapiles, Los Diamantes,
collected 26-12-2000, cultivated at Finca La Gavilana, Rio


Savagres basin, N. Byrd 11-A-1-5 PANAMA: PANAMA:
Vine climbering up trees in low thicket. Farfan Beach;
roadside thicket adjacent to beach. 30 March 1969, J.D.
Dwyer L.N. Durkee & J.R. Castill6n 4691 *MO(1980689)!
Balboa, "Tapia", alt sea level, grows out in much sunlight,
sepals & petals pale green; lip white with stain of brownish
color at edge, flowers in April, 1924, C.W. Powell 412
*AMES(28219)! Ancon Hill, June 4, 1923, W.R. Maxon
6779 *AMES(33689)! *US(1180005)! Bismark above
Panama, Fls. nearly white except greenish apex; climbing on
trees several yards, March 12, 1908, R.S. Williams 588 NY!
*US(678159; det by Schlechter as V pittieri)! Common,
sea level, February 1924, epiphyte, Hort. Powell 137, C. W.
Powell 3507 *AMES(28290)! Climbing vinelike on the
Farfan Beach area, forward dune-high beach community,
18 June 1971, R.L. Lazor 5358 SCZ(2262; sterile)! Fort
Clayton, Farfan Beach area, vine with fl cream, 29 May
1966, E.L. Tyson 4106 MO(1917579)! SCZ(2263)! [cf.]
Moist thickets; herbaceous vine; leaves very fleshy. Along
the old Las Cruces Trail, between Fort Clayton and Corozal,
December 3, P.C. Standley 29103 AMES(31440, sterile)!
[cf.] Brushy slope; large herbaceous vine, common, Tumba
Muerto Road, near Panama, January 6, 1924 P.C. Standley
29731 AMES(31439, sterile)! VERAGUAS: Carretera
Panamericana, 4.3 km al W de El Higo, ca. 20 km al W de
La Mesa, sabana de Curatella, Clethra y Roupala con selva
riparia de Andira, Pithecellobium, Enterolobium y Acacia,
bejuco con frutos jvenes, extendido sobre algunos arbolitos
de la sabana, hojas muy pequefias, tallos delgados, M. Soto
9920 PMA(ITS sequence)! DOUBTFUL LOCALITY:
CANAL ZONE or ALAJUELA. Photo, April 15, 1925 O.F.
Cook *SEL(011359; specimen; 111357 photo, the photo in
envelope is of a membranaceous species)!

12. Vanilla planifolia G.Jacks., in Andrews, Bot.
Repos. 8: t. 538. 1808.

LECTOTYPE: West Indies, without proper
locality. Introduced into England by the Marquis of
Blandford and flowered in the collection of the Rt.
Hon. Charles Greville. The plate 538 was prepared
from Greville's specimen and was chosen as the
lectotype (Garay & Sweet, 1974).
Lobus oblongus aromaticus Clusius, Exoticorum
Libri Decem.: 72. 1605.
Araco aromaticus Hemandez, Thes. Rev. Med.
Nov. Hisp.: 38, fig. 1651.
Myrobroma fragrans Salisb., Parad. Lond. 2: t.
82. 1807, nom. illeg.
Based on the same specimen that served as the
type of V planifolia G. Jackson.

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LANKESTERIANA


Vanilla fragrans (Salisb.) Ames, Sched. Orch.
7: 36. 1924; Stehle, Fl. Descr. Antill. Fr. 1: 109.
1939; Porteres, Enc. Biol. 46: 234. 1954.
Vanilla sativa Schiede, Linnaea 4: 573. 1829.
Holotype: "Baynilla mansa Hispano-
Mexicanorum". Hab. sponte Papantlae,
Misantlae, Nautlae et Colipae inqque iisdem
pagis colitur"; Schiede, May 29 [single leaf]
BM!
Vanilla sylvestris Schiede, Schiede, Linnaea 4:
573. 1829.
Type: "Baynilla cimarrona, Hispano-
Mexicanorum. Hab. Papantlae, Nautlae,
Colipae", not located.
?Vanilla carinata Rolfe, J. Linn. Soc. London 32:
446. 1896. [or probably V gardneri but the type
is not useful]
Vanilla duckei Huber, Bol. Mus. Goeldi, Para,
5(2): 327. 1908.
Type: A. Ducke GOELD.
Vanilla bampsiana Geerinck, Bull. Jard. Bot.
Nat. Belg. 52: 345, fig. 1. 1982.
Type: Zaire, Bikoro: Lebrun 1459 BR!

COMMON NAMES: "Vainilla mansa", vanillal"
(Veracruz), "Xanat", "shanat", "caxixanath"
(Totonaco, Veracruz), vanillaa colibri" (Oaxaca),
"tlilx6chitl" antique Nahuatl (Badianus manuscript),
"Kuoley gm" (Chinanteco, Oaxaca), "zizbic" (Maya,
Yucatan); "juju" (Tabasco).

Hemiepiphytic or rupicolous vine, much branched,
leafy, up to 10 m high. Stems flexuous, terete, smooth,
dark green, whitish dotted, sometimes the new
stems covered with a whitish wax (Oaxacan plants),
(6.5)11-12(13) mm thick; intemodes 8-11 cm long.
Terrestrial roots pubescent, 1.4-1.6 mm thick; aerial,
free roots terete, 2.5-3.0 mm thick; ,.ll.i.cii, aerial
roots semiterete, flat on the surface in contact with the
substrate, 3 mm wide. Leaves subsessile, the blade,
elliptic, oblong, narrowly oblong, usually with parallel
margins, abruptly acuminate to subacuminate, somewhat
oblique (variable), upper surface deep to pale green,
paler on lower surface, 9.5-23 x 3.5-7.6 cm, ca. 1.3-2.4
mm thick in fresh. Inflorescence a 7-18(-70) flowered
raceme, up to 26.5 cm long. Flowers successive, 1-2
open at once, segments variably spreading, most
frequently the tepals ca. 40 with respect to the column;

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


ephemeral, showy, tepals pale green to whitish green,
lip pale cream-yellow to yellow with ochre in the throat
and papillae; ca. 62-64 mm high, 39 mm wide, fragrance
weak, sometimes a cinnamon can be appreciated, herbal,
or white (e.g. Hymenocallis-like). Ovary arcuate at
base, upper part straight, terete, smooth, basally white,
the rest green, with extrafloral nectaries, 57 mm long, 4
mm thick. Dorsal sepal narrow elliptic to oblanceolate,
apex subacute, rounded, subcalyptrate, base attenuate-
subclawed, slightly concave, canaliculate basally, ca.
9-veined, 55-60 x 10-12 mm. Lateral sepals obliquely
narrowly elliptic, to oblanceolate, apex subacute,
subcalyptrate, base attenuate- subclawed, almost flat,
slightly canaliculate basally, margins slightly involute,
ca. 10-veined, 54-60 x 12-13.5 mm. Petals obliquely
long oblanceolate, somewhat arcuate, apex slightly
reflexed, obtuse-rounded, oblique, notched, slightly
thickened, base attenuate; concave, canaliculate at
base; with a very conspicuous, axial elevated, flat keel
at the abaxial surface, finishing in a cylindrical, free,
ascending, free process, ca. 2 mm long; ca. 13-veined,
55-58.5 x 10-11 mm. Lip attached to the column along
the margins of the basal half (ca. 31-35 mm), tubular,
trumpet-shaped, very concave to cymbiform, slightly
sigmoid, the apex abruptly flared, deflexed-recurved;
opening subtriangular, 18 mm wide, axially grooved on
the abaxial surface, the groove well-defined and deep;
when spread out 49-55 x 24 mm; long unguiculate, the
claw pubescent, trichomes short, yellow to ochre, 15 x
4 mm; the blade obovate-flabellate, trilobed in outline,
ca. 36-veined, the veins branched in the distal third, and
slightly thickened; the lateral lobes obliquely triangular-
flabellate, margins widely undulate, denticulate towards
the midlobe; teeth less than 0.6 mm long, ca. 27 x 10.5
mm; midlobe approximately subquadrate-transversely
oblong, emarginate to deeply bilobed, the margin
undulate-crenate, 5 x 9 mm; penicillate callus at 30-
37 mm from the base, 5.8-6 x 4-4.5 mm; made up by
ca. 8 imbricate, flabellate-praemorse, retrorse, lacerate
scales, pale yellow, the distal ones bigger, sometimes
united each to other along the lateral margins,
continuous towards the lip apex with 2 conspicuous
rows of papillae, and 2-4 inconspicuous rows, apical
papillae bright green. Column very elongate and
slender, trigonous-semicylindrical, 39-42 mm long, 2.5
mm wide; with vertical, flabellate to obscurely lobed-
erose, acute to rounded, membranaceous wings, 2 mm








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


long, 4 mm wide. Stigma trilobed, the lobes emergent;
midlobe a trapezoidal, convex blade; ca. 3 x 2 mm;
covering and parallel to the lateral lobes, longitudinally
oblong, slightly divergent each to other, ca. 1 x 1 mm.
Anther versatile, attached to the clinandrium margin
by a laminar, broad filament; body ovate-cordiform, 3
x 3 mm. Fruit straight to strongly arcuate, cylindrical-
subclavate, thickened towards the apex, often 2-3
conspicuously sulcate in wild specimens; smooth in
cultivated and many wild specimens; 10-30 cm long,
7-10 cm thick. Fig. 16E-F, 17D.

DISTRIBUTION: Imperfectly known. Mexico (Veracruz,
Oaxaca, Chiapas, Tabasco, Quintana Roo, and
probably escaped in Yucatan and southern Oaxaca),
Guatemala, Belize, Honduras, and Costa Rica. It
is unknown if the plants in other areas are native or
escaped from cultivation. For example, in Panama it
is known mostly from Barro Colorado Island and San
Blas Province, perhaps an indication that it is escaped
there.

The species seems to be escaped in Florida (cf.
Luer, 1972), and in Jamaica (Fawcett & Rendle, 1910),
two areas well botanized by experienced collectors.
Foldats (1969) indicated that the species is common
in Venezuela, but did not cite records; neither did
Garay and Dunsterville include it in their works on
Venezuelan orchids. South American wild specimens
previously identified as V planifolia and others from
elsewhere outside Central America have proven to be
misidentifications. On the other hand, there are some
collections from Ecuador that match closely the Central
American, cultivated material. It is doubtful that V
planifolia is native in regions outside of Mesoamerica.
Suposedly wild specimens of V planifolia from Rio
Palenque Center, Ecuador, have proven to be V hartii.
The original distribution in Mexico is also
uncertain. Northernmost, wild-collected specimens
come from the region of Cordoba, in Central Veracruz,
some hundreds of kilometers southwards from the area
where the species supposedly was domesticated in
northern Veracruz and Puebla. Most collections come
from northern Oaxaca, where great morphological
variation has been observed; a single record from
southern Oaxaca, on the Pacific slope, has not been
confirmed by recent collections. The species is very
rare in Chiapas. There are several old collections from


Yucatan, in the xerophytic thorn scrub of the northern
part of the peninsula; we suspect that they represent
escaped, old relicts from cultivated plants, since this
habitat is very different from the moister forests where
it is wild at present.

ECOLOGY: Dwelling in tall tropical evergreen or
semievergreen forests, from 150 to 900 m, rarely to 1300
m altitude. The species seems to prefer moist forests,
seasonally dry in spring, on calcareous terrain. It is absent
in volcanic areas and in the wet tropical rainforests of
Mexico. In moister areas it can be found in secondary,
very young forests, or in large gaps. It flowers mainly
in March to April, but varies from year to year, and
flowering is rather gregarious in a region. Occasionally
it flowers from February to May. Flowering seems to
be a response to low winter temperatures, followed by
strong sunny conditions in early spring.
Vanilla planifolia faces severe conservation
problems in the wild. There are very few and scattered
locations known at present; the largest populations in
northern Oaxaca have been completely removed as a
source of cuttings to establish new plantations; it is
also almost extinct in Veracruz, where only two clones
have been located in recent years. Only four isolated
specimens have been located in Chiapas.

Vanilla planifolia is the vanilla of commerce, and
the most widely planted species for its aromatic fruits.
The nomenclature of this species is very complex,
since its history began before the Linnean system
of nomenclature. The problem has been solved by
Garay & Sweet (1974), who lectotypified the species
with a plate based on the original plant (leaving aside
the statement of the identity of an earlier Plumier's
polynomial); it seems pertinent to reproduce here
their observations: "Jackson regards Plumier's
unpublished drawing of "Vanilla flore albo, fructa
breviori, corallino" as representing his V planifolia.
A study of this plate, however, convinces us that it
represents Vanilla eggersii Rolfe. Salisbury based
his Myrobroma fragrans also in Greville's cultivated
specimen, but cited in synonymy Epidendrum rubrum
Lam. Since in his publication Epidendrum rubrum is
an integral part of the protologue, he should have made
a transfer of Lamarck's epithet. It is of further interest
that Salisbury equates or rather confuses Epidendrum
rubrum Lam. with Plumier's polynomial description
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LANKESTERTANA


referable to Vanilla eggersii Rolfe." Therefore, the
widely used name Vanilla fragrans must be rejected,
because it was based on an illegitimate name.
The plate selected as the lectotype is not accurate
enough to permit the recognition of the species;
especially the much fringed lip suggests another
species of the group, such as V odorata or V insignis.
Hooker (1891) also expressed concern about the
identity of this plate, and recognized the differences
among Andrews' and Salisbury's plates. However, the
same Greville's individual plant was illustrated and
precisely described a year earlier by Salisbury, when
he published his Myrobroma fragrans, the plate and
description are much more accurate, and undoubtedly
belong to the widely cultivated, "Mexican vanilla",
discarding the possibility of it being another related
species. The history of Vanilla planifolia has been
reviewed by Dillon (1942) and Bruman (1948).
The published plate (Flore d'Afrique Centrale, pl.
39) of the recently described V bampsiana Geerinck,
from Zaire, is indistinguishable from our concept of
V planifolia. There are no vanillas with lamellose
transverse callus, fimbriate margins, long unguiculate
lips and papillose apices outside of Tropical America.

VARIATION AND CULTIVARS: Vanilla planifolia has proven
to be somewhat variable in flower size, degree of
flower opening, lip color, lip concavity, development
of the lip papillae, intensity of fragrance, development
of grooves on the fruit, self-compatibility, etc. This
variation initially led us to consider the existence of
more than one species involved; especially because of
the uniformity of plants in cultivation (the cv. "mansa")
compared with wild-collected specimens. However, it
is possible that the narrow morphological variation
seen in cultivated plants must be attributed to reduced
genetic variation (Cibrian, Soto Arenas, ). Studies of
gas chromatography indicate that the floral fragrance
of different forms is composed primarily of the same
compounds (1-2-dimethyl-cyclopentane, ethyl acetate,
1-8-cineol and ocimene-trans); 1-8-cineol especially
is well known to be a strong attractant for euglossine
bees. Since V planifolia is pollinated by Euglossa
viridissima and maybe other species of the same genus,
the fragrant, attractive compounds can be considered
as evidence for the specific integrity of all the known
forms.
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


The growers in Veracruz recognize different
types in V planifolia, and these perhaps represent
different genetic individuals (clones), since no
genetic manipulation or breeding programs have been
conducted in Mexico. These clones are better treated
as cultivars.

cv. "mansa" or "dura"
This is the form widely cultivated in northern
Veracruz, and probably in Africa and Asia; the plants
from Asia (except the Philippines) and Africa are
descendant of the original type specimen grown by
Grenville and distributed to many botanic gardens
during the 19th Century. It is unknown if more than
one clone is involved, but the plants are very uniform
in morphology. Different from many wild specimens
are the non-sulcate fruits.

cvs. "acamaya", "rayada", and "variegata".
The stems and leaves have yellowish stripes,
alternating with the common deep green of the other
cultivars (see Roullet, 1990). There are at least two
different genetic individual with variegated plants
(SchlUter et al. 2007). There seem to be no differences
in molecular markers between one of this clones and
'Mansa', suggesting that it can be a sport (due to
somatic mutation).

cv. "albo-marginata"
A cultivar with beautiful leaves, margined with
white (see Roullet, 1990). It has never been found in
Mexico, and it is probably also a sport of Madagascan
origin.

cv. "oreja de burro"
? Vanilla sylvestris Schiede
The cv. "oreja de burro" is occasionally grown as
a curiosity in the plantations of the region of Papantla.
It is very similar to the cv. "mansa", and difficult
to distinguish except by experienced growers; the
stems are slightly thinner, the leaves rather hanging,
because the more elongate petiole, elliptic, abruptly
acuminate, concave, with more prominent veins, and
more greenish flowers. Once pollinated it forms a large
fruit up to 30 cm long, usually bisulcate, but almost
all the fruits are aborted in July, except sometimes
when the pollen came from a different clone or from V
pompona; therefore, it is self-incompatible. Additional
watering or feeding does not seem to maintain the fruits








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


on the plant. The occasional fruits are longitudinally
sulcate; it is probable that this cultivar was the base for
Schiede's V sylvestris, since the sulcate fruits are cited
in the protologue.
It is said that the cv. "oreja de burro" grows faster
and with more vigor than other cultivars of Vplanifolia;
furthermore the growers say that it is not severely
affected by fungal diseases. Some of the indicated
differences between "mansa" and "oreja de burro"
are statistically significant (Castillo and Engleman,
1993); however, the vegetative propagation of vanillas
in the plantations has made many of them genetically
uniform, and it is very probable that the observed
differences are among clones and not between different
taxa (species, subspecies or varieties) or indicative of
a genetic substructure in V planifolia. The "oreja de
burro" has a floral fragrance similar to the cv. "mansa"
and it is also visited by Euglossa viridissima.
Cuttings of "oreja de burro" are sold occasionally
by unscrupulous growers to discourage the
establishment of new plantations. We have observed
hectares of plantations that were nonproductive due to
large quantity of vines belonging to the cv. "oreja de
burro".
Several wild collected plants are notoriously
different from the specimens cited above, and it is
very probable that they belong to locally differentiated
populations. Among them, there are several specimens
collected in Belize that look somewhat different, with
small leaves, and flowers with a poorly-defined midlobe
of lip. These specimens have also rather elongate rachis
and the flowering time is from December to April.
Schipp S-971, M.A. Soto 8355, P.C. Standley 25064
are from Belize, Izabal, and Chiapas, from riparian
habitats and they seem to be wild; however, leaf
aspect is much more elliptic, and M.A. Soto 8355 has
more greenish flowers and much longer fruits. DNA
sequences of MAS 8355 is nested within the variation
of V planifolia, although with several autapomorphies,
but we do not know if they represent only sequencing
problems. Schlfiter et al. have shown that V. planifolia
has three rather different genetic groups, the cultivated
group, the wild specimens from Mexico to Quintana
Roo and the wild specimens from Costa Rica.
A specimen from Costa Rica [Esquinas forest.
Area between the Rio Esquinas and Palmar, sea level.
May 2, 1950. Flowers pale green. P.H. Alien 5532 SEL


(011362)! ] is somewhat strange, the leaves are clearly
petiolate and elliptic, acuminate, but not abruptly and
the inflorescence is very dense with bracts smaller
than typical. The flowers are not well-preserved, but
the lip does not seem lobed, and the warts on the apex
of the lip are very obscure. Our impression is that its
morphology is intermediate between V hartii and V
planifolia, and therefore it may be a hybrid.

OTHER RECORDS: MEXICO: VERACRUZ: Rancho El
Coscolin, El Escolin, Cerro Grande, Mpio. Papantla de
Olarte, 350 m s.n.m., 2029'N, 9733'W, clima calido
seco, suelo arcilloso-arenoso, plantaci6n de vanilla. Hierba
trepadora de 2 m, flor y fruto, abundante. Flores con la base
del perianto amarillenta. Fruto con la base adelgazada.
Aborta 50-100% de los frutos. vanillaa oreja de burro", 25
Abr 1989, R. CastilloM 211 & H. Cdlix de Dios AMO(x2)!
MEXU(678227)!

Other names applied in northern Veracruz, such as,
"negra", "verde" or "amarilla" are apparently due to
different exposure to light intensity.

OTHER RECORDS: UNITED STATES: FLORIDA: Dade Co.
growing on trees in an extensive area in Cutler Hammock, on
Biscayne Bay. Fls. greenish. May 15, 1976. D.S. Correll & J.
Popenoe 47216 NY! Brickell Hammock, Miami, Dade
County, J.K. Small, 19 Feb-March 22, 1915, NY! MEXICO:
SAN LUIS POTOSI: Tanjasnec, Mpio. San Antonio, bosque
tropical perennifolio, dooryard, vine, vanillala, planted, 4
May 1979, J.B. Alcom 3006 *LL! MEXU! PUEBLA:
Bejuco, fruto verde, vanillala, en cafetal, fecundada por
jicotes, Yancuictlalpan, Cuetzalan, el fruto seco se usa como
aromatizante y saborizante de atole y otros alimentos, F
Basurto y R. Patr6n 263 MEXU! VERACRUZ: Rancho El
Coscolin, El Escolin, Cerro Grande, Mpio. Papantla de
Olarte, 350 m s.n.m., 2029'N, 9733'W, clima calido seco,
suelo arcilloso-arenoso, plantaci6n de vanilla, hierba
trepadora de 2 m, floor verde amarillenta, abundante, vanillaa
mansa o fina", fruto alimenticio y para condimento, 25 Abr
1989, R. Castillo M. 210 & H. Cdlix de Dios AMO(x2)!
MEXU(678228)! [cultivated] Predio Escolin, 12 km al NE
de Papantla, 370 m s..n.m., trepadora, abundante, fruto verde,
selva alta, secundaria, vainillal, suelo rendzina de color
negruzco, 22 Junio 1987, A. Garcia 3226 MEXU! [cultivated]
Papantla de Olarte, 2027'N, 9719'W, vanillala; usos, para
hacer figures de vanilla, F Rosas C.I.P. 709 *XAL! [cf] San
Andres Tuxtla, El Vigia, al E de Cerro Blanco, 28 May 1967,
M. Peila 108 *AMES(11744, sterile)! Estaci6n de Biologia
Tropical de Los Tuxtlas, ca. 200 m s.n.m., selva alta
perennifolia, G.A. Salazar 2247 AMO(drawing)!
AMO(exsiccata, en FAA)! Volcano San Martin, on trees in
dense forest, 5 May 1936, 9511', 18035', 600 m, J. Gonzdlez

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


subE. Oestlund5790 *AMES(51851, sterile)! Mpio.Atoyac,
a un lado de la via del tren, cerca del puente, selva alta
subcaducifolia de Bernoullia flammea y Dendropanax
arboreous, terreno carstico, sobre piedras y arboles, 2 mayo
1995, ca. 400 m altitud,M. Soto 7745 *AMO! Vainillal del Sr.
Rafael Cardena, Diaz Mir6n y Rio Nautla, Papantla, Veracruz.
180 m s.n.m. Muestreada para fragancia floral. 31-III- 1995.
M. Soto 7648 AMO! [cf, fruit] A 800 m al norte de la casa de
Genaro, ejido Vista Hermosa, Mpio. Tezonapa, 18 40'N,
96 40'W, alt. 500 m, selva median subperennifolia, primaria,
suelo oscuro, arcilloso, bejuco perenne, escasa, fruto verde,
nom. vul. vanillala, 4-8-1986, R. Robles G. 977 *XAL!
[cultivated] Papantla, suelo poroso bien drenado con
abundante material organica asociada a Gliricidia sepium,
Erythrina berteroana, Bursera simaruba, bejuco, flor verde
amarillenta, nom. vul. vanillala, "caxixanath" (Totonaco),
17-02-1985, P. Reyes, N Carcamo & N. Garcilazo s.n.
*XAL! OAXACA: Vine 10 ft high in woods between Los
Llanos de Ozumazin and Rio Chiquito, District of Choapam.
Lat. 17 35', Long. 95 55', Alt. 700 ft., 1, May, 1939, R.E.
Schultes andB.P Reko 727 *AMES(9034)! Cultivada en los
vainillales de Santiago Tlatepuzco, Mpio. Usila, ca. 500 m
s.n.m., R. Medinilla s.n. AMO(en FAA)! Cerro Machete,
Distr. Pochutla, 800 m, Feb. 1941. B.P. Reko 6279
*AMES(60425)! orquidea epifita, flor verde con la base
blanquecina, acahual derivado de selva alta perennifolia, 3
km al W de La Esperanza, Mpio. Ixtlan de Juarez, Sierra
Norte, R. L6pez G. 2 AMO(16698)! same data E. Morales 1
AMO(16697)! Usila, Mpio. Usila, cultivada en los vainillales
de la familiar Bautista, cerca de la pista de aterrizaje, 90 m
s.n.m. 25 abril 1992,M. Soto 6672 &M. Herndndez *AMO(x
2; also in spirit)! mismos datos, aparentemente un clon con
olor a canela y labelo teiido de caf6, M. Soto 6673 & M.
Henmdndez AMO! mismos datos M. Soto 6674 & M.
Henmdndez AMO! Dto. Tuxtepec, Mpio. San Jose
Independencia, Cerro Clarin, en el extreme SW de la Presa
Temascal, ca. 120 m s.n.m., selva alta perennifolia, 1 mayo
1994. M. Soto 7630, U. Guzmin & Carlos *AMO! Institute
Tecnol6gico Agropecuario, San Bartolo Tuxtepec, plantaci6n
a cargo de Jesus Perez Meza. Abril 1996. MSoto 8040AMO!
Ejido La Gran Lucha, Mpio. Valle Nacional. Vainillal
particular de Nemesio Miguel Martinez. Plantaci6n
proveniente de un s61o clon que originalmente crecia en las
afueras de la comunidad. Plantaci6n de 1 ha, 2500 plants, de
3 anos de edad. Plantaci6n a 200 m s.n.m., en acahual
derivado de selva alta perennifolia en ladera karstica. 17-III-
1997. "Ligfiey" (Chinanteco). M. Soto 8117 & A. Cibridn
*AMO! La Cueva, cerca de Cerro Verde, km 25.3 del camino
Jalapa de Diaz-Usila, Oaxaca, selva perennifolia de montana,
780 m s.n.m., muy humeda, ca.18 57'N, 94 44'W. Plantaci6n
de vanilla de Jose Roldan (aparentemente hay tres clones en
la plantaci6n). 21-IV-1997. Tal vez el mismo clon que Soto
8477. Flor muy c6ncava que huele a canela. M. Soto 8483 &

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


M. Herndndez AMO! Mun. Santa Maria Jacatepec, predio El
Aguila, al O de San Agustin, entrando por la Reforma, 28 km
SO de Tuxtepec, carr. a Matias Romero. 1750'N, 96 06'W.
Elev. 550 m. Selva alta perennifolia. 19 Ene 1988. R. Torres
11027 & E. Martinez AMO(sterile)! MEXU(x3; sterile)!
TABASCO: cultivada en el Jardin Botanico de Puyacatengo,
en Teapa, fotografia! (Alderete y Capello, 1988); [cf, sterile]
Teapa, 0.34 km al E de la Universidad Agraria de Chapingo,
17 31'31"N, 92 55'33"W, selva alta perenifolia, 28 enero
2002, J. Cal6nico 21157 et al. MEXU! Poblado de Nacajuca,
20 m s.n.m, selva median perennifolia, primaria, suelo negro
arcilloso con abundante material organica, bejuco perenne, 3
m, escaso, nom. vul. "juju", uso comestible, 3-10-1978, R.
Ortega, W. MdrquezyB. Guerrero 858 *XAL! CHIAPAS: 17
km NO de Ocozocuautla, carretera nueva a la presa de Mal
Paso, selva alta perennifolia, ca. 900-1200 m, J Castillo 833
(specimen in cultivation)! Palenque, abril de 1892, Alt. 450
m, R.M. Comisi6n Geografico-Exploradora, Secci6n de
Historia Natural No. 7725 *AMES(21309)! 6 km al E de
Emilio Rabasa, 1 km al NE de La Reyna, Ocozocuautla,
16 55'N, 93 38'40", 720 m, alrededores de una casa, huerto
familiar y cultivos de maiz, S. Ochoa 3659 CHIP! [cf, fruit]
Reserva Ecol6gica "El Ocote", Mpio. Ocozocuautla, Alt. 750
m, selva alta perennifolia, primaria, suelo calizo, perturbada,
hierba perenne, abundante, fruto vaina, trepadora, 14-02-
1986, M. Palacios-Rios 2823 *XAL! Alrededores de la
Laguna Ocotalito, cerca de Monte Libano, selva baja
perennifolia, 950 m altitude, M. Soto 2836, 2840 AMO(slides,
in spirit)! Mpio. Ocosingo, Estaci6n de Biologia de Chajul,
vereda que conecta la vereda a La Granja con la de la Sabana
I, a unos 500 m del Rio Lacantim, selva perturbada con
bejucos y Bactris, terrenos ondulados y arenosos pero
ocasionalmente inundables, ca. 200 m s.n.m. 16o08' N,
90o53'O. 13-IV-1997.M. Soto 8355AMO(sterile)! Estaci6n
de Biologia de Chajul, monticulo 1, ruinas El Zapote, ca. 1
km de la estaci6n. 200 m s.n.m. 16o08'N, 90 53'. En acahual
de selva alta perennifolia con cania brava, muy j6ven, muy
escasa. 20 junio 1996. M Soto 7947 AMO(sterile)! Estaci6n
de Biologia de Chajul, junto al Rio Lacantun, acahual con
cana brava, muy j6ven, monticulo 1, Ruinas El Zapote;
aparentemente es el mismo clon que Soto 7947, sobre
Cecropia obtusifolia, muy escasa, vegetativamente igual a
Soto 8355, que es otro clon. Flor mas verde que las plants
cultivadas, hojas mas anchas no plans y fruto corto y trigono,
muy largo. Probablemente silvestre. 15 de abril de 2000, M.
Soto 9728, S. Maldonado, L. L6pez & P. Schluetter
*AMO(x3)! Selva del Ocote, km 44.7 de la carretera
Ocozocuautla-Malpaso, cerca de la presa, manch6n
perturbado de selva alta perennifolia con Poulsenia armata,
Ficus glabrata, Reinhardtia gracilis y Zamia splendens, muy
hfimeda, sobre areniscas, 275 m s.n.m., probablemente
silvestre. 21 abril de 2000, M. Soto 9922, S. Maldonado, P.
Schluetter, L. L6pez & M. Soto *AMO! Naha, Mpio. de








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Ocosingo; 27 km al SE de Palenque por la carretera fronteriza
hasta el Crucero Chancala, despu6s 55.6 km por el camino de
terraceria hacia Monte Libano, 1656'-17ol' y 9133'-
91038'; 900 m de altitude; bosque tropical perennifolio: Planta
trepadora de unos 4 m de alto con flores amarillas. 6 mayo
1999, A. Durdn Ferndndez 1166 MEXU! YUCATAN:
[sterile] "Vainilla" Yuc., "Sisbic" Maya, Cenote de Mayana,
Yucatan, 1866, A. Schott 215 F(276993)! In virgin xerophytic
forest, April 1916, forests of Xbac, G.F Gaumer & Sons
23352 AMES(*71360, 71369)! *F(446854)! MO(948146)!
NY! In virgin xerophytic forests, Izamal, April 1917, G.F
Gaumer & Sons 23909 *F(466398)! *US(1268044; sterile)!
[cf, poorly preserved] "Sisbic", June 4, 1899, G.F Gaumer
& sons F! Virgin forest near Chemax, on tree trunks, 29 Sep
1935, 8756', 2039', alt. 30 m, O. Nagel sub E. Oestlund
5050 *AMES(51846, fruit)! OUINTANA ROO: Isle of
Cozumel, Cenote Cedral, on tree trunks in mixed forest of
Achras zapota, 2 sep 1935, 8659', 2021', 5-10 m, alt., O.
Nagel sub E. Oestlund 5001 *AMES(51852, fruit)! Ejido
Hermenegildo Galeana, predios de Artemio y Constantino
Lopez Pascual, 1810'33.2"N, 8915'08.4"W selva alta
subperennifolia alternada. 10-V-2008. M. Soto 11370, L.
Ibarra& C. L6pezAMO! WITHOUT LOCALITY: "1815 de
Mexico, Epidendrum vanilla" [Sess6 and Mocifo collection]
Pav6n G! "Mexico, Mr Cowan G! BELIZE: Machaca,
forest shade, altitude 50 ft., height 35 ft., flower cream,
common, 16 May 1934, A. Schipp &-844 *AMES(40477)!
TOLEDO: [cf] "Vanilla". Vine, creamish colored flowers,
scented, in broken cohune ridge, Feeders Road, 14 Miles, San
Antonio-Punta Gorda Road, April 20, 1949, PH. Gentle 6721
*F(1599303)! *LL(x2)! NY! "Vianilla" Vine, flower
greenish-white, scented, in acahual, on hill slope beyond San
Antonio, January 30, 1952, P Gentle 7557 MEXU(511464)!
*LL(x2)! "Vainilla", Fls. pale green, in cohune ridge, Bolo
Camp, uper reach of Golden Stream, April 12, 1944, P.H.
Gentle 4521 *LL! "Vianilla", vine, flowers white, scented; in
cohune ridge, near Jacinto Hills. December 30, 1944, P.H.
Gentle 5106 *LL(x3)! Quite common along river bank here
in wet swampy places, still its difficult to secure flowering
material as the squirrels get them. Flowers greenish yellow.
Temash River. 100 ft alt., 30 ft, in. diam., 13th Mar 1935,
WA. SchippS-971 *AMES(43554)! Maskal Pine Ridge road,
high ridge, yellow flowers, 3 March 1934, [?C.L. Lundell &]
P.H. Gentle 1234AMES (*41360, *41361)! GUATEMALA:
BAJA VERAPAZ: Bis 6 m lang. Bl. dichfleischig. gellegriin.
Blt. mattschwefelgelle, sehr himfalling. Friichte. Blitht sehr
reich. Am Rio Polochic Oiber Tucurn, 900 m, Selten!
30.4.1882. EC. Lehmann 1436 G! ALTA VERAPAZ:
Flowers pale greenish; lip slightly finely fringed with slight
crests down middle. Vicinity of Finca Yalpemech, near Alta
Verapaz-Pet6n boundary line, alt. 100-120 m. March 24,
1942, J.A. i ..,... 45286*AMES(62981)! *F(1195546)!
PETEN: "Vianilla", vine, flowers green; Dolores, in low


forest about 2 km. 100 m south of village, May 3, 1961; E.
Contreras 2239 *LL! IZABAL: Large herbaceous vine, wet
thicket, frequent, vicinity of Puerto Barrios, at sea level, June
2-6, 1922, P.C. Standley 25064 *AMES(22674, fruit)! EL
SALVADOR: Finca Las Canoas, 700 m alt. terr., climbing
trees, mostly oaks, April llth, 1969, H.C. Clason sub F
Hamer 268 MO(3092269)! same data, near Ahuachapan,
April 9th, 1978H.C. ClasonsubEHamer 776*SEL(049262)!
"Vainilla", San Salvador, in garden, Vanilla seems to be
cultivated in Salvador only as a curiosity, 1922, S. Calder6n
518 *AMES(22663, sterile) *US (1151498; sterile)!
HONDURAS: MORAZAN: [cultivated] vanillala, racemes
15-20 greenish-yellowish flowers, fragrant, vine 8 m long or
more on Cupressus lusitanica in my yard, campus of EAP El
Zamorano, alt 800 m. March 28 1990, A. Molina 34346
*MO(4248893)! ATLANTIDA: [cf] "Vainilla". Wet forest.
Large herbaceous vine climbering over trees. Fls. pale green
throughout. Lancetilla Valley, near Tela, altitude 20 to 600 m,
Dec. 6. 1927, Mar. 20, 1928, P.C. Standley 55583
*AMES(36947, sterile)! F(581081)! Climbing on tree,
Lancetilla Valley, 159 ft alt., 8/12/1934, TG. Yuncker 4993
*AMES(105299, sterile)! COMAYAGUA: [cf, fruit]
"Vainilla", frutos muy aromaticos cuando secos. Planta
trepadora sobre el matorral en el bosque de montaniuela, alt.
1300 m, Marzo 31, 1963, A. Molina 11777A F(1620024)!
NICARAGUA: ZELAYA: Logging camp near Quebrada La
Talolinga. Elev. ca. 170 m. ca. 1151-52'N, 8426-27'W.
Tropical wet forest. Climbing vine fruit green, J.S. Miller &
J.C. Sandino 1138 MO! COSTA RICA: LIMON: "Vainilla".
Large epiphytic vine; leaves thick; fruit green; Hamburg
Finca, on Rio Reventaz6n below Cairo, altitude about 55
meters. Febr. 19, 1926, P.C. Standley & J. Valerio 48904
*AMES(32697, fruit)! *US(1309657)! in region Atlantica:
,,Hamburg", 9 km ab ore fluminis Reventaz6n, inter Musas
cultas, epiph! fl. viridi-lutei, altitude: 15 m, Die: 2.V.1930, G.
C,,ri...-F.i, 735 *AMES(37507)! W(1770)! Forest between
Punta Manzanillo and Punta Nona, E of Manzanillo de
Talamanca. Elev. ca. 8-12 m. 938'N, 8238'W. Vine
appressed-climbing in lower part, the fertile branches
pendent, festooning small tree in understory of open forest
atop coral bluffs along coast just E of Punta Manzanillo;
borne ca. 3-4 m above ground. Flowers whitish-green,
without fragrance (11:30 AM). Fruits green. Leaves thick,
subsucculent. Flowers and fruits, 2 May 1985, *M.H. Grayum
& G. Schatz 5257 CR(119858)! Pococi Canton. Refugio de
Vida Silvestre Barra del Colorado, Cerro del Tortuguero,
summit ridge. 1035'N 8332"W, ca. 100-119 m. Vining to
ca. 3.5 m above ground on small tree in secondary forest.
Leaves very thick and fleshy. Corolla light greenish, the
labellum longitudinally pleated internally and more or less
lacerate marginally. 22 January 1997, M.H. Grayum, B.
Hammel, J. Schipper & L. Merrill 11141 INB! Canton de
Talamanca, PN. Cahuita. Faja costefia de Limon. Sendero a

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


Puerto Vargas, 09044'20" N, 8249'30" W, 2 m. Planta
trepadora, creciendo a orilla de sendero. Frutos inmaduros
verde -claro y maduros verde amarillento. Comiin. 9
December 1993, E. Lpiz 87, L. Poveda & VH. Ramirez INB!
PUNTARENAS: Canton de Golfito. Valle de Coto Colorado.
Ribera del Rio Esquinas. Boca Rio Esquinas. 0844'00"N
8320'00"W, 30 m. Bejuco flores blancas, frutos verde-
amarillentos. 22 December 1993, M. Segura 264 & F
Quesada INB! CARTAGO: Pejivalle, Very common in forest
on ridges, seen also by riverside. No open fls found, May
1924, G.H. Lankester 847 AMES(28561)! PANAMA:
PANAMA: [sterile] Gross Lathrops trail, Barro Colorado
Island, June 26, 1931, L.H. Bailey & E. Zoe Bailey 374
AMES(37507)! [sterile] Wet forest; large herbaceous vine;
occasional, Barro Colorado Island in Gatiun Lake, altitude
120 meters or less, Nov. 18-24, 1925, P.C. \i..,,.k r 40897
*US(1251676)! Barro Colorado Island, Lutz Trail 300, vine,
much scandent, seldom branching, sterile, July 10, 1970, TB.
Croat 11177 SCZ(2261)! [cf] Wheeler Trail, Barro Colorado
Island, Oct. 26 1931, 0. Shattuck230AMES(71355,juvenile,
sterile)! [cf, sterile] Wooded swamp; large herbaceous vine.
Between France Field, Canal Zone, and Catival, Province of
Colon, January 9, 1924, P.C. \i.,,. 1 r 30160 US(1225400)!
SAN BLAS: Flowers pale green, April 28, 1933, G. Proctor
Cooper 287 AMES(*40380, *71354)! *F(771391)! NY!
*US(1557502)!
REFERENCES: Hooker, Bot. Mag. 117: t. 7167. 1891;
Rolfe, Journ. Linn. Soc. Bot. 32: 463. 1896; Duss,
Ann. Inst. Col. Marsaelle 3: 601. 1897; Cogniaux in
Urb. Symb. Antill. 6: 322. 1909; Ames & Correll,
Orchids of Guatemala, Fieldiana: Bot. 26(1): 57, fig.
16. 1952; Cabrera, T. 1999. Orquideas de Chiapas:
39; Luer, Nat. Orch. Florida 72. 1972; Garay & Sweet,
Fl. Less. Antill. p. 44, fig. 7. 1974; Hamer, Orq. El
Salvador 2: 372-373. 1974; Halle, Fl. Nov.-Caledonie
8: 407. 1977.

13. Vanilla pompona Schiede, Linnaea 4: 573. 1829.

TYPE: "Baynilla pompona Hispano-
Mexicanorum"; Mexico, Papantla et Colipa,
Schiede 1043, "Vanilla pompona nob. interim
Vaynilla pompona Papantlensis in sylvis Papantla,
Ja. 29" BM (lectotype; Garay Fl. Ecuador Orch.)!;
isolectotype W (sterile)!

Vanilla pompona is a variable, widely distributed
species. We consider it rather to be a species complex.
The specific name is based in a Mexican specimen
collected in central Veracruz. Mexican populations are
apparently isolated from the rest of the populations of

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


the species because there are no known populations in
Chiapas, Guatemala, Belize, or El Salvador. The taxon
reappears only in Central Honduras and Nicaragua.
This geographical isolation is correlated and probably
the reason that Mexican samples in the ITS tree form
a clade with high bootstrap support. The Mexican
plants also have slightly different morphology, since the
flowers and leaves are smaller, and the midlobe of the
lip deeply emarginate, but the general aspect of the vine
and flowers is very similar to the plants from Central
and South America.
Perhaps a more detailed analysis of the flower may
reveal more differences, but there is a notable lack of
well-preserved flowers in the Vpompona group.
The rest of the samples of V pompona have a
paraphyletic relationship with the Mexican samples,
but are more or less clustered into two groups. One of
the groups encompasses the samples from Honduras,
Nicaragua, Costa Rica and Panama. Differently
from the Mexican and the South American vines,
these samples sometimes have flowering shoots with
rather small leaves, that is, they keep the juvenile
stem morphology. Even though they do not form a
monophyletic group within the tree it seems better to
consider them another group within the species.
All the South American samples have commom
synapomorphic DNA characters, like peculiar
nucleotide bases in certain positions and common
deletions or insertions, although they also form a
paraphyletic group.
We have considered recognizing these three groups
in the Vpompona complex as different species, but the
topology of the ITS tree, with the Mexican samples
forming a monophyletic, derived group, and the other
two geographic groups being rather paraphyletic,
and the lack of constant, morphological traits have
led us to give them subspecific ranks. Therefore,
subspecies pompona, pittieri, and griiml/itlora are
here recognized.

13a. Vanilla pompona Schiede subspecies pompona

COMMON NAMES: vanillaa platanillo", "platanillo",
vanillaa pompona", "platano", vanillaa cimarrona",
"bania" (oaxaca), "nuguyu" or "nejuyu" (Oaxaca,
Zoque), vanillala, "litsmoya" (southern Veracruz;
?Popoluca); vanillaa gruesa" (Guerrero and S
Oaxaca).








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Hemiepiphytic, massive vine, little branched, leafy,
up to 16 m high. Stems terete, smooth, pale green dotted
with whitish, 10-24 mm thick; intemodes 11-14.5 cm
long. Aerial, free roots thick; terrestrial roots pilose, 4
mm thick; attaching roots conspicuously flattened on
the lower surface, whitish-greenish. Leaves subsessile;
blade variously shaped, ovate, widely elliptic, oblong-
falcate, base widely rounded to subcordate, apex acute
to subapiculate, coriaceous, very fleshy, brittle, pale
green, 22-29 x 8-13.7 cm, 3 mm thick. Inflorescence
7-20-flowered raceme, up to 200 mm long. Bracts
widely ovate, obtuse, rounded, concave to cymbiform,
apex somewhat conduplicate, ca. 8 x 10 mm. Flowers
successive, 1-2 open at once, ephemeral, tubular or
with spreading segments, 58-83 (69.679.96) mm long;
7.5-10 cm in diameter; pale yellow, lip orange yellow,
very showy; fragrance very strong, spicy, mint-like.
Ovary arcuate, oval-subtrigonous in cross section,
dorsiventrally flattened, arcuate, somewhat twisted,
very slightly grooved on one face, smooth, green, 37-45
(42.292.71) mm long, 5-7 (5.850.78) mm wide, 3-4
(3.290.36) mm thick. Dorsal sepal long oblanceolate
to spatulate, apex obtuse, rounded, subcalyptrate and
minutely warty on the apex of the outer surface; base
long unguiculate, long attenuate, canaliculate, the
claw ca. 17-30 mm long, 8 mm wide in the broadest
part, blade concave to somewhat conduplicate, 11-
16.5 (14.061.73) mm wide, ca. 14 veined, smooth,
thick and fleshy, total length 52-75 (66.376.42) mm.
Lateral sepals long oblanceolate, oblique, apex obtuse,
rounded, subcalyptrate and minutely warty on the
apex of the outer surface; base long unguiculate, long
attenuate, strongly canaliculate, the claw ca. 28 mm
long, 9 mm wide in the broadest part, blade concave, 13-
16 (14.871.13) mm wide, ca. 13-veined, smooth, thick
and fleshy, total length 56-75 (68.86.79) mm. Petals
oblanceolate-oblong, arcuate, slightly sigmoid and
oblique, the upper margin more straight, apex truncate-
rounded, base attenuate, basally slightly canaliculate,
blade flat to lightly concave, with an elevated, axial, flat
keel on the lower surface, ending in a triangular, acute,
flat process, 2 mm long; conspicuously canaliculate
at the apex of the outer surface, ca. 12-veined, 59-75
(67.835.58) x 11-17 (14.252.15) mm. Lip attached
to the column along the margins of the basal half 30-
40 mm (35,40,30,37,39) tubular, trumped-shaped,
cymbiform, deepest near the middle, axially grooved


on the lower surface; when spread out 60-80 (718.52)
x 29-40 (35.84.26) mm; unguiculate, the claw
canaliculate, with rows of trichomes, 22-30 (24.22.92)
x ca. 5 mm at apex, 3 mm at base; the blade flabellate to
very obscurely trilobed, margin undulate, ca. 40 veined,
apex deeply emarginate-bilobed; lateral lobes long
triangular-ovate, oblique, rounded, 30-39 (35.23.48)
x 10-15 (12.62.06) mm; midlobe inconspicuous, the
margins fairly reflexed, revolute, deeply bilobed, ca.
6-10 (82) mm x 13-22 (173.4) mm; penicillate callus
at 32-46 mm from the lip base, made up by ca. 10-14
congested, retrorse, approximately trapezoidal, laciniate
scales, the scales regularly united each to other along
the lateral margins, ca. 5.5-7 x 5-5.5 mm, continuous
with 3 thick keels that soon become fused in a cushion
like, claviform, rounded, rugose callus ending at the
very apex; 18-23 (20.61.85) x 3.5-4.5 mm. Column
elongate, conspicuously sigmoid, 43-59 (535.54) mm
long, 3.2 mm wide; ventral surface lanuginose on the
apical half, below the stigma; apex dilated, 4.5-6.5 mm
wide; vertical wings triangular-flabellate, the lower
margin acute, uncinate, ca. 2 x 4.5 mm. Stigma trilobed,
the lobes emergent, with a convex rostelum, ca. 3 x 4.5;
lateral lobes divergent each to other. Fruit variable,
usually arcuate, trigonous, thick, green, blackish when
rippen and strongly aromatic, 7.3-15 cm long, ca. 14
mm thick (n = 6). Fig. 13.
DISTRIBUTION: Mexico (Nayarit, Jalisco, Colima,
Michoacan, Guerrero, Oaxaca, Veracruz). Specimens
cited as V pompona from Guatemala and Belize, and
most from Honduras have turned out to be other taxa.
Vanilla pompona has a disjunct distribution. The
type came from Veracruz, Mexico and this group of
populations extends into N Oaxaca until the Tehuantepec
Istmus, but not reaching Chiapas. Subspecies pompona
occurs in W Mexico (Nayarit to S Oaxaca), where
characteristically small flowers whose segments do not
spread widely are very common.
In the ITS tree all the samples of V pompona from
Mexico (both from the Atlantic and Pacific watershed)
form a nested subclade with high bootstrap support
(97%). Another clade within V pompona includes
material cultivated in Guadaloupe, and from Suriname
and Guyanas, whose bootstrap value is moderate (70%).
The rest of the samples of the complex have unresolved
relationships at the base of the V pompona clade,
which has a 100% bootstrap value and is sister to V
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.




LANKESTERTANA


5mm


5cm


g::- 1v:


FIGURE 13. Vanilla pompona Schiede subsp. pompona. Based onM. Soto 7746. Drawing by M. Soto.
LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


z


b








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


calyculata, a clearly different species.
The distribution, molecular, and morphological
data suggest that subsp. pompona may be the result of a
relatively recent colonization of Mexico through a rather
long-dispersal event from Central American plants. We
believe that we can recognize at least two taxa in the
complex: V pompona subsp pompona and V pompona
subsp. r, uiiilitlorii (Lindl). Soto Arenas.
ECOLOGY: from sea level up to 1200 m, more common
at 300-900 m; in several types of tropical forest
(deciduous, subdeciduous, evergreen, lower mountain,
gallery forests, in savannas or warm pine-oak forest,
also in flooded areas) common in seasonally very
dry areas; this species is usually absent from the
wet forests, but occasionally can be located there in
limestone outcrops. Flowering period from April to
early June.

Vanilla pompona was described from sterile
specimens collected by Schiede near Papantla and
Colipa, in northern Veracruz, about 1829. The big plants,
withvery fleshy and thick stems bear huge, fleshy leaves,
very variable in form, but always big in flowering shoots
of Mexican plants. The large, deep yellow, and strongly
fragrant flowers are very showy when they spread their
segments (rarely in western Mexican populations); the
short, thick fruit is strongly aromatic and very much
appreciated by peasants, and also by vanilla conoisseurs.
The odor of the cured fruits is typically heliotropin-like
(Ehlers and Pfister, 1997). It is cultivated in small scale
in the V planifolia plantations, as a curiosity or charm,
rarely offered to the commerce. On the Pacific slope,
where it is known as "vanilla gruesa", is very common
in the coffee plantations, protected, but used only for
local consumption.
Vanilla pompona subsp. pompona is somewhat
variable; the specimens from the Pacific slope usually
have bigger leaves and thicker stems, smaller flowers,
and the perianth segments apparently never spreading;
vegetative features probably represent a plastic
response to drier conditions, and the flower size of
some specimens of southern Oaxaca approach those of
the Gulf watershed; the smallest-flowered specimens
are found in Nayarit and Jalisco.

OTHER RECORDS: MEXICO: NAYARIT: Camino de Ruiz a
Jesuis Maria, ca. 200 m altitude, cafetal derivado de selva
median subdecidua, abril 1986, G. Salazar & M. Soto
2009, 2010, 2011, 2013, 2017 AMO! El Caiman, 100 m


sobre la brecha hacia El Naranjo, que sale del km 10 del
camino El Cuarenteno El Cora, encinares calientes
mezclados con selva median subcaducifolia en las caiadas,
Quercus cf. conspersa, Byrsonima crassifolia, Bursera
acuminata, 490 m s.n.m., 21 26' 22.2" y 105 06'09.6".
Nos gui6 el Sr. Benito Decena Jimenez, de El Cora. 26/
junio/1998. M. Soto 8613 y E. Huerta AMO! M. Soto 8617
y E. Huerta AMO(x4)! JALISCO: La Playa (oeste de Los
Llanitos), Mpio de Puerto Vallarta, bosque subdeciduo,
trepadora, 550 m alt., flores amarillas, muy escasa, R.
Gonzdlez Tamayo 884 *AMO! *LL! Mpio. Puerto Vallarta,
entire Los Llanitos y Los Almacenes, Gonzdlez Tamayo s.n.
dibujo! Mpio. Cabo Corrientes. above El Tuito, road to
Mina de Cuale, R. McVaugh 26397 MICHOACAN: Mpio.
Aquila, San Pedro, Coalcoman, 550 m, 6-19-39, "Vanilla",
forest, vine, G.B. Hinton 13815 *AMES(57796)! NY!
*US(1805771)! Aquila, Tizupan, Coalcoman, 4-8-41,
"Vanilla", forest, G.B. Hinton 15910 *AMES(60655)! NY!
GUERRERO: Vallecitos, Montes de Oca, 9-13-1937,
"vanilla", on a tree, G.B. Hinton et al. 11387 K!
*US(2020421)! Sierra Madre, Guerrero- Michoacan, 400-
1000 m alt., Mars 1900, Pl. grimpante & attachant aux
troncs d'arbres- Recl6te des fruites d'Octobre a Novembre.
Endroits ombrag6s,"Vainilla", E. Langlasse 941
*AMES(71358)! G(x2)! K! *US(386288)!. OAXACA
(specimens from the Pacific slope): Pacific slopes of Sierra
Madre, near Copalita and below Pluma Hidalgo, climbing
on tree trunks, 26 Jul 1936, 500-1000 m, O. Nagel & J.
Gonzdlez sub E. Oestlund 6102 *AMES(51840)! km 190
Oaxaca-Pto. Angel. 14 mayo 1990. Flor prensada de
material en liquid 0. Sudrez 1153 AMO(15906; also in
spirit)! Sanguijuela, Mpio. Juquila, 1200 m s.n.m., 21 sept.
1992, V Meza & M. Tovar s.n. AMO(only fruit)! ca. San
Gabriel Mixtepec, selva median subcaducifola en caiadas
con encinar seco en las lomas. ca. 600 m s.n.m. 17 de abril
de 1992. M. Soto 6607, N. Perezy E. Martinez AMO! Cerca
de San Gabriel Mixtepec, cafetal en la zona de selva median
subcaducifolia y encinar caliente, 800 m s.n.m. 11 mayo
1994. Fotografiada, flores semicerradas. M. Soto 7632, A.
Ryan, E. Sandoval A. Rojas & A. Martinez *AMO!
OAXACA (specimens from the Gulf slope): Consoquilla,
/42, Liebmann 288 W(11761, drawing)! 1859, Cuming s.n.
G! Near Mogofi, climbing on trees, Isthmus ofTehuantepec,
fruit very fragrant, 9501' 170', 100 m, 18 Feb 1935, 0.
Nagel sub E. Oestlund 4574 *AMES(51841)! F(1257235)!
*MO(1145434)! Isthmus of Tehuantepec. Near Mogofe,
climbing on trees, 9501' 170', 100 m, 20 Apr 1935, H.
Knape sub Oestlund 4781 *AMES(51842)! F(1257236)!
*MO(1145435)! *SEL(011366)! *US(1809788)! San Juan
Guichicovi, vanillala, hierba, pachine seco, en la orilla del
camino, uso, con la fruta machacada, VIII-1986, Nereyda
Antonio & M. Heinrich UI 188 MEXU(431242)! Mpio.
Sta. Maria Chimalapa: Arroyo Huahuagtza, ca. 5.5 km al E
LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERIANA


de Sta. Maria; lat. 16 55' N, Long. 94 39' O, alt. 250 m;
acahual en area plana en orilla de arroyo (anteriormente
selva); suelo negro. Bejuco; fruto verde, came amarilla-
verdosa con semillas negras; comun; n.v. "bainia", "nuguyu"
(en zoque); se usa en forma de aceite para el cabello para
que de mas olor, 5 septiembre 1984, H. Herndndez G. 396
*AMO(8094, fruit)! CHAPA! Mpio. Sta. Ma. Chimalapa:
Arroyo Rancho Viejo, ca. 2-4 km al E de Sta. Maria; lat
16 54'N, long. 94 39'30"0; alt. 230 m; cafada con selva
(rodeada por encinar) con Poulsenia, Ficus, Brosimum etc.,
suelos color caf6 cascajoso. Epifita; fl. amarilla; "vainillo"
(en castellano), "nejuyu" (zoque); frecuente; se usa para
darle fragancia al cabello (se seca primero), 6 abril 1987, H.
Hernmndez 2418, AMO(16964)! 4 km al W de la frontera
Oaxaca-Chiapas, carretera a Tapanatepec, Dto. Juchitan,
epifita, flor amarilla, poco frecuente, veg. riparia, suelo
rocoso, 27 Marzo 1984 [botones], R. Torres 4807 & C.
Martinez MEXU! NY! Mpio. Santa Maria Guienagati, Loc.
Dto. Tehuantepec. 3 km al N de Santa Maria Guienagati,
carretera a Guevea de Humboldt. 16 43'N, 95 22'W; elev.
460 m, riparia con cafetal. Epifita, fruto verde, poco
frecuente, 27junio 1991,A. Campos 3721 MEXU(578082)!
ca. Valle Nacional, 1996, J. Prez Mesa s.n. AMO(in spirit)!
Terrenos comunales de San Agustin, Mpio. Sta. Ma.
Jacatepec; a unos 2 km al W de San Agustin, selva humeda
conManilkara, Brosimum, Pseudolmedia, con cafetal, a 350
m s.n.m. Planta silvestre de ca.18 m de alto, protegida, con
flores. 22-IV-1997.M. Soto 8495, M. Herndndez, F Sanchez
y M. Tensohua AMO! VERACRUZ: Near Zacuapan. On
trees, 12 feb 1932, 96 52' and 19 12' ca. 900 m, O. Nagel
sub E. Oestlund 2684 *AMES(41477, sterile)! From
Mexico. Secured by Prof. PH. Rolfs, in charge of the
Subtropical Laboratory, Miami, Fla., while traveling in
Mexico as agricultural explorer of the Office of S&PI in
April, May, and June, 1905. This species is a very strong-
growing vanilla. Produces the largest fruits and in
considerably quantity. Secured near Papantla, Person
interested: David Fairchild. P.I. No. 14440 SEL(011358;
photo)! Valle de Cordoba, 20 april 1865-6, Bourgeau 2332
K! Region of Zacuapan, in brush wood, half shady, ca. 800
m, Jun 1935, C.A. Purpus sub E. Oestlund 4852
*AMES(51839)! H. de la Higuera, 510 m s.n.m., vegetaci6n
riparia, bejuco perenne, abundancia regular, fruto aplanado
[sic], septiembre 8, 1966, M. Sousa 2721 *AMO(3443,
fruit)! MEXU(277445,296430,277427; x3)! MO(4273614)!
Caiada del Rio Guazuntlan, 380 m s.n.m., selva alta
perennifolia primaria, perenne, "litsmoya", colectada en
fruto. Usos: seca se pone en vaselina y se unta en el pelo,
septiembre 30, 1967, M. Sousa 3276 *AMO(8592, fruit)!
MEXU(fruit, 284554)! a 1 km al NE de Soteapan, 400 m
s.n.m., encinar, epifita, escasa, perenne, 30 Sept. 1967
[est6ril], M. Sousa 3272 MEXU(284553)! vanillala, La
Ceiba, Municipio de Puente Nacional, bosque de arboles
LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


various en canada, 100 m altitude, F Ventura 12355
*AMO(785, fruit)! MEXU(fruit, 235724)! Barranca de
Monterrey, entire el manantial y la cueva del abono, Mpio.
Axocuapan, alt. 380 m s.n.m., selvamediana subperennifolia,
bejuco, fruto verde, nom. vul. vanillaa cimarrona", trepador,
02-01-1987, M. Chdzaro, L. Robles & J Mdrquez 4376
*AMO(9026, fruit)! XAL(fruit)! El Remudadero, por la
carretera Conejos-Totutla, Mpio. Paso de Ovejas, (19 17'N,
96 33' W), selva median subcaducifolia primaria, suelo
arcilloso, pedregoso, en la barranca, bejuco perenne, 10 m,
escasa, fruto verde, nom. vulgar "platanillo", fruto
comestible, bejuco trepador, 01-11-1982, G. Castillo C.
2747 *XAL(fruit)! Miraflores, 9 km al NW de Atoyac,
Mpio. de Atoyac, 18 57'N, 96 49'W, alt. 900 m s.n.m.,
selva median subperennifolia, primaria, suelo carstico,
bejuco perenne, 8 m, flor amarilla, nom. vulgar vanillala,
18-05-1985, R. Acevedo & R. Acosta 195 *XAL! [cf, fruit]
Limites de Veracruz con Tabasco, Mpio. Coatzacoalcos,
18 03'N, 9606'W, selva baja inundable, primaria, suelo
negro pantanoso, hierba trepadora, perenne, 6 m, fruto verde
en vainas, nom. vulgar vanillala, 18-05-1980, J.I. Calzada
6096 F(1975502)! *XAL(fruit)! Alrededores de San
Fernando, Mpio. Soteapan, 18 18'N, 94 53'W, alt. 720 m
s.n.m., pinar-encinar, primaria, suelo rojo arcilloso, bejuco
perenne, 1 m, escaso, fruto verde, 18-12-1978, R. Ortega et
al. 1021 F(1963648, flower)! *XAL(fruit)! 1 km al S de
Palmillas, Mpio. Puente Nacional, 1913'N, 96 46'W, 600
m s.n.m., selva baja caducifolia, primaria, suelo delgado,
arcilloso, pedregoso, caf6, con poca material organica,
bejuco perenne, 4 m, fruto verde, 13-03-1985, G. Castillo &
M.E. Medina 4284 *XAL(fruit)! El Pochote, Mpio. Zentla,
hierba, floraci6n en mayo, N. Demeneghi G. 558
*XAL(fruit)! Mecayapan, region de Los Tuxtlas; acahual
derivado de encinar-pinar, 13/01/1998, S. Cruz Ramirez sub
M. Soto 8777(cultivated plant)! Miraflores, cerca deAtoyac,
ca.750 m s.n.m. Acahual de selva median perennifolia
submontana muy himeda. Protegida o cultivada. Planta
muy vigorosa, tallos 13-26 mm de diametro, hojas hasta de
30 x 12.5 cm, flores de 9 cm de largo, no abiertas, s6palos y
p6talos verdes exteriormente, parte internal amarillo-verde
lim6n, labelo amarillo yema cocida con algunas lines ocre,
garganta profunda con la parte apical amaillo-naranja,
aroma especioso, hierbabuena y algo mas. 29-IV-1995. M.
Soto 7746 & J Arguijo *AMO! El Angostillo, 34 km de
Huatusco, ca. 14 km de Manuel Gonzalez, Congregaci6n El
Pochote, La Reforma, Mpio.de Zentla, 500 m s.n.m.; selva
baja caducifolia en barrancas basalticas; aroma fuerte a
mental, visitada por Eulaema en la casa de Julio Arguijo.
30-IV-1995. M. Soto 7747, M.A. Flores, J. Arguijo, D.
Demeneghi, M. Demeneghi *AMO(x2)!
REFERENCES: McVaugh, Fl. Novo-Galiciana. 16:
Orchidaceae: 351-353,fig. 115, 1985.
13b. Vanilla pompona Schiede subsp. grandiflora








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


(Lindl.) Soto Arenas, comb. et stat. nov.

BASIONYM: Vanilla iiilnditlorai Lindl.,Gen. Sp.
Orchid. Pl. 435. (1840).
TYPE: Guyana Francesa, Martin s.n. holotypee,
K-L!).
Illustr. Dunsterville y Garay Venez. Orchid. Ill.
1: 435 (1050, as V pompona). N.v.: Palanda
vanilla, Ecuador: vainill6n.
Vanilla claviculata Duss non Sw.; V lutescens
Moq. ex Dupuis

COMMON NAMES: "vainill6n" (French and British
Antilles),

DISTRIBUTION: Apparently naturalized in the West
Indies.
There is no adequate material from NW South
America, but the specimens from the Amazon
basin, Guyanas, Coastal Venezuela, and S Brazil
are considered here to belong to V gidiilitloiri. It is
unknown if the plants from W Ecuador are Vpompona
or V grandifolia.
ECOLOGY: Lubinsky, Van Dam & Van Dam
(2006) report the pollination of V (pompona subsp.)
grindlitloir by male Eulaema meriana, one of the
largest of the euglossine bees. They also photographed
a male Eulaema cingulata gathering scents from the
mature fruit of this species. There has been at least one
other report of euglossine bees brushing on Vanilla
fruits (Madison, 1981). It is quite possible that the
Euglossini may play a role in the dispersal of Vanilla
seeds, as well as the pollination of the flowers.

Vanilla pompona is very similar to V. gunditlora
Lindl. ("French Guiana: without precise locality,
Martin s.n., K-L!), and the latter is has been recognized
as a synonym by most orchid taxonomists. However, V
griulitloir is different because it has a rather oblong
lip blade, not flabelate to obscurely trilobed as in V
pompona. In V grindulitlor the flowers are larger, with
the lip > 80 mm long, the callus 9-10 mm long and
originated at ca. 60 cm from the base and the column
is 60-75 mm long; also the apical thickening is less
conspicuous in V ,linditloiri and the flowers have
usually spreading segments, while V pompona is
usually closed, although occasionally, specimens with
spreading segments on sunny days are found, as the
illustrated in fig. 13.


Vanilla lutescens Moq.-Tand. described from
material of Paraguay [M. Coudert, cultivated at the
Faculty de Medecine de Paris, the type, not seen;
probable duplicate at W(35455)!] is also similar to V
,rundiitlorai and maybe conspecific with it.
REFERENCES: Chiron, G.R. & R. Bellone. 2005. Les
orchidees de Guyane Frangaise: 138, photo p. 93.

13c. Vanilla pompona Schiede subspecies pittieri
(Schltr.) Dressier, comb. et stat nov.
BASIONYM: Vanillapittieri Schltr., Fedde Rep. 3:
106. 1906.
TYPE: Costa Rica: In der Waldem an Ufern
des Rio Ceibo bei Buenos Aires, c. 200 m. M--
H. Pittier no. 6600, bliihend im Januar 1890,
B(destroyed); AMES(24329; drawing of the
holotype)! BR!

DISTRIBUTION: Honduras, Nicaragua, W Costa Rica
and the Pacific side of Panama.

The available Costa Rican material is not very
well-preserved, but seems to be more similar to
the Mexican populations of the Pacific slope; the
name V pittieri Schltr. is available to these Costa
Rican populations. The type of V pittieri was lost
in Berlin bombing; a sketch of the type made under
R. Schlechter's supervision, and a rather crude
illustration of the flower is kept at AMES; the lip is
shown as entire, with a very elliptic blade, as it had
been prepared from an immature bud. There are
additional specimens from the area of Buenos Aires,
Puntarenas, that match our concept of Vpompona.

OTHER RECORDS: NICARAGUA: CHONTALES: ca. 2.8 km
above (N of) Cuapa; ca. 12017'N, 8523'W, elev. 400-500
m; roadside, pastures, disturbed evergreen forest on hillside,
and bank of small stream. One seen, climbing up trunk of
large tree in disturbed forst near road, section of lower stem
collected, WD. Stevens, B.A. Krukoff 3690 *SEL(049330;
sterile)! SEGOVIA: OerstedW(19359)! Plant Large thick
(up to 2 cm) climbing leafy stems. Flowers- Large light
yellow sepals and petals, yellow-orange lip. Light yellow
column, orange on anterior face [mounted with description,
notes an analytical drawing] A.H. Heller s.n. *SEL(003850)!
NUEVA SEGOVIA: Quilali, ca. 1334?N, 8601'W, elev.
430 m, cultivated, said to have been collected wild in the
vicinity about 10 years earlier; flowers yellow-green, fragrant,
heavely visited by euglossine bees, fruits fleshy, dark brown,
aromatic, 16 March 1980, W.D. Stevens, B.A. Krukoff 16829

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*SEL(049331)! ZELAYA: Cerro Livico, 7 km northeast of
Siuna, forest slope; elev. 500 m. Climbing trunk oftree 28April
1978, D. Neill 3670 *SEL(047695; sterile)! COSTA RICA:
GUANACASTE: Flores amarillas. Enredadera subiendo
tronco de arbol, en chaparral o bosque poco denso dentro de
sabana cerrada. A 5 millas al sur de La Cruz, Liberia. Alt. 200
m, febrero 11, 1963, A. Jimenez M 313 *F(1606969)! 550
m, Calvo 1136 F(1456238)! Parque Nacional Guanacaste,
Estaci6n Maritza, sendero a la cima del Volcan Orosi. Bosque
primario y secundario, 1057.6'N, 8529.6'W, 600 m. Planta
epifita, creciendo sobre un arbol a 5 m de altura y a la orilla
de una quebrada. Frutos verdes. 2 julio 1989, INBio 132
INB! *CR(fruit)! PUNTARENAS: Canton de Buenos Aires.
Along the Rio Ceibo, Ujarras. 0914'00" N, 8318'00'W,
500 m. Thick stemmed vine; climbing high on trunk of tree
in riparian forest. Lvs. succulent, deflexed. Just one young
bud and one old, dry flower seen. 9 March 1993. M. Grayum
10237 INB! Esparza Macacona, margen derecha, quebrada
La Turbina. Bejuco trepador, botones florales verde tenue,
flor de corola amarilla, aromatica, 15 enero 1987, G. Herrera
& E. Herrera 466 MO(3594340)!WITHOUT PRECISE
LOCALITY: "Vanilla reichenbachii", Endres 270 W(16231,
16175, 16176)! Sepals and petals greenish yellow. Lip orange
yellow. Pie del Turrubares, 150 m, March 6, 1926 A. Alfaro
269 *AMES(31487, *31487)! In forest between Colonia
& Las Huacas, May 25 1903, O.F Cook & C.B. Doyle 742
*US(474659)! "Costa Rica", dried leaf fromM. Godefroy-L.,
Paris, 1895 K! PANAMA: PANAMA: Low places contiguous
to City, Alt Sea level. Grows in damp places. Climbs trees up
to 40-60. Flowers yellow, with orange throat. Flowers Feb-
March. 1919, C. W Powell 137AMES(*23763; AMES 28290
is V phaeantha)! Moist woods; large herbaceous vine, Rio
Paraiso, above East Paraiso, January 7, 1924, PtC. Standley
29919 *AMES(31437, sterile)! *US(1225393)! Vine on
west shore, Barro Colorado Island, March 11, 1931, C.L.
Wilson 119 *F(636191)! Barro Colorado Island, shores of
Gatum Lake, South of Lab., Aug 28th 1929, WN. Bangham
458 *AMES(71357, fruit)! Near Canal Zone, C.V Piper
s.n. *AMES(36948)! Low thick scrub along R. Tecumen,
north of Chepo road, up to 30 m. March 10, 1935, A.A.
Hunter andP.H. Allen 852 *AMES(42162)! *F(1329667)!
*US(1976167)! Coastal thicket; large herbaceous vine;
common, Punta Paitilla, January 12, 1924, PC. ,.i.,.,,,
30788 *US(1229496)! Parque Nacional Altos de Campana.
Epifita con botones florales verdes. Fuera del parque, a 1 km
de la oficina del parque. Sendero de interpretaci6n, 1 km al
este del campamento de los guardabosques de INRENARE.
Bosque huimedo tropical premontano. Elevaci6n entire 800 y
900 m. 840'N; 7955'W. 23 de abril de 1993, M.D. Correa,
E. Montenegro & E. Hidalgo 9460 PMA! Cerca de Cerro
Azul, 14.8 km al N de la carretera Cd. de Panama-Tocumen,
camino Cerro Azul-Cerro Jefe, vegetaci6n secundaria con
Terminalia y Byrsonima, ca. 600 m s.n.m., trepadora, est6ril,

LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


toda la poblaci6n con hojas elipticas, grandes y tallos gruesos,
4junio 2001,M. Soto 9921, G. Salazar J. LinaresyR. Gregg
PMA! in cult., at Smithsonian Tropical Research Institute in
Balboa. Voucher: fragrance collection. 18 Mar 1979, J.D.
Ackerman 1354 SEL(039338)! In Powell's garden; large
vine, Balboa, Non., 1923-Jan., 1924, P.C. Standley 28570
*US(1225365; sterile)! COCLE: ca. 5 km al N de El Cope,
entrada del Parque Nacional Omar Torrijos, vegetacion
secundaria abierta, derivada de bosque humedo tropical
premontano con Vochysia, ca. 700 m s.n.m. 2junio 2001, G.
SalazarJ. Linares, M. Soto yR. Gregg PMA!
REFERENCES: Hamer, Orch. Nicaragua, Ic. Pl. Trop. pl.
1195. 1984.

14. Vanilla sarapiquensis Soto Arenas, sp. nov.

TYPE: COSTA RICA: HEREDIA: Sarapiqui.
Llanura de San Carlos. Sardinal. Bosques residuales
y tacotales camino a Finca Dos Loas. 1031'45"N,
8405'32", 70 m. Bejuco herbaceo. Sepalos verde
crema, labelo blanco, column con la punta negra,
10 May 1995, holo. J.F Morales 4082, E. Lepiz,
VH. Ramirez & A. Rojas INB! iso. apparently at
MO.

Vanillae costaricense aemulens differt
inflorescentiis bracteisque minoribus, floribus
longioribus, labello suborbiculari, 3-lobato, callo
duobus carinis incrassatis prominentibus in apicem
confluentibus.

Hemiepiphytic vine, leafy. Stems 4 mm thick;
intemodes 6.5-7.8 cm long. Aerial roots flattened,
ca. 2 mm wide. Leaves obscurely petiolate, the
petiole canaliculate, ca. 7 mm long, 3.5 mm wide;
blade elliptic, long acuminate, somewhat cuneate at
base, membranaceous-chartaceous in dried condition,
15.9-19.5 x 4.8-7 cm. Inflorescence very different
from the vegetative shoots, a 4-flowered raceme 5.5
cm long, intemodes up to 1.5 cm long; peduncle ca.
26 mm, rachis ca. 2 mm thick. Bracts scale-like,
ovate to lanceolate, acuminate, concave, 9 x 4 mm.
Flowers successive, with rather spreading segments,
showy, ca. 7 cm in diameter; tepals green-cream
colored, lip white; of heavy substance. Ovary rather
sigmoid, terete, somewhat thickened at base, smooth,
inconspicuously sulcate, 42-53 mm long, 3-4 mm
thick. Dorsal sepal rather spreading, convex, the
margins reflexed, the apex slightly incurved, slightly
undulate; elliptic, apex acute-subacute, extreme apex








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Icm 4cm


10 cm










FIGURE 14. Vanilla sarapiquensis Soto Arenas, based on the type specimen, J.F Morales 4082, Drawing by M. Lopez.

IANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.








LANKESTERTANA


conduplicate-subcalyptrate, rounded; base obtuse-
rounded, ca. 10 veined, sparsely warty on the dorsal
surface, 44 x 17 mm. Lateral sepals basally concave,
convex at apex, the margins near the middle reflexed,
the apex recurved, widely elliptic, obtuse, base obtuse-
rounded, margins scarcely undulate, sparsely warty
on the dorsal surface, 11-veined, 41 x 20 mm. Petals
spreading, recurved at apex, slightly undulate on the
upper margin, oblong-elliptic, oblique, subacute, base
obtuse-rounded, 8-veined, smooth but axially sulcate
on dorsal surface, 45 x 16 mm. Lip attached to the
column 3 mm, ovate-suborbicular, obscurely trilobed
in outline, constricted at 2/3; 32 x 28 mm, the base
subcordate, slightly concave, with the lateral margins
erect forming a throat around the column; midlobe
concave, margins inflexed, incurved at apex, rounded,
8 x 16 mm; the lateral lobes semielliptic, the margins
minutely pleated, 21 x 10 mm; callus arising above the
middle, builted up by a pair of broad, elevated apically
confluent keels, the keels somewhat sulcate from the
botton to the crest, more elevated at distal part, widely
triangular in cross-section on their distal part, 18 mm
long, 5 mm wide. Column arcuate, semiterete, with a
dilated apex, smooth, 21 x 4 mm. Anther protruding,
semierect the distal part perpendicular to the column
body; the anther separated from the surface of the lip.
Fig. 14.

DISTRIBUTION: Known only from northern Costa Rica,
in the Llanura de San Carlos; it is to be expected from
adjacent Nicaragua.
ECOLOGY: Apparently in rainforest, at 70 m elevation.
It was collected with flowers in May.

This species of the membranaceous group is known
from a single collection from N Costa Rica. Vanilla
costaricensis, is known from the same geographic
area (Llanura de San Carlos), but V sarapiquensis is
distinct in its much shorter inflorescences with scale-
like bracts (V costaricensis has big bracts like those
of V inodora), larger flowers with much broader
tepals, suborbicular, trilobed lip, and especially by
the very different callus made up of two elevated,
thick, sulcate keels, confluent in the apical part. In
V costaricensis the lip is subentire, ovate-flabellate,
with the apex truncate or somewhat trilobed, but
the lateral lobes at the apex are subequal or slightly
longer than the mid lobe. In V costaricensis the callus
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


is made up of a pair of flat, broad keels along the axial
line, from the base to the beginning of the apical third,
then separated in 3 low, erect, congested keels ending
at the apex.
Vanilla mexicana has a distinctly trilobed lip, with
the midlobe more protruding, 3-keeled, with the keels
parallel. Also related is V martinezii with much larger
flowers, straight sepals, more rugose, basal keels,
although the short inflorescence, with reduced bracts
is similar.

15. Vanilla trigonocarpa Hoehne, Arq. Bot. Estado
Sao Paulo, nov. ser. 1(6): 126, t. 139. 1944.

TYPE: BRAZIL:PARA: Matas de terra fire,
Belem do Para, flores em 23-11-1926 e frutos em
7-10-1926,A. Ducke s.n.. holo., HEPF(19445)!
Vanilla pic ittorai Dressier, Orquideologia 13(3):
229-232. pl. 1979.
TYPE: PANAMA: Prov. Panama, carretera El
Llano-Carti, 10-15 km al norte de El Llano;
3 marzo 1976; s6palos verde palido, p6talos
crema verdoso, labelo blanco, lamina cafe-
amarillo, fragante, R.L. Dressier 5290, holo.
US(not seen).

Hemiepiphytic vine, leafy, up to at least 8 m high.
Stems somewhat fleshy, 7-10 mm thick; intemodes
4.8-7.5 cm long. Adventitious attaching roots
dorsiventrally compressed, at least up to 6 cm long,
ca. 4 mm wide; adventitious free roots subterete, ca.
3 cm long, 2 mm wide. Leaves petiolate, the petiole
1.5-2.5 cm long; blade elliptic or elliptic-oblong to
obovate, long acuminate, leathery, relatively thin, 15-
30 x 5.5-7 cm. Inflorescence usually on the lower
part of the stem, a 1-2(-4) flowered raceme, 13-15 mm
long. Bracts lanceolate, 15-20 mm long. Flowers with
spreading segments, sepals pale green, petals greenish
cream, lip cream, the blade yellow-brown with a white
margin, the throat with yellow-brown veins. Ovary ca.
4 cm long. Dorsal sepal lanceolate, obtuse, subclawed,
the claw ca. 40 mm long, 8 mm wide, 11-11.5 x 1.6-1.8
cm. Lateral sepals elliptic, subclawed, the claw ca.
40 mm long, 8 mm wide; 10.5-11 x 1.8-2 cm. Petals
narrowly oblanceolate, obtuse, with a prominent rib on
the outer surface, 10.8-11 x 1.6-1.8 cm. Lip attached to
the column along the margins of the basal half(ca. 6.5-
7 cm long), when flattened 11.5 x 5.5 cm; unguiculate,









SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


10cm
10 cm


5cm


FIGURE 15. Vanilla trigonocarpa Hoehne. Based on McPherson 11286 (plant, flower dissection), Dressier 4750 (fruit),
Collingram & Ugoccioni 912 (flower, lateral view) and Dressier s.n. published picture. Drawing by M. Lopez and M.
Soto.


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LANKESTERIANA


claw ca. 4 cm, then expanding abruptly and somewhat
campanulate, inflated blade; the blade entire, margin
strongly ruffled, ovate, ca. 11.5 cm long; penicillate
callus made up by several laciniate scales, 5 x 5 mm;
with a verruculose ridge running toward the apex; lip
apex strongly veined. Column elongate, ca. 80 mm
long, slightly arched, ventral surface pilose toward the
apex (for ca. 3 cm); with small wings. Stigma trilobed,
the lobes emergent, the mid-lobe the largest. Anther
ventral, 6 x 4 mm. Fruit cylindrical, 20-23 cm long,
ca. 1.8 cm wide; somewhat triangular in cross-section,
dehiscent; the seeds subspheric, somewhat flattened,
black and shiny, ca. 0.6 mm long. Fig. 15.

DISTRIBUTION: Costa Rica, Panama, and Brazil, and
probably in the intermediate areas.
ECOLOGY: From very wet rainforest, from 120 to 350
m altitude; it can be common in certain areas. Flowers
in Central America in January-February, and July;
perhaps all year. Hoehne (1945) indicates that the
flowers have a disagreeable fragrance, but that of the
fruits is pleasant. Dressler (pers. com.) has observed
males ofEuglossa asarophora pollinating this species
in Panama.

This species has been known in recent years
in Central America as Vanilla \l,,Iititorn Dressier,
but the Central American specimens are apparently
conspecific with the Brazilian plant earlier described
by Hoehne as V trigonocarpa.
Vanilla trigonocarpa is somewhat variable in
the form of the apical lobe of the lip, deeply retuse
in the type and truncate to slightly emarginate in the
Central American specimens. It is also variable in
the development of the lacerate margins. The type
specimen is minutely lacerate, as it is the specimen T.
Antonio 3645, however, other specimens are deeply
lacerate, as G. McPherson 11286.
Vanilla trigonocarpa is distinguished by the very
large flowers on very short racemes with only 1-4
flowers with lanceolate, long acuminate bracts; the
inflated lip, and strongly ruffled margins are also
characteristic. It is probably related to V espondae (and
perhaps to V sprucei), from Colombia, with flowers of
the same color, but with shorter and broader segments,
and the lip with an oblong-rounded midlobe covered
with digitiform papillae, up to 3.5 mm long. Molecular
analyses show that V trigonocarpa is the most basal

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.


species among the American penicillate Vanillas.

OTHER RECORDS: COSTA RICA: PUNTARENAS: at base
of Osa Peninsula, ca. 25 km E of Rinc6n along newly
improved road; elev 350 m, climbing on tree in mature
rainforest, 4 Jan 1986, P.M. Catling & V Brownell C24.1
AMES! PANAMA: PANAMA: El Llano-Carti road, 10.1
miles from highway, on trail to west; 9015'N, 7900'W; c.
350 m. Epiphytic on trunk; perianth green-white, the frilly
portion of lip brown, 16 July 1987, G. McPherson 11286
*MO(3499115)! about 10 km north of El Llano, 29 January
1974; vine, flowers borne from leafless stem, sepals and
petals greenish cream, lip cream, streaked with brown-
yellow within, center of limb brown-yellow, R.L. Dressier
4571 (PMA, SEL); El Llano-Carti highway, 10-12 km north
of El Llano, 12 September 1974; climbing vine, about 8 m,
R.L. Dressier 4750 PMA, SEL *US(2952908)! COCLE: On
the Atlantic side, ca. 5 hr walk from sawmill at El Cope.
Along slopes above Norte Rio Blanco near small village of
Caio Sucio; elev. 400-500 ft. Epiphyte vine on side of tree;
flowers a beautiful cream white; lip yellow very fragrant,
2 Feb 1980, T Antonio 3645 *MO(2928700)! WITHOUT
LOCALITY: leg.: Collector unknown. Selby Accession:
89-0281. Growing in Selby Display House. 26 April 1991.
Coll. Ingram & Ugoccioni 912 *SEL(065176)!
REFERENCES: Hoehne, Fl. Bras. 13(2): 28, t. 20. 1945.


Excluded species

Vanilla barbellata Rchb.f., Flora 48: 274. 1865.
A horticultural specimen grown in Austria,
supposedly from Cahuita, Costa Rica, seems to be the
West Indian V barbellata. There is a plate prepared by
the artists of the Royal Botanical Expedition to New
Spain, circa 1800 (Hunt Library; tracing at Library of
MEXU (as "112 Vanilla mexicana"), G, and tracings
probably elsewhere) that represents this species.
The flower has scale-like leaves and hirsute lip. The
Mexican origin of the plant is doubtful, since there are
no confirmed records of leafless vanillas in mainland
Mesoamerica. Many other paintings labeled as from
Mexico ("de Mexico") are actually Cuban plants,
prepared during the stay of the Expedition at the island.
As far as we know, Vanilla barbellata has been found
only in Bahamas, Florida, Cuba, Puerto Rico, and
theVirgin Islands.

Vanilla mexicana Miller, Gard. Dict. ed. 8: n. 1. 1768.

TYPE: Haiti, Plumier s.n., the holotype isPlumier's
original drawing at P; reproduction Plumier, Nov.







SOTO ARENAS & DRESSLER -A revision of Central American Vanilla

E~~,~ -


N A


FIGURE 16. A Vanilla cribbiana. B Vanilla dressleri. C Vanilla hartii. D Vanilla phaeantha. E, F Vanilla
planifolia.
IANKESTERIANA 9(3), January 2010. 0 Umversidad de Costa Rica, 2010.


AI









LANKESTERIANA


FIGURE 17. A Vanilla calyculata. B -Vanilla guianensis. C -Vanilla insignis. D -Vanilla phaeantha.

IANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


P1. Amer. Genera 25, pl. 28, 1703!
Epidendrum vanilla L., Sp. P1. ed. 1, 2: 952.
1753.
Lectotype (icon designated by Cribb, Taxon:
48(1): 47. 1999): "Volubilis siliquosa
plantaginis folio" in Catesby, Nat. Hist.
Carolina 2 (3): app. t. 7. 1747.
V vanilla (L.) Britton in Britton & Wilson, Bot.
Porto Rico 5: 185. 1924.
Syn: V aromatica L. (based on the same
specimen).
V aromatica Sw., Nov. Act. Soc. Sc. Upsal. 6:
66. 1799.
TYPE: Haiti, Plumier s.n., the holotype is the
original drawing by Plumier.
V anaromatica Griseb. Fl. Brit. W. Ind. 638. (?)
V epidendrum Mirbel, Hist. P1. ed. II. 9: 249.
1802-6.
V vanilla Karst. Deutsch Fl. ed. 2, 1: 474. 1895.
V vanilla Huth, Helios 11: 136. 1893.

Vanilla mexicana has been repeatedly cited from
Central America because the prevailing confusion
between this species and Vanilla inodora. In spite of its
name, it is a plant from the West Indies and northeastern
South America. It is recognized by the large flowers
with apple-green tepals and white, distinctly trilobed,
3-keeled lip. Other specimens labeled as V mexicana
from Costa Rica belong to the new species here
proposed as V costaricensis.

Vanillaplanifolia Jackson x Vpompona Schiede x V
planifolia Jackson

This artificial hybrid was created in Madagascar.
It was imported to Costa Rica, where is rather
common, and it is said that it is very resistant to
root rot disease (David Gardella, com. pers.). ITS
sequences are polymorphic and similar to both V
pompona subsp. gr riilitoir and to Vplanifolia. MatK
and rbcL sequences are grouped with V planifolia
sequences, which suggest that V planifolia was used
as the maternal progenitor, since the plastid genome is
inherited maternally.

Vanilla x tahitensis J.W.Moore, Bishop Mus. Bull.
Bot. 102: 25. 1933.

TYPE: Flora of the Society Islands, Island of


Raiatea, altitude 20 m; on trees, escaped from
cultivation; 3rd valley south of Faaroa Bay, 2-3
dm high. Native name Vanira tahiti, Nov. 11, 1923,
J. W Moore 294, holo. Bemice P Bishop Museum
Herbarium (559503)!
V hirsuta M.A. Clements & D.L. Jones,
Lasianthera 1(1): 47. 1996
TYPE: Papua New Guinea, M.A. Clements 6742
CANB(holo.) iso(NCBG) not seen, photo!
ADDITIONAL SPECIMENS: FRENCH POLYNESIA:
Raiatea: H. St. Johns 17310 K! Mongareva Is. H. St.
Johns 14459 K,'.lh., ii i,
Vanilla tiarei Constantin & bois, Compt. Rend.
1915 161: 202, nomem
TYPE: Tahiti.

DISTRIBUTION: Known from French Polynesia (Society
Islands), and New Guinea. The Tahitian vanilla
certainly is not known in Central America.

Vanilla tahitiensis J.W. Moore was described
from material from Raiatea, Society Islands. The type
corresponds to the plant widely cultivated in French
Polynesia and known as "Vanille de Tahiti", and the
second most important source of commercial vanilla. The
same taxon was also recently redescribed by Clements
& Jones (1996) as V hirsuta, based on a specimen
collected in Papua New Guinea (M.A. Clements 6742
CANB(holotype), NCBG(isotype) not seen, photos!).
There are also other different plants in Tahiti, including
Vplanifolia, and some variants of V tahitiensis that may
represent different cultivars or different species (e.g. a
plant labeled as cv. Tahita is apparently V bahiana).
Porteres (1954) indicated that the origin of V
tahitiensis was very enigmatic. It has been suggested
also that the distribution of V tahitiensis is far from
other vanillas, that its occurrence in Tahiti is perhaps
due to a human introduction, and that it is very similar
to Vplanifolia and not to the species from New Guinea,
which is the area with native vanillas closest to Tahiti,
such as V giulianettii F.M. Bayley, V kaniensis Schltr.,
V kempteriana Schltr., V ramificans (J.J. Sm.) J.J.
Sm., V wariensis Schltr.
Except for V tahitiensis, all the Vanilla species with
aromatic fruits are restricted to the Neotropics (Soto,
2003). Occasional reports of fragrant fruits in Asia (e.g.
Heyne, 1922) are probably based in wrong observations,
or in odors unlike the typical vanilla fragrance. Vanilla

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


lineages are confined to particular geographic areas, and
disjunctions are better explained by ancient vicariant
events than by long distance dispersal (Soto et al., in
prep).
Vanilla (in the generic sense) was introduced to
Tahiti from Manila about 1848. Petard (1986: 123)
claimed that Admiral Hamelin was responsible for
bringing the plants to the Botanic Garden at Papeete,
whereas Herman et al. (1989: 20) asserted that William
Ellis, a missionary from London Misionary Society,
brought cuttings from the Philippines (Smith et al.,
1992). Ryan (1986) agrees that the origin of V tahitiensis
is uncertain, she said that it may be that some Vanilla
planifolia plant stock was hybridized or that it mutated,
allegedly in the Philippines and that V tahitiensis is not
native of the South Pacific Islands, but she did not give
more data about this statement, although she indicates
Tahitian growers as the source of the information.
Morphological variation in Vanilla planifolia is
rather great, but it is clear that traits exhibited by
V tahitensis depart from the known variation of V
planifolia. It is difficult to think of a somatic mutation
than could cause such conspicuous differences. Sexual
reproduction of vanilla is almost impossible under
plantation conditions, and the life span is long enough
to discard rapid artificial selection as a probable source
of such "novelties". Therefore, the hybrid hypothesis is
more plausible.
The hybrid origin of V tahitiensis was suspected by
Porteres (1954) and Petard (1986), who suggested that it
arose from a cross between Vfragrans (= V planifolia)
and V pompona. Vanilla tahitiensis is morphologically
very similar to V planifolia and its allies, especially
V odorata and V insignis, but not to V pompona.
The hybrid between [V pompona x V planifolia] x
V planifolia has thicker stems, leaves, and different
flowers than V tahitensis.
Compared to V planifolia, V tahitensis has retrorse
warts on the midlobe of the lip. Since this trait is a
synapomorphy of the group of V odorata (including
V odorata, Vfimbriata, V helleri, and V insignis, and
also V cristagalli and V ribeiroi from South America),
if of hybrid origin one of these taxa is the candidate for
parental species in addition to V planifolia. V helleri
has a very short lip claw, and widely elliptic leaves,
broader than either in V planifolia or V tahitensis. V
insignis has conspicuously sulcate, papillose stems,

IANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


and leaves similar in shape to V planifolia. Since V
tahitensis has slender stems, narrower leaves, longer
perianth segments and column, the best candidates for a
parental species are V odorata and Vfimbriata (if this is
specifically distinct from V odorata), both with slender
stems, narrower leaves, longer perianth segments and
column, than Vplanifolia and V tahitensis.
Vanilla odorata is an important source of vanilla
fruits from the wild. Most vanilla beans collected by
peasants in Chiapas, Mexico are from V odorata, since
wild Vplanifolia is extremely rare or absent. Ames also
reported that it was the best quality species in Ecuador;
and herbarium records from Bolivia indicate that "it
was the best one there". Furthermore, V odorata has
some interesting features that make it a good candidate
in a Vanilla hybridization program, such as the higher
tolerance to more open habitats, its origin from places
with higher annual rainfall and more equatorial
climates, tolerant to occasionally flooded terrains, and
its resistance to the attack of"chinche roja" (Tentecoris
confusus) (M. Soto, pers. observations); nothing is
known about its resistance to fungal and bacterial
diseases, but it seems not as susceptible as V planifolia
(M. Soto, pers. obs.). Vanilla planifolia and V odorata
may be sympatric in several regions of Central America.
In Northern Oaxaca, Mexico, both species have been
found intermingled and flowering at the same time.
Both species are probably pollinated by Euglossine
bees (apparently Euglossa viridissima in the case of
V planifolia) involving a pollination system involving
deception. Fragrance analysis
Vanilla tahitiensis is the only Vanilla species
from which there are reports of diploid and tetraploid
chromosome counts. Tonier (1951) reported 64 somatic
chromosomes for "Vanilla haapape"(sic), considered to
be a variety of V tahitiensis (R. Porters, 1954), while
Heftimiu-Heim (1941) reported 2n=32 for V tahiti (sic)
and V papeno (sic), the later a common name of V
tahitiensis var. haapape (Porteres op. cit.).
ITS sequences of a specimen of V tahitensis
cultivated in France, and another gathered from the
Gene Bank as V hirsuta are nested into one of the clades
of V odorata with 100% of bootstrap support (other
specimens of V odorata have an ITS sequence more
similar to that of V helleri).
Although we do not have flowering material of
V tahitensis to make a quantitative analysis of the








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


intermediate morphology character of it between V
planifolia and V odorata, qualitative data support the
hybrid origin between these species. ITS sequences
indicate an extremely close relationship with V odorata.
However, Schluter found that was no evidence of a
hybrid origin analysing.
In conclusion, it is suggested that V tahitensis
is a taxon of hybrid origin between V odorata and
V planifolia introduced to Tahiti from Phillipines.
Lubinsky et al. (2008) provide strong support for
the hybrid origin of V tahitensis from crosses) of V
planifolia and V odorata. It is not improbable that
this cross was taken from Mexico to the Philippines
when both countries were ruled by the Spanish crown
and subject to a large commercial exchange thanks
to the Manila Galleon, that connected Acapulco with
the far east. The large variation observed (Maurice
Wong, pers, com.) and detected by RAPDs in the
tahitian vanilla is an additional point that supports
its hybrid origin. Lubinsky et al. (2008) have
suggested that V. xtahitensis can be an artificial
hybrid and even postulated a postclassic origin.
Orchid hybridization and subsequent germination
of the sclerotic seeds is almost impossible under
primitive conditions. Even now, Vanilla seeds are
planted from immature fruits in which the testa have
not become hard and it is thought that mature seeds
in ripe fruits must pass through the digestive tract of
a bat to germinate under natural conditions. Given
the sympatricity of V planifolia and V odorata,
the same flowering season, and the prevalence of
the similar herbal aroma with cinnamon-like notes
in both species, is not unlikely that they share the
same pollinator, apparently Euglossa viridissima
in V planifolia. Therefore, a spontaneous hybrid
between V planifolia and V odorata may be the
source of V x tahitenss, with a later selfing or back-
crossing with V planifolia, already in Tahiti, that
gave origin to the variation in cultivars, genotypes
and chromosomic numbers.
The following specimens are sterile, but their leaf
morphology is so different to the rest of the Central
American Vanilla, that we are convinced that they
represent a different, undescribed species. The leaves
are rather thin, but not as thin as those of subsect.
Membranacea; the leaves are conspicuously obovate
and abruptly mucronate.


COSTA RICA: CARTAGO: Sin flores ni frutos, Instituto
[Interamericano de Ciencias Agricolas], Turrialba, en el
bosque, 600 m alt., 23-VIII-49, J. LIon 1692 US(2021492)!
GUANACASTE: Large epiphytic vine in moist forest, El
Silencio, near Tilaran, altitude about 750 m, Jan. 13, 1926,
P.C. \'.,',.r & J. Valerio 44791 US(1309513)!

There are very few records on vanilla dispersal,
but bat dispersal had been suggested in V insignis
(M. Soto, unpublished data) and recently observed in
V pompona (N. Byrd, pers. com.). Bird dispersal is
expected in some Asian species, as V ahllnlitloir J.J.
Smith and V. ,-, iiilii Rchb. f., and also in the closely
related genera Cyrtosia.

ACKNOWLEDGMENTS. Thanks are due to Bush Boake Allen
Ltd. (BBA), London, for the financial support to develop this
work, especially to Drs. John Janes, Angela Ryan and Neil
C. Da Costa. Dr. Phillip J. Cribb, Royal Botanic Gardens,
Kew, encouraged us to conduct this work. The Comisi6n
Nacional para el Conocimiento y Uso de la Biodiversidad,
Mexico (CONABIO, Project J-101) financed part of the
analysis of nucleotidic sequences of Mexican taxa. We are in
debt to the keepers of the following herbaria, for permission
to study the collections in their charge: AMO, AMES, BM,
BR, CHAPA, CR, EAP, ENCB, F, G, GOEL, HEPF, INB,
K, LL, MA, MEXU, MO, NY, P, RJ, SCZ, SEL, UCR,
US, UV, W, WU, and XAL. Dr. Kenneth M. Cameron,
Wisconsin, for sharing his ITS sequences of Epistephium
and Lecanorchis. To the Union de Vainilleros de San Felipe
Usila, Oaxaca, to Ing. Heriberto Larios, for the facilities
to visit and work in plantations in Oaxaca and Veracruz,
Mexico. To Mariana Hernandez and Angelica Cibrian,
who assisted MAS in the field work. To Eric Hagsater who
placed at our disposal the facilities of AMO Herbarium.
To the Direcci6n General de Conservaci6n Ecol6gica,
Secretaria de Medio Ambiente, Recursos Naturales y
Pesca (now SEMARNAT), Mexico, for the permits for
collecting vanillas in Mexico, particularly in regions under
the National System of Natural Protected Areas. To the
staff of the Biosphere Reserve of Calakmul, Campeche,
particularly to Esteban Martinez, who made it possible
to study the populations of Vanilla insignis. Julio Arguijo
guided MAS in Central Veracruz to locate wild plants of
V planifolia and V pompona. To the staff of the Station
of Tropical Biology, Chajul, Chiapas, for the facilities to
work in the reserve. To Ulises Sanchez, who collected spirit
preserved material of the region Mazateca, in Oaxaca. To
Dr. Thomas Wendt and the National Geographic Society for
the logistics for the expedition to the region of Uxpanapa-
Sierra Tres Picos. To Dr. German Carnevali (CICY, Merida,
Mex.) and Joao Batista da Silva, from Belem, Brazil, who

LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rca, 2010.









LANKESTERIANA


gave us material from Venezuela and Brazil, extremely
useful to confirm the limits of the Central American
taxa. To Jose Linares, Escuela Agricola Panamericana,
El Zamorano, Honduras, for information and material of
V calyculata, and other species. To Febronio Tun Lopez,
for information on the vanillas of northern Guatemala. To
Prof William Stern, Florida State Museum for material of
V trigonocarpa. To Dr. Thomas Croat, Missouri Botanical
Garden, for access to material grown at the garden. To Mr.
Neal Byrd, Cia. Agricola La Gavilana Ltd., Costa Rica,
and Professor Franco Pupulin for invaluable material and
information on Costa Rican species. To M.Sc. Elvira Salas,
Universidad de Costa Rica, for her help with Costa Rican
materials. To Rolando Jimenez and Marco Lopez, who
prepared several illustrations. To Dr. Gerardo A. Salazar for
constant help, support and comments during the preparation
of this work. To Msc. Laura Marquez Valdelamar, Lab.
de Bologia Molecular, Instituto de Biologia, UNAM, by
sequencing labour. Finally, to our team in the Laboratorio
de Genetica Molecular y Evoluci6n Instituto de Ecologia,
UNAM, especially to Ms. Angeles Cortez, M.Sc. Rosalinda
Tapia, Drs. Alicia Gamboa, Caroline Burgeff, and Francisca
Acevedo for all their help in the laboratory.



LITERATURE CITED
Acufa, G.J. 1939. Catalogo descritivo de las orquideas
cubanas: 22.
Ames, O. 1934. An addition to the genus Vanilla. Bot. Mus.
Leafl. 2(8): 101-103.
Ames, O. and D.S. Correll. 1952. Orchids of Guatemala.
Fieldiana: Bot. 26: 54-60.
Bold6, B. & J. Estevez. 1990. Cubensis prima flora.
Fontqueria29: 160-161.
Bouriquet, G. 1954. Le vanillier et la vanille dans le monde.
en R. Port6res [ed.] Encyclopedie Biologique 46. Paul
Lechavalier. Paris.
Britton, N.L. & C.F. Millspaug. 1962. The Bahama flora:
83-84.
Bruman H. 1948. The culture history of mexican vanilla.
Hisp. Am. Hist. Rev. 28(3), 360-376.
Cameron, K.M. 2000. Gondwanan biogeography of
Vanilloideae (Orchidaceae). Southern Connections
Congress, Program and Abstracts: 25-26. Lincoln, New
Zealand.
Cameron, K. 2003. Vanilloideae. Pp. 281-285 In: Pridgeon,
A.M., P.J. Cribb, M.W. Chase & F.N. Rasmussen. (eds.),
Genera Orchidacearum vol. 3. Orchidoideae (Part two),
Vanilloideae. Oxford University Press. Oxford.
Camevali F.-C., G., J.L. Tapia M., R. Jimenez M., L.
Sanchez S., L. Ibarra G., I. M. Ramirez & M. P Gomez.
2001. Notes on the flora of the Yucatan Peninsula II:
A synopsis of the orchid flora of the Mexican Yucatan

LANKESTERIANA 9(3), January 2010. 0 Universidad de Costa Rica, 2010.


Peninsula and a tentative checklist of the orchidaceae of
the Yucatan Peninsula Biotic Province. Harv. Pap. Bot.
5(2): 383-466.
Castillo Martinez, R. & E.M. Engleman. 1993.
Caracterizaci6n de dos tipos de Vanilla planifolia. Acta
Bot. Mex. 25: 49-59.
Chase, M.W. 2001. The origin and biogeography of
Orchidaceae. pp. 1-5 In: A.M. Pridgeon, PJ. Cribb,
M.W. Chase & F.N. Rasmussen. (eds.), Genera
Orchidacearum vol. 2. Orchidoideae (Part one). Oxford
University Press. Oxford.
Chase, M.W. & H.G. Hills. 1991. Silica gel: An ideal
material for field preservation of leaf samples for DNA
studies. Taxon 40(2); 215-220.
Childers, N.F., H.R. Cibes & M.E., Hernandez. 1959.
Vanilla- The orchid of commerce. Pp. 477-508 in: C.L.
Withner (ed.), The Orchids. A scientific survey. Ronald
Press Co. New York.
Cibrian, J.A. 1999. Variaci6n gen6tica de Vanilla planifolia
en Mexico. Thesis. Faculty of Sciences, National
University of Mexico, Mexico City. 60 pp.
Correll, D.S. 1944. Vanilla: its history, cultivation and
importance. Lloydia 7, 236-264.
Correll, D.S. 1965. Supplement to orchids of Guatemala
and British Honduras. Fieldiana: Bot.: 31(7): 725.
Croat, T.B. 1984. Flora of Barro Colorado Island. Stanford
University Press.
Dix, M.A. & M.W. Dix. 2000. Orchids of Guatemala.
A revised annotated checklist. Monogr. Syst. Bot.
Missouri Bot. Gard. 54 p.
Dressier, R.L. 1979. Una Vanilla notable de Panama.
Orquideologia 13(3): 229-232.
Dressier, R.L. 1993. Field guide to the orchids of Costa Rica
and Panama. Comstock Publishing Associates. Ithaca
and London. Pp. 325-326.
Ehlers, D. & M. Pfister. 1997. Compounds of Vanillons
(Vanilla pompona Schiede). J. Essent. Oil Res. 9: 427-
431.
Fawcett, W. & B. Rendle. 1910. Orchidaceae. In: Flora of
Jamaica.
Foldats, E. 1969. Orchidaceae. In: Flora de Venezuela. vol.
15: 123-125, fig. 42.
Garay, L.A. & H.R. Sweet. 1974. Orchidaceae. In: Flora of
the Lesser Antilles.
Hagsater, E., M.A. Soto Arenas, G.A. Salazar Chavez, R.
Jimenez Machorro, M.A. Lopez Rosas & R.L. Dressier.
2005. Las orquideas de Mexico. Institute Chinoin A.C.
Mexico City.
Hamer, F. 1974. Las orquideas de El Salvador. Ministerio de
Educaci6n. San Salvador.
Hamer, F. 1984. Orchids of Nicaragua, part 5. Ic. P1. Trop.
fasc. 12. The Marie Selby Botanical Gardens, Sarasota,
Florida.








SOTO ARENAS & DRESSLER -A revision of Central American Vanilla


Heller, A.H. & A.D. Hawkes. 1966. Nicaraguan orchid
studies. Phytologia 14(1): 1-37.
Hermann, B., J.C. Celhay, M. Guerin, J.M. Maclet & J.
Rentier. 1989. Fleurs et plants de Tahiti. Les Editiones
du Pacifique, Sangapore.
Hoehne, F.C. 1945. Flora Brasilica XII, II. Orchidaceae.
(Texto e Ilustracao). Secretaria daAgricultura, Industria
e Com6rcio de Sao Paulo. Brasil. Tab. 20.
Kenny, J. 1988. Native orchids of the Eastern Caribbean.
London, Macmillan Caribbean.
Leon, F.S.C. & H. Alain. 1946. Flora de Cuba 1: 351.
Lubinsky, P, M. Van Dam & A. Van Dam. 2006. Pollination
of Vanilla and evolution in Orchidaceae. Lindleyana
[Orchids] 76: 926-929.
Lubinsky, P, K. M. Cameron;, M. C. Molina, M. Wolng, S.
Lepers-Andrzejewski, A. G6mez-Pompa, y SK. Seung-
Chul. 2008. Neotropical roots of a Polynesian spice: the
hybrid origin of Tahitian vanilla, Vanilla tahitensis
(Orchidaceae). Amer. J. Botany 95(8): 1040-1047.
Luer, C.L. 1972. The native orchids of Florida. The New
York Botanical Garden.
McLeish, I.M., N.R. Pearce & B.R. Adams. 1995. Native
orchids of Belize. A.A. Balkema. Rotterdam.
Murray, M.G. & W.F. Thompson. 1980. Rapid isolation of
high weight plant DNA. Nucl. Acids Res. 8: 4321-4325.
Petard, P. 1986. Quelques plants utiles de Polyn6sie
frangaise et Raau Tahiti. Editions Haere Po No Tahiti,
Pepeete.
PortBres, R. 1954. Le genre Vanilla et ses especes. In:
Lechevalier, P. (ed.). Le vanillier et la vanille dans le
monde. Paris.
Purseglove, J. W. 1975. Tropical crops: Monocotyledons. J.
Wiley, New York.
Purseglove, J. W., Brown, E. G., Green, C.L. & Robbins, S.
R .J. 1981. Spices, vol. 2. Longman, London.
Putz, F. E. & Holbrook, N. M. 1986. Notes on the natural
history of hemiepiphytes. Selbyana 9, 61-69.
Rain, P 1986. Vanilla cookbook. Celestial Arts. Berkeley.
Ray, T. S. 1990. Metamorphosis in the Araceae. Amer. J.
Bot. 77: 1599-1609.
Rodriguez C., R. L., D.E. Morta, M.E. Barahona, N.H.
Williams. 1986. G6neros de orquideas de Costa Rica:
70-71.
Rolfe, R.A. 1896. A revision of the genus Vanilla. J. Linn.
Soc. 32: 439-478.
Roullet, J. Fleur noire-Legende-Voyage & alea (Suite).
L'Orchidee 12:28.
Roubik, D.W. & J.D. Ackerman. 1987. Long-term ecology
of euglossine orchid-bees (Apidae: Euglossini) in
Panama. Oecologia (Berlin) 73: 321-333.
Savolainen, V, P Cu6noud, R. Spichiger, M.d.P Martinez,
M. CrBvecoeur & J.-F. Manen. 1995. The use of
herbarium specimens in DNA phylogenetics: evaluation


and improvement. P1. Syst. Evol. 197: 87-98.
Schlfiter, PM., M.A. Soto Arenas & S.A. Harris. 2007.
Genetic variation in Vanilla planifolia (Orchidaceae).
Econ. Bot. 61(4): 328-336
Schultes, R.E. 1960. Native orchids of Trinidad and Tobago:
39-40.
Smith, N. J. H., Williams, J. T., Plucknett, D. L. & Talbot.
J. P. 1992. Tropical forests and their crops, pp.357-364 .
Cornell University Press. Ithaca.
Soto Arenas, M.A. 1989. Listado actualizado de las
orquideas de Mexico. Orquidea (M6x.) 11: 233-277.
Soto Arenas, M.A. 1994. Vanilla odorata, una especie de
amplia distribuci6n. Orquidea (M6x.) 13(1-2): 295-300.
Soto Arenas, M.A. 1999. Filogeografia y recursos gen6ticos
de las vainillas de Mexico. Proyecto J-101. Mexico
City.
Soto Arenas, M.A. 2003. Vanilla (generic treatment). Pp.
321-334 in: A.M. Pridgeon, P.J. Cribb, M.W. Chase
& F.N. Rasmussen. (eds.), Genera Orchidacearum
vol. 3. Orchidoideae (Part two), Vanilloideae. Oxford
University Press. Oxford.
Soto Arenas, M.A. 2003. Vanilla insignis Ames. Pl. 700
in: E. Hagsater & M.A. Soto Arenas (eds.), Orchids of
Mexico, parts 2 and 3. Icones Orchidacearum fasc. 6
and 6. Herbario AMO. Mexico D.F.
Soto Arenas, M.A. & P.J. Cribb. 2010. A new infrageneric
classification and synopsis of the genus Vanilla Plum.
ex Mill. (Orchidaceae, Vanillinae). Lankesteriana 9(3):
355-398.
Soto Arenas, M.A. & R.L. Dressier. 2003. Vanilla. Pp. 583-
587 in: B.E. Hammel, M.H. Grayum, C. Herrera & N.
Zamora (eds.), Manual de plants de Costa Rica, vol.
3 Monocotiled6neas (Orchidaceae-Zingiberaceae).
Missouri Botanical Garden,-INBio-Museo Nacional de
Costa Rica.
Soto-Arenas, M. A. & A.R. Solano-G6mez. 2007. Ficha
tecnica de Vanilla planifolia. In: Soto-Arenas, M. A.
(ed). Informaci6n actualizada sobre las species de
orquideas del PROY-NOM-059-ECOL-2000. Institute
Chinoin A.C., Herbario de la Asociaci6n Mexicana de
Orquideologia A.C. Bases de datos SNIB-CONABIO.
Proyecto No. W029. Mexico. D.F..
Toledo, V M. 1982. Pleistocene changes of vegetation
in tropical Mexico. Pp. 93-111 in: G.T. Prance (ed.)
biological diversification in the tropics. Columbia
University Press, New York.
Wendt, T. 1989. Las selvas de Uxpanapa, Veracruz-Oaxaca.
Una evidencia de refugios floristicos cenozoicos.
Anales Inst. Biol. Univ. Nal. Auton. M6x. ser. Bot. 58:
29-54.
Wendt, T. 1993. Composition, floristic affinities, and origins
of the canopy tree flora of the Mexican Atlantic slope
rain forest. In: T.P Ramamoorthy, R. Bye, A. Lot and J.

LANKESTERIANA 9(3), January 2010. 0 Umversidad de Costa Rica, 2010.









LANKESTERIANA


Fa (eds.), Biological diversity of Mexico. Origins and Williams, N. H. 1982. The biology of orchids and euglossine
distribution. Oxford Univ. Press bees. Pp. 119-171 in: J. Arditti (ed.), Orchid Biology,
Williams, L.O. 1951. The orchidaceae of Mexico. Ceiba 2: Reviews and Perspectives, 2. Cornell University Press,
34-35. Ithaca, New York.
Williams, L.O. 1956. An enumeration of the Orchidaceae of Wood, J.J. 2003. Orchids of Borneo, vol. 4. Revised
Central America, British Honduras and Panama. Ceiba classification and selection of species. Sabah Society
5(1): 18. and Kew Publishing.


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LANKESTERIANA9(3) 355-398 2010


A NEW INFRAGENERIC CLASSIFICATION AND SYNOPSIS OF THE

GENUS VANILLA PLUM. EX MILL. (ORCHIDACEAE: VANILLINAE)


MIGUEL A. SOTO ARENASt1 & PHILLIP CRIBB2

1 Laboratorio de Gen6tica Molecular y Evoluci6n, Instituto de Ecologia,
Universidad NacionalAut6noma de Mexico and
Herbario AMO, Montafias Calizas 490, Mexico D.F. 11000, Mexico
2 The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, UK


ABSTRACT. An updated, annotated checklist of the species of the genus Vanilla is presented, with information
on nomenclature, distribution, literature and examined specimens. A key for the determination of the species
is included. A new proposal of infrageneric classification of Vanilla is provided.

RESUMEN. Se present un listado actualizado, comentado de las species del g6nero Vanilla, done se
proporciona information nomenclatural, distribucional, literature y especimenes examinados. Se incluye una
clave para la determinaci6n de las species. Finalmente se present una nueva propuesta de la clasificacion
infragenerica de Vanilla.

KEY WORDS /PALABRAS CLAVE: Orchidaceae, Vanilla, checklist, listado, infrageneric classification, clasificacion
infragenerica


The pantropical genus Vanilla Plum. ex Mill. belongs
to the subtribe Vanillinae and the subfamily Vanilleae,
it comprises some 106 species of hemiepiphytic vinous
orchids (Soto Arenas, 2003). Recent phylogenetic
analyses have shown that it is closely related to the
South American genera Dictyophyllaria Garay and
Epistephium Kunth and to the achlorophyllous genus
Lecanorchis Blume (Cameron, 1996; Cameron et al.,
1999, Soto Arenas & Alvarez-Buylla, in press).
Information on vanillas has been collected for a
phylogenetic analysis, but since no modem revision
exists, it has been necessary to evaluate the status
of all the proposed taxa. The results of this survey
are presented here, pending the production a formal
revision.
Vanilla has proved to be a taxonomically difficult
genus (Soto Arenas, 2003). The first species were
named before the establishment of the binomial
system of nomenclature (Linnaeus, 1753) and their
nomenclature and typification is confusing. Some
taxa were described from sterile material, and like
other hemiepiphytic plants, vegetative traits vary
greatly, rendering such characters difficult to use for
determination. A particular problem is that several
taxa have been described from immature buds, in


which measurements, and shape of the segments can
be substantially different from the morphology of
the same features at anthesis, making identification
uncertain. Furthermore, numerous species are known
from a surprisingly small number of specimens,
mostly due to the scarcity of available flowering
material because of their irregular flowering and
the ephemeral nature of the flowers, the positioning
of the inflorescences in the canopy of the rain
forests, and the low density of some populations.
Vanillas from densely populated areas in the tropics
apparently face severe conservation problems and
their populations are often depleted. Additional
complications are the poor preservation of the
flowers in herbarium specimens, the flowers often
rotting before they dry, and the presence of some
complexes in which interspecific differentiation
is subtle. A very large group of American species
is pollinated by male Euglossine bees attracted by
chemical compounds (Roubik and Ackerman, 1987;
Dressler 1993; Soto Arenas & Alvarez-Buylla,
pers. obs.). In other orchids with similar pollination
systems, there may there be little morphological
differentiation, yet the species can be effectively
separated by barriers imposed by the fragrance-








LANKESTERIANA


collecting behavior of their pollinators (van der Pijl
& Dodson 1966).
Our knowledge of Vanilla is therefore far from
adequate to produce a good, modem revision of
the group, although there have been important
contributions, such as two revisions of Vanilla (Rolfe,
1896; Porteres, 1954), the treatment by Klotzsch
(1845) and regional accounts by Hoehne (1945),
Correll (1951), Cribb & Hunt (1984), La Croix &
Cribb (1995), Luer (1972), Summerhayes (1968),
Seidenfaden (1992), and Szlatchetko and Olzewski
(1999). The revision by Porteres, the last full account
of the genus, although useful, is somewhat out-of-
date because, although we accept a similar number of
species in this checklist compared with his revision
(110 species), there are substantial differences.
The description of several new species, improved
collections of some poorly-known taxa clarifying their
status, and our understanding of the distribution of
the species and their phylogenetic relationships (Soto
Arenas, 2003; Soto Arenas et al., unpublished results)
have improved our understanding of the genus.

The fragmentary or poor material of some taxa
available to Porteres (1954) led him to propose
affinities between taxa that belong to different lineages
of the group. Therefore, the aims of this paper are to
present a new classification of the genus based on
cladistic analysis of morphological and molecular
data presented elsewhere (Soto Arenas, 2003; Soto
Arenas & Alvarez-Buylla, in press), to present an
updated checklist of the species and to stress where
more information is needed. This revision will also
allow the better identification of species and constitute
a base for future research.

Material and methods

A bibliographic revision was conducted.
Specimens were examined in the following herbaria:
AMES, AMO, BM, F, G, K, L, NY, P, W. Special
attention was paid to type and critical specimens, for
example flowering material from type localities where
the species were described from sterile or immature
material. When possible, we examined cultivated
and living wild specimens, spirit-preserved flowers,
botanical illustrations and photographs. Affinities
of many taxa have been assessed with the aid of
LANKESTERIANA9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


molecular data, published elsewhere (Soto Arenas,
2003; Soto Arenas et al., in press).

Infrageneric classification of Vanilla

The genus Vanilla is a rather diverse
assemblage of species of pantropical distribution.
Despite the lack of morphological synapomorphies
in the genus, there is strong molecular evidence that
it is monophyletic (Soto Arenas & Alvarez-Buylla,
in prep., Cameron, unpubl. data). The following
combination of characters are diagnostic for Vanilla:
the hemiepiphytic vinous monopodial growth
habit, with roots produced at each intemode (these
characters are found in some Galeolinae, a subtribe
that may be considered part of Vanillinae, and
Clemastepistephium N.Halle), fleshy fruits and the
wingless seeds with a hard seed coat (similar seeds
are apparently found in Dictyophyllaria).
Vanilla is an ancient group, and most morphological
groups within it are monophyletic and have defined
geographic distributions. In many ways it is a very
diverse genus compared with others in the Orchidaceae.
Some authors might favor the idea that the genus
should be split into more uniform assemblages with
a more restricted geographic distribution. The type
species, Vanilla mexicana, and most of Porteres' section
Membranacea could be easily considered as the genus
Vanilla with the rest of the species separated in a
different genus. Strong molecular and morphological
information supports this view, although it is equally
correct (in terms of recognizing monophyletic taxa) to
consider all of them in a single genus, although it has
few morphological synapomorphies.
The inner lineages of the genus may be supported
by molecular characters, but most of them lack
morphological synapomorphies that permit their
recognition and we found that their acceptance as
separate genera is not favoured by other authors. One of
the main reasons to keep Vanilla in its traditional sense
is to keep the traditional use of the name. The type
species, V mexicana, is in one of the two main clades,
and the species cultivated for their aromatic fruits fall
in the other one. To recognize both lineages as different
genera would produce the following problems:

SThe proposal of a new genus to accommodate
the economically most important species in the








SOTO ARENAS & CRIBB -Infrageneric classification and synopsis of Vanilla


Orchidaceae, to which the name "vanilla" or
vanillal" has been attached since pre-Linnean
times.
SA new lectotypification of the genus, whose
nomenclature has been historically complex, and
conservation of the name for the cultivated species.

We believe that to keep Vanilla in its traditional
sense is congruent with the phylogeny of the group
and to maintain the nomenclatural stability that has
now been reached.
Rolfe (1896) proposed a formal infrageneric
classification of Vanilla, recognizing two sections:
Foliosae and Aphyllae. Porteres (1954) accepted
Rolfe's sections and, in Foliosae, proposed
3 subsections: Papillosae, Lamellosae and
Membranacea. However, his subsections are invalid
as he provided them with neither a Latin diagnosis
nor a type. Furthermore, his sections and subsections
are polyphyletic, except for subsection Membranacea
which is paraphyletic (Soto Arenas, 2003).
Section Aphyllae is polyphyletic, because the
leafless habit evolved independently at least four times
in the genus, or maybe is the plesiomorphic condition
for the clade containing the Caribbean-Paleotropical
taxa, in which leafy species are also included. The
leafless condition evolved independently at least once
(V penicillata) in the clade of V planifolia. If sect.
Aphyllae is polyphyletic, then sect. Foliosae is also
polyphyletic. These sections are also hierarchically
incongruent, because they do not reveal the existence of
the two main clades in the genus: the membranaceous
species and all the others. Porters included both V
wrightii (= V bicolor) and Vpalmarum, the most basal
species in the clade containing the American non-
membranaceous leafy species, and V weberbaueriana
(= V ruiziana) and V pleei, to which he had no access
to appropriate material (see comments under these
species in the checklist) in subsect. Membranacea.
Subsections Papillosae and Lamellosae are very
heterogeneous and incongruent with our cladistic
analyses.

In order to be congruent with the phylogenetic
analyses, we propose Vanilla to be formed by two
subgenera: Vanilla and Xanata. As defined here
subgenus Xanata comprises two sections: Xanata and
Tethya.


Proposed classification of Vanilla

The following classification is proposed here.
Species groups are indicated in an informal way. They
are easily recognized morphologically, but there is
little evidence of their monophyly, and in some cases
they may well be paraphyletic.

A. Vanilla subgenus Vanilla

Vanilla subsect. Membranacea Porteres in Bouriquet,
Le Vanillier et La Vanille: 159 (1959). nom. illeg.

Plants with membranaceous leaves. Inflorescences
scarcely or not at all differentiated from vegetative axis.
Lip without a penicillate callus. Column united to the
lip only at the base. Column with a subperpendicular,
protruding anther (incumbent only 90), concave
stigma, and smooth ventral surface. Type species: V
mexicana Mill.

DISTRIBUTION: 15 species in the Neotropics, most
diverse in southern Brazil.
Subgenus Vanilla has never been used as a name
for this taxon, but it is automatically applicable to the
group of V mexicana Mill., the type species of the
genus.
The genus Dictyophyllaria is practically identical
in floral morphology to the V parvifolia group. Its
recollection and comparison with this subgenus and
other Vanillinae is much needed.

Two groups can be recognized, but their limits are not
clear-cut:

1. V mexicana group: Petals undulate; sepals usually
inrolled; lip very wide, 3- to 5-lobed, with a wide,
obtuse apical lobe and numerous high keels or an
elevated, very fleshy callus; and column lacking a
basal triangular keel on the abaxial surface.
Species placed here include Vanilla costaricensis,
V guianensis, V inodora, V martinezii, V methonica,
V mexicana, V oroana and V ovata.

2. Vparviflora group: Petals not undulate; sepals not
inrolled; lip 3-lobed, with a rounded-quadrate apical
lobe, a blade with axial rows of warts; and column
with a basal triangular keel on the abaxial surface.
Species included here are V angustipetala, V
parvifolia and V verrucosa. Similar to these, but
flowers larger with smooth keels, and an acuminate

IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


midlobe of the lip are V bertoniensis, V bradei, V
edwallii and V organensis.

B. Vanilla subgenus Xanata Soto Arenas & Cribb
subgen. nov. a section typical plants foliosis
vel aphyllis; inflorescentiis elongatis ab surculo
vegetato valde differt; callo penicellato labelli
deficienti; column ad labello medio adnata, latus
adaxialis glaber vel pubescens; anthera ventralis ad
column parallel; lobis stigmatibus emergentibus
distinguenda.

Typus: Vanilla planifolia G. Jackson.

Plants leafy or leafless. Leaves coriaceous-fleshy.
Inflorescence a specialized axis, very different from
vegetative ones. Lip with or lacking a penicillato
callus. Column united to the lip usually more than a
half of its length. Column with a parallel, somewhat
ventral anther, emergent stigmatic lobes, and with a
smooth or often hairy ventral surface.

DISTRIBUTION: pantropical.
DERIVATION OF THE NAME: from the Mexican Indian
Totonaco name for Vanilla, "xanat" or "shanat".
Totonaco Indians are thought to have brought Vanilla
planifolia to cultivation.

a. Vanilla subgen. Xanata sect. Xanata

3. Vanilla palmarum group: Ovaries calyculate;
flowers without a penicillate callus; lip rather entire
and united to the column only about a half of column
length, with longitudinal pubescent to hirsute lines on
the distal half of the lip.

DISTRIBUTION: West Indies and South America.
Species included here are V bicolor, V palmarum
and V savannarum.

4. V trigonocarpa group: Lip very inflated with
much crenulate margins; vegetative habit mesophytic;
racemes few-flowered; flowers very fragrant, small to
huge; fruit trigonous, very long and fleshy. Probably a
grade rather than a clade.

DISTRIBUTION: Tropical America, from Costa Rica to
Amazonia.
Species included here are V espondae, V hartii, V
spruce and V trigonocarpa.
IANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.


5. V planifolia group: Flowers greenish; lip concave
and with small papillae in the apical region. The South
American species have an apical callus connected
with the penicillate callus in the middle of the lip by
rows of papillae and with the lip apex usually recurved

DISTRIBUTION: Widespread in Tropical America,
including the West Indies.
Species included here are V appendiculata, V
bahiana, V cristagalli, V denticulata, V dubia, V
dungsii, Vfimbriata, V helleri, V insignis, V odorata,
Vphaeantha, Vplanifolia, V ribeiroi, V schwackeana,
V tahitiensis and V uncinata.

6. V penicillata group: Plants leafless, but related
to the American leafy, penicillate species. Petals
subunguiculate, lip long-unguiculate, with penicillate
callus, and villose apex. Fruit calyculate. Similar to the
leafy V ribeiroi and, except by the leafless condition,
belonging to the Vplanifolia group.

DISTRIBUTION: Guyana-Amazonia.
The only species included here is Vpenicillata.

7. V hostmanii group: Flowers large, usually white
with a yellow-orange lip; lip with thickened, radiating
veins on the distal half that can be somewhat warty
(the warts rounded and flattened); inflorescences with
rather distichous bracts; Ovary, sepals, and petals
granulosely keeled, very conspicuously so in some
species; Lip surface puberulent-cellular-papillose
in some species; perianth segments sometimes with
crystal druses included.

DISTRIBUTION: Widespread in Tropical America, except
for the West Indies.
Species included here are V cribbiana, V dressleri,
V gardneri, V hostmannii and V ruiziana.

8. V pompona group: Leaves very fleshy; stems
thick, xerophytic; lip rather smooth, except for the
penicillate callus, and with a slightly thickened axial
cushion running from the penicillate callus to the
apex; flowers strongly fragrant, yellow with an orange
lip; fruits thick, trigonous, banana-like.

DISTRIBUTION: Widespread in Tropical America, except
for the West Indies.
Species included here are V calyculata, V








SOTO ARENAS & CRIBB -Infrageneric classification and synopsis of Vanilla


chamissonis, V columbiana, V gr iilitiora. V
pompona, Vpseudopompona and V vellozii.

b. Vanilla subgen Xanata sect. Tethya Soto Arenas &
Cribb, sect. nov. a section Xanata foliis absentiis vel
praesentis, camosis vel coriaceis; ovario non calycoso;
floribus saepe labelli callo penicillato omatis; labello
manifeste vel obscure trilobato, ad column liberate,
in partim apicali lines pubescentibus destitute
distinguenda.

Typus: Vanilla phalaenopsis Rchb. f. ex van Houtte.

DERIVATION OF THE NAME: From the ancient Tethys Sea,
thought to be the migration route of Vanilla into the
Palaeotropics.

9. V kinabaluensis group: Leaves usually very broad,
elliptic; stems terete; inflorescences multi-flowered, in
some species with up to 150 flowers (V ahbimlitiloir
and V kinabaluensis); ovary stout, especially in New
Guinean species; fruit thick and relatively short; tepals
concolorous, greenish-yellowish, usually obtuse; lip
cuneate or cuneate-flabellate in some New Guinean
species, trumpet-shaped, not unguiculate, more or less
trilobed, with the midlobe often recurved and with
rows of reddish subulate papillae that may be very
long and flattened in V kinabaluensis, or whitish and
longer in V siamensis. Most species of this group are
seldom collected and many are imperfectly know, but
several of them seem to be closely related or maybe
conspecific. The group is much more developed in the
Malay Archipelago and New Guinea.

DISTRIBUTION: From S China to New Guinea, most
abundant in Indonesia.
Species included here are V albiunitlorii.
V giullianetii, V kaniensis, V kempteriana, V
kinabaluensis, V klabatiensis, V ovalis, Vplatyphylla,
V ramificans, V siamensis, V seranica, V sumatrana,
V. utteridgei and V wariensis.

10. V albida group: An Asiatic group with flowers
and plants similar in appearance to the American V
planifolia group; flowers greenish white; lip long-
unguiculate; tepals long; and lip with short apical
papillae. Some species, such as V moonii and V
yersiniana, and perhaps other have sulcate stems
in the way of V insignis. V albida, V andamanica,


V montana and V yersiniana are closely related,
differing in morphological details and distribution.
V havilandii seems to belong here, but has a very
different inflorescence, bearing very small flowers.
V moonii has buff-colored tepals and yellow lip with
apical papillae arranged in two rows, representing
perhaps a shared character with V annamica and V
somai.

DISTRIBUTION: Sri Lanka to SE Asia and the Malay
Archipelago across to Borneo.
Species included here are V albida, V andamanica,
V havilandii, V montana, V moonii, V sanjappae and
V yersiniana.

11. V annamica group: Inflorescences very distinct,
paniculate, bearing biflorous cymes; flowers trumpet-
shaped; and the lip has two apical, parallel rows of
papillae.

DISTRIBUTION: S China, Taiwan and Indochina.
Species included here are V annamica and V
somai.

12. V griffithii group: V palembanica is very
imperfectly known, but it seems to be allied to V
-, ir,l,, Leaves broadly elliptic; flowers with very
obtuse tepals, a lip with a deeply refuse midlobe,
woolly rather than scaly penicillate callus, and
truncate buds.

DISTRIBUTION: Sumatra, Borneo and peninsular
Malaysia.
Species included here are V -, ,iT dli and V
palembanica.

13. V borneensis group: Callus penicillate,
continuous with the pilose lip apex; plants with small,
narrow leaves and short inflorescences; lip with a pair
of corpuscles at base. They seem to be leafy versions
of the V aphylla-V calopogon-V wightii group.

DISTRIBUTION: From India and SE Asia to Indonesia.
Species included here are V borneensis and V
diabolica.

14. V imperialis group: Flowers large; lip trumpet-
shaped, similar to that of the V planifolia group but
shorter; callus absent; lip apex papillate.
IANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.








LANKESTERTANA


DISTRIBUTION: Continental Africa.
Species included here are V grandifolia, V imperialism,
V ochyrae and V polylepis.

15. Vfrangoisii group: Flowers very small, whitish
green with the lip rose-tinted at the base and with
purple margins; lip entire or six-lobed, densely
covered with fleshy hairs or papillae in middle and
front.

DISTRIBUTION: Madagascar.
Species included here are V frangoisii and V
coursii.

16. V chalotii group: Lip sacciform; midlobe reduced,
recurved; side lobes well developed.

DISTRIBUTION: West and equatorial Africa.
Species included here are V chalotii, V nigerica
and V seretii.

17. V africana group: Lip not sacciform; side lobes
of lip united to the basal 1/2-2/3 of the column and
forming deep sinuses with the mid-lobe; callus formed
by scales or warts; column apex exposed.

DISTRIBUTION: West and equatorial Africa, with a single
species in Eastern Africa.
Species included here are V acuminata, V africana,


V crenulata, V cucullata, V hallei, V heterolopha, V
ramosa and V zanzibarica.

18. V barbellata group: Flowers green to greenish
with a reddish-purplish lip; callus incipient, penicillate.

DISTRIBUTION: Caribbean.
Species included here are V bakeri, V barbellata,
V claviculata, V dilloniana and Vpoitaei.

19. V aphylla group: Lip with very long trichomes
and sometimes an incipient penicillate callus. Scarcely
different from V borneensis group, except for the lack
of leaves.

DISTRIBUTION: India to the Philippines.
Species included here are V aphylla, V calopogon
and V wightii.

20. V phalaenopsis group: Lip flabellate with two
rows of hairs besides the axial line; petals obtuse, very
wide.

DISTRIBUTION: Indian Ocean basin (East Africa,
Comores, Mascarene Islands, Madagascar, with a
single species in Sri Lanka and S India).
Species included here are Vanilla decaryana, V
humblotii, V madagascariensis, V phalaenopsis, V
perrieri, V roscheri and V walkeriae.


KEY TO VANILLA

1. Plants leafless, sometimes the new stems bearing small scars or short leaves (much shorter than the intemodes,
fugaceous and absent in older stems, bracts of the inflorescence ....................... .......................................................... Key I
1. Plants leafy, the blades well-developed and persisting several years, bracts persistent until the fruit ripens............ 2
2. Leaves soft, membranous, very thin in herbarium specimens; lip margins united to the column only at base for
less than 4 m m ; penicillate callus ab sent ....................................................................................... .................. ................................... K ey II
2. Leaves coriaceous, fleshy, often chartaceous in dry condition; lip margins united to the column more than 1/3 of
column length, usually for more than 8 mm; penicillate callus absent, wanting or often present ............................ 3
3. Lip deeply saccate at base .....................Key III................................................................................................................................................................ Key III
3 L ip n o t sa c c a te a t th e b a se ................................................................................................................................................................................................... 4
4. Stem 4 mm in diameter; leaves very small, 2 cm or less long, much shorter than the intemodes................................. 5
4. Stem stouter, 5 mm or more in diameter; leaves much larger, usually more or less as long as the intemodes or
lo n g e r .................................................................................................................................................................................................................................................... 6
5. Lip side lobes lacerate or deeply erose on front margin, the central callus on lip circular, densely papillate, the
m id-lob e of lip den sely p apillate ......................................................................................................................................................... V p en icillata
5. Lip crenulate towards apex, the disc densely hairy ......................................................................................................... Vfran oisii
6. Lip hexagonal in outline ........................................ coursi.............................................................................................................................................. V coursii


LANKESTERIANA 9(3), January 2010. 0 Unversidad de Costa Rica, 2010.








SOTO ARENAS & CRIBB -Infrageneric classification and synopsis of Vanilla


6 L ip e n tire to 3 -lo b e d ............................................................................................................................................................................................................... 7
7. Lip 3-lobed in basal half ......................Key ................................................................................................................................................................... Key IVKey
7 L ip en tire o r 3 -lo b e d in ap ic a l h a lf ............................................................................................................................................................................. 8
8. Inflorescence paniculate, each branch a 2-flowered cyme ................................................................................. ........................... 9
8. Inflorescence racemose or, if branched, then branches not 2-flowered ................................................... ............................. 11
9. L ip hexagonal ........................................................................................................................................................ ................................ coursii
9 L ip n o t h e x a g o n a l .................................................................................................................................................................................................................. 1 0
10. Lip entire, broadest in middle, callus densely papillose, running from base of lip almost to tip ..................V annamica
10. Lip 3-lobed, broadest across the apex; callus on disc, elongate-elliptic, papillose, with a few scattered papillae
on the midlobe ............................................... somai
11. Lip disc lacking a central almost quadrate tuft of erose or lacerate callus; callus papillose, hairy or of several
erose or papillose ridges running the length of the lip..................................................................................................... Key V
11. L ip w ith a central tu fted or con vex callu s ....................................................... ........ ............... ................................................................ 12
12. Lip cuneate or cuneate-flabellate, lacking a prominently narrowed claw when spread flat, usually 1.5 times as
long as broad or less; species from O ld W world ................................. ...... ........ ............................................................................. K ey V I
12. Lip with a prominent narrowed claw when spread; lip usually more than 1.5 times as long as broad ............Key VII


KEY I

1. Inflorescences subcorymbose, very long; bracts deciduous; flowers white, yellow or greenish with pink or
maroon blotches deep in the throat of the lip; petals much broader than sepals; penicillate callus absent, but with
rows of longitudinal papillae; species from the Indian Ocean basin: E Africa, Madagascar, Sri-Lanka, India,
Seychelles, Comoros ......................................................... 2
1. Inflorescences racemose; flowers greenish-white, often flushed or blotched with purple; lip with a penicillate
callus and other trichomes towards the apex; species from America ................................................................................................ 8
2. Column of 14-17 mm long; sepals 25-30 mm long ................................................... ...................... V decaryana
2. Column 18-30 mm long; sepals 50-70 mm long ..................... ........ ................... .......... 3
3. Flowers white with a yellow throat; veins of the petals and lip thin, almost straight ..................................... walkerae
3. Flowers white with an apricot or pink throat or yellow with orange or maroon markings in the throat of the lip;
veins of the petals and lip partially or totally sinuous-undulate ............................................ ..................... ................................... 4
4. Column 27-30 mm; lip covered i ili i. ._ scattered hairs in the middle, veins slightly flexuous in the petals and
the upper half of the lip; flowers yellow with the throat of the lip maroon or orange .............. ....... ............. 5
4. Column 18-23 mm; disc of the lip adorned with 2 lines of long papillae or fleshy hairs; veins of the petals and
lip all very sinuous-flexuous; flowers white, the lip a pink-reddish blotch on the throat............................................. 6
5. L ip papillose; column n 20 m m long ...................................................................................................................................... V hum blotii
5. L ip not papillose; colum n 27-30 m m long ......................................................................................................................................... V p errieri
6. Sepals 75-80 m m long; stem 12-20 m m in diam et ...................................................................................................................... V roscheri
6 Sepals 50-65 m m long; stem 8-10 m m in diam ................................................................................................................................................ 7
7. L ip oblong, acute, the m argins undulate ................................................................................................................................ V phalaenop sis
7. Lip obovate, obtuse, the margins not undulate ............................................ ............................ ..................... madagascariensis
8. Stems slender, usually less than 5 mm in diam. when mature in living condition; penicillate callus well-defined;
column fused to the lip margins until stigmatic region; margins of the obscure lateral lobes of the lip lacerate-
fimbriate, lip more than 50 mm long; Amazonian plants ............................ .............................................. V penicillata
8. Stems thick, about 1 cm or more in diam. when mature in living condition; lip less than 50 mm long; lip apical
margins crenulate, undulate or entire, not lacerate-fimbriate; Caribbean plants ......................... ......... ................ 9
9. Leaf present at each internode, persistent, to half or more the length of the internode, leaves plane, apices
hooked; lip green with maroon margin and veins, hairs yellow, margins irregular ........................................ V poitaei


LANKESTERIANA 9(3), January 2010. 0 Universdad de Costa Rica, 2010.




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This issue of Lankesteriana is dedicated to the memory of the great Mexican botanist and orchidologist, MIGUEL NGEL SOTO ARENAS (1963), tragically murdered in Mexico on August 27th, 2009

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INTERNATIONAL JOURNAL ON ORCHIDOLOGYISSN 1409-3871VOL. 9, No. 3 JANUARY 2010Miguel ngel Soto Arenas (1963-2009) EDUARDO A. PREZ GARCA & ERIC HGSATER In memoriam: Miguel ngel Soto Arenas (1963-2009) ERIC HGSATER Miguel ngel Soto Arenas: publications and main conferences (1983-2009) A new species of Vanilla from South America MIGUEL A. SOTO ARENAS A revision of the mexican and Central American species of Vanilla Plum. ex Mill. with a characterization of their ITS region of the nuclear ribosomal DNA MIGUEL A. SOTO ARENAS & ROBERT L. DRESSLER Vanilla Plum. ex Mill. (Orchidaceae, Vanillinae) MIGUEL A. SOTO ARENAS & PHILLIP CRIBB Comparettia sotoana (Orchidaceae: Oncidiinae), a new Ecuadorian species FRANCO PUPULIN & GILBERTO MERINO Confusion in Epidendrum brenesii Schltr., and a new Costa Rican species: Epidendrum sotoanum (Orchidaceae: Laeliinae) ADAM P. KARREMANS & ERIC HGSATER Oncidium ornithorhynchum, para una vieja especie: Oncidium sotoanum (Orchidaceae) ROLANDO JIMNEZ MACHORRO & ERIC HGSATER Two new species of Lepanthes from Costa Rica close to L. schizocardia (Orchidaceae: Pleurothallidinae) FRANCO PUPULIN, DIEGO BOGARN & CHRISTINA M. SMITH Stanhopeinae Mesoamericanae, V. Las Stanhopea de Mxico GNTER GERLACH 269continues273 276 285 355 399 403 411 423 431 281

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Crossoglossa sotoana (Orchidaceae: Malaxideae), a new species honoring the Mexican botanist, Miguel ngel Soto Arenas FRANCO PUPULIN & ADAM P. KARREMANS Dos especies nuevas de Pleurothallidinae (Orchidaceae) para Mxico RODOLFO SOLANO G”MEZ Masdevallia sotoana (Orchidaceae: Pleurothallidinae), a new species from Ecuador HUGO MEDINA & FRANCO PUPULIN New species of Porroglossum (Orchidaceae: Pleurothallidinae) from Ecuador GILBERTO MERINO, ALFONSO DOUCETTE & FRANCO PUPULIN Myoxanthus scandens (Orchidaceae: Pleurothallidinae), with a new species FRANCO PUPULIN, DIEGO BOGARN & MELANIA FERNNDEZ Some new Sobralieae from Costa Rica and Panama ROBERT L. DRESSLER & DIEGO BOGARN Lepanthes arenasiana (Pleurothallidinae: Orchidaceae), a new species from Costa Rica DIEGO BOGARN & MELANIA FERNNDEZ Sotoa, a new genus of Spiranthinae (Orchidaceae) from Mexico and southern United States GERARDO A. SALAZAR & CLAUDIA BALLESTEROS-BARRERA A new species of Odontoglossum (Orchidaceae: Oncidiinae) from Ecuador STIG DALSTRM & GILBERTO MERINO A New Ornithidium (Orchidaceae: Maxillariinae) from the Massif de la Hotte of Haiti JAMES D. ACKERMAN & W. MARK WHITTEN GUSTAVO A. ROMERO-GONZLEZ, GERMN CARNEVALI FERNNDEZ-CONCHA & PAUL ORMEROD A new species of Lophiaris of Mexico RICARDO BALAM NARVEZ, WILLIAM CETZAL IX & GERMN CARNEVALI FERNNDEZ-CONCHA A la tercera se gana: the validation of Benzingia (Orchidaceae: GUSTAVO A. ROMERO-GONZLEZ & CALA W AY H. DODSON LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.ii LANKESTERIANA443 447 455 459 467 475 487 491 505 509 513 521 526

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Epidendrum jalcanse (Orchidaceae), a new species from Northern Peru MIGUEL CHOCCE, STIG DALSTRM, ERIC HGSATER & JORGE ARNAIZ Anlisis de la seleccin de sustrato por parte de Dendrophylax lindenii (Lindl.) Benth. & Rolfe (Orchidaceae) en Cabo San Antonio, Pennsula de Guanahacabibes, Pinar del Ro, Cuba ERNESTO MJICA, JOSEP RAVENT”S & ELAINE GONZLEZ estado Monagas, Venezuela CARLOS LEOP ARDI El redescubrimiento de Mexipedium xerophyticum (Soto Arenas, Salazar & Hgsater) V.A. Albert & M.W. Chase EDUARDO A. PREZ-GARCA Book Review SAMUEL SPRUNGER SAMUEL SPRUNGER Reviewers of the manuscripts submitted to Lankesteriana, vol. 8 LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. INDEX iii529 533 541 557 564 566 568 567

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LANKESTERIANAINTERNATIONAL JOURNAL ON ORCHIDOLOGY Copyright 2010 Lankester Botanical Garden, University of Costa Rica Effective publication date: January 21, 2010 Layout: Jardn Botnico Lankester. Cover: Vanilla helleri A.D.Hawkes. Drawing by M. Lpez and M.. Soto Arenas. Printer: Litografa Ediciones Sanabria S.A. Printed copies: 500 Printed in Costa Rica / Impreso en Costa RicaR Lankesterian a / International Journal on Orchidology No. 1 (2001)-. -San Jos, Costa Rica: Editorial Universidad de Costa Rica, 2001-v. ISSN-1409-3871 1. Botnica Publicaciones peridicas, 2. Publicaciones peridicas costarricenses

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MIGUEL NGEL SOTO ARENAS (JUL Y 12, 1963 – AUGUST 27, 2009) 1 EDUARDO A. PREZ GARCA 2 & ERIC HGSATER 32 Departamento de Ecologa y Recursos Naturales, Facultad de Ciencias, Universidad Nacional Autnoma de Mxico, Ciudad Universitaria, Coyoacn, Mxico D.F., eduardo.perez-garcia@ciencias.unam.mx3 AMO Herbarium, Instituto Chinon, Mexico City, herbamo@prodigy.net.mx269. 2010. LANKESTERIANA 9(3): Miguel ngel Soto was born in the city of Torren, Coahuila, on July 12, 1963. In that city he studied elementary and secondary school at the Colegio at the Universidad Autnoma del Noreste (1978became involved in the world of plants. His parents loved plants and he inherited great part of this culture. Living in a semi-arid region allowed him to get to know closely one of his favorite groups of plants, about which he acquired ample knowledge: the cati and succulents. In spite of living in the desert, Miguel ngel already knew and cultivated his favorite plants, the orchids. Numerous visitors came to his house, even journalists from the local communication media, to see his orchid house. Later on, Miguel ngel moved to Mexico City to follow university studies; he majored in Biology at the Faculty of Sciences /UNAM) from 1982-1987. undergraduate thesis on the orchids of Bonampak, Chiapas. It is suitable to point out that in those times access and with great extensions of the poorly known Lacandon Jungle. From there, Miguel ngel and Jorge generated by the eruption of the “El Chichn” (or Chichonal) volcano. Miguel ngel was a perfectionist thesis, although he did publish an article on that results of that sojourn in the heart of the Lacandon Jungle: Soto Arenas, M.A., (1986), Orqudeas de Bonampak, Chiapas. Orqudea (Mex.) 10(1): 113-132. Miguel graduated some-time later with a new thesis topic, this time on the genus Lepanthes. He wrote the thesis with Gerardo Salazar, and their work served as the basis for the publication of the book: The genus Lepanthes Sw. in Mexico, where they published 32 new species, doubling the number of known species for Mexico. projects and publications on orchids. During this period, Miguel spent his days between the Ecology Laboratory (of the Faculty of Sciences, UNAM) and the AMO Herbarium since its inauguration (at that time it was located in La Herradura, in the outskirts of Mexico City, at Eric Hgsaters house), and his herbaria of the United States and Europe, studying all the Mexican material on orchids; these voyages being very fruitful in the search of types of numerous species native to Mexico. Miguel taught some undergraduate courses of the biology, in the Faculty of Sciences, UNAM. Among them: Bio-geography on 10 times (from 1983-1993), General Ecology twice (1987-1988), Systematics (1999), and Natural Resources (2000-2001). He was the Montane Cloud Forest of the Sierra Madre in the South of Guerrero” (1983-1984) and “Sinecological Analysis of the Montane Cloud Forest of the State Park of Omiltemi, Guerrero.” (1984-1985). He directed or co-directed around ten students both of undergraduate as well as graduate levels. Some of them received recognitions for the quality of their thesis; like Rodolfo Solano Gmez whose thesis “The Genus Stelis Sw. (Orchidaceae: Pleurothallidinae) in Mxico” (E.N.E.P. Iztacala, UNAM, written in 1993 in co-direction with E. Hgsater, AMO Herbarium) earned an Honors Mention in the Undergraduate 1 Translation by Carlos Ossenbach.

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Thesis Contest of the Botanical Society of Mexico. Similarly, Mariana Hernndez Apolinar received First Place in the Undergraduate Thesis Contest of the Botanical Society of Mexico with the thesis “Population Dyanmics of Laelia speciosa (HBK) Schltr. (Orchidaceae)” (Faculty of Sciences, UNAM, written in 1992 with the co-direction of Irene Pisanty). In October of 1993 he organized very successfully the 5th Latin American Meeting of Orchideology in Xalapa, Veracruz, with the participation of the best known specialists and scholars of orchids of tropical America. In this event an important international exhibition of orchids was also held and of which he was President of the Organizing Committee (Exporqudea Xalapa ). Miguel ngel was Vice-President of the Latin American Orchideology Commission (C.L.O., 1991-1993) and President of the same in 1993. Miguel ngel was admitted into the PostGraduate Program in Ecology of the Ecology Institute in 1994. He approved all the credits and passed the doctoral candidacy examination. However, due to his perfectionism, he postponed sitting for his graduation examination until he had the publications he considered were necessary. The thesis he developed was titled “Evolution in Vanilla (Orchidaceae): Phylogeny, biogeography and evolution of characters” and he prepared it under the direction of Dr. Elena lvarez-Buylla (Laboratory of Molecular Genetics and Evolution, Ecology Institute, UNAM). Miguel ngels studies on vanillas were not limited to the aspects mentioned in the title of his thesis, since, for example, he also included the study of diversity, genetic variation and the uses historic of the vanillas. From this work interesting results derived, such as the routes and dates of the human dispersion of the cultivation of Vanilla planifolia around the world. Unfortunately he did not see the publication of the formal description of several new species and varieties of this genus, like Vanilla costarricensis (ined), V. cribbiana (ined), V. dessleri (ined), V. martinezii (ined), V. sarapiquinsis (ined), V. pompona subsp. (Lindl.) Soto Arenas (ined), and V. espondae (ined). which are now published in this issue of Lankesteriana, and the corresponding species are included in his contribution to Flora Mesoamericana, which had already been delivered to the editors at Missouri Botanical Gardens. From his studies on vanillas the following were published or were left to be published: perspectivas de su cultivo. (Vanilla: Challenges and Perspectives of its cultivation). Biodiversitas 66: 1-9. Vanilla from South America. Lankesteriana. A revision of the Mexican and Central American species of Vanilla Plum. ex Mill: Conspectus of morphological and molecular data. Lankesteriana. lvarez-Buylla. (in preparation) Phylogenetic analysis of Vanilla Plum. ex Mill. (Orchidaceae: Vanillianae) from congruent morphological and molecular data. the genus Vanilla Plum. ex Miller (Orchidaceae, Vanillinae). Mexican Vanilla (Orchidaceae) and the evolution of pollination systems in the genus. (in preparation) Bio-geographic history of the Pantropical genus Vanilla and the history on the Gondwanic tropical biota. Vanilla planifolia: what morphology, isozimes, RAPD’s, and nuclear DNA sequences indicate. Miguel ngel was one of the most knowledgeable of all of Tropical America. He described, alone or as co-author, many new orchids, among many others are: Phragmipedium xerophyticum, Barkeria fritzhalbingerii, Rossioglossum hagsaterianum, Sobralia macdougalii, Stanhopea dodsoniana, S. whittenii, Stelis greenwoodii, Elleanthus teotepecensis, Encyclia calderoniae, E. rzedowskiana and Oncidium leleui. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.270 LANKESTERIANA

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of Laelia anceps, author) numerous species of different genera, among them Barkeria, Elleanthus, Rhynchostele, Prosthechea, Dichromanthus, etc. He had ample knowledge on the Pleurothallidiinae (Stelis, Acianthera, Pleurothallis, etc.). According to the Missouri Botanical Garden database (W3TROPICOS), there are more than 160 species and sub-species described by him, including previously published taxa. Miguel collected more than 11,000 different samples (collection numbers) of plants in Mexico, Guatemala, Costa Rica, Panama and Brazil, which include almost 150 type collections. The main set of his collections as well as his collecting notebooks and personal notes are deposited in the AMO Herbarium (Chinoin Institute, Mexico City). Miguel was a tireless traveler and few people knew the natural habitats of orchids like he did. This is one of the reasons why he was one of the authors that wrote most in the book “The Orchids of Mexico”, and personally supervised its design and edition; the work describes a journey through the ecosystems of Mexico and its orchids, including cultural and conservation chapters. Together with the digital Catalogue (CD) of The Orchids of Mexico it is the most complete popular work there is about Mexican orchids. One of his already classic works was the publication: Updated Listing of the Orchids of Mexico [ Orqudea (Mexico City.) 11: 231-275 (1989)]; this list was the basis of the most recent lists about the orchids of Mexico. Jointly with Federico Halbinger he coauthored the book Laelias of Mexico, which is one of the most widely cultivated genera by those fond of orchids. Miguel ngel was executive editor of the journal Orqudea (Mexico City.) from 1985 to date. He was also executive editor of some volumes of the Icones Orchidacearum (Mexico) , which is probably the best technical reference of the Mexican orchids. He had two other volumes in preparation. Other issues deal with the genus Epidendrum throughout the neotropics. Also, Miguel ngel collaborated with several of the great current orchideologists including Robert Dressler, Gerardo Salazar, Eric Hgsater, Germn Carnevali, Mark Chase, Cssio van den Berg, Mark Whitten, Phil Cribb, and Ed Greenwood, among many others. Miguels publications are a mandatory reference for orchid scholars, but are also very useful the Neotropics. Due to his great work as a botanist, several species have been dedicated in his honor like: Lepanthes sotoi Archila, Maxillaria sotoana Carnevali et Gmez-Jurez, Mormodes sotoana Salazar, Stelis sotoana R. Solano, and some others that are being published in this issue of Lankesteriana. Miguel was a conservationist since an early age; for example, he was one of the most participative students in the creation of the Reserve of the Pedregal of San ngel in Mexico City. More recently, he published one of the most complete works on the current situation of orchid conservation in Mexico and participated in many forums related with the conservation of orchids. He was a prominent member of the “Orchid Specialist Group, Species Survival Commission, IUCN” (19931997; 1998-2009) and a member of the “Conservation ex-situ Committee” of the same commission (2000-2003). He also participated as counselor of various government agencies like SARH (Ministry of Agriculture and Water Resources), SEDESOL (Ministry of Social Development), SEMARNAT (Ministry of Natural Resources), CONABIO (National Comission of Biodiversity), establishing the most important criteria for the national orchid conservation strategies of Mexico. Miguel ngel was a botanist who loved plants and various families, among which stand out the Crassulaceae, the Cycadaceae and of course, orchids. He was able to form the most important collection of live plants of Mexican orchid species, much of which is now located in the live collection at AMO samples, he helped form a DNA bank for research in molecular biology of almost 500 orchid species, and in addition 500 Vanilla samples. In collaboration with E. Hgsater and Cssio van den Berg, he was preparing a phylogeny of the genus Epidendrum based both on molecular data as well as morphological and on the vegetative architecture; for that he had sequenced more than 300 species from throughout the neotropics, mostly collected by Hgsater and cultivated at AMO, always careful to have voucher specimens prepared.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PREZ-GARCA & HGSATER — Miguel ngel Soto Arenas (1963-2009)271

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He stood out for his attitude, always cooperating with hobbyists and other biologists, which led him sessions of the Mexican Orchid Association, and with farmers, particularly those that cultivate vanilla in the states of Veracruz, Oaxaca and Chiapas. He seminars, he delivered more than 150 conferences in universities, in botanical gardens, and in botanical and horticultural associations. Those of us who had the good fortune of having vegetation of Mexico. His acute vision allowed him or in the darkness of the closed forest canopy. It is suitable to point out that he performed as a human altimeter, since he could calculate very exactly the altitude where we found ourselves by only looking at the vegetation around us. Nora Esponda, the associate secretary of Instituto Chinon, worked with Miguel for 25 years and developed a close friendship/professional relationship, and describes him as passionate in his work and everything he did, his conversations, his friendship, he was also fun, and worked late into night, sending mails and instructions in the early morning. She claims he described himself as “neurotic and ill-tempered”, but she never saw him loose control, his character was strong but affable, very demanding of himself, but also very sensitive. His sister Miriam describes him as a great Chef, it was Miguel who invited the family over for Christmas or some special occasion. He enjoyed cooking a chicken in “mole”, or a leg of venison with herbs in white wine for his friends. His specialty, vanilla ice-cream, was delicious. He moved back to Torren a couple of years ago in search of tranquility to continue writing about orchids of Mexico, and spend much more time with his parents and two sisters. He had recently traveled with his sisters on a month-long trip through the Huasteca region of San Luis Potos, Veracruz and into Chiapas; they were amazed how well he knew the country and the most spectacular landscapes. A couple of weeks before his assassination, he had discovered what appeared to be a new species of Pine-tree, on an excursion with his sister Miriam and other friends. His tragic death occurred on August 27, at his home in Torren, Coahuila. Miguel ngel Soto was a controversial character, but without a doubt, he was an extraordinary human being who leaves behind a him. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.272 LANKESTERIANA

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IN MEMORIAM : MIGUEL NGEL SOTO ARENAS (1963-2009) ERIC HGSATER 273. 2010. LANKESTERIANA 9(3): gatherings of the Asociacin Mexicana de Orquideologa in the early 1980’s. He and Gerardo A. Salazar were studying biology at the Faculty of Sciences at the UNAM, the National University in Mexico City. Both made their social service mounting specimens and looking after the live orchid collection at the recently founded Herbario AMO (then a part of the Asociacin Mexicana de Orquideologa). They both proved to have a deep interest in orchids, and decided to pursue a career in orchid research. Miguel chose to specialize in ecology and of Bonampak, Chiapas, were he became particularly well acquainted with the orchids of this tropical, lowland rain-forest. He later participated in several studies of the mesophilous forests of Omiltemi, Guerrero, another site of high orchid diversity, but this time in the higher mountains of southern Mexico, undergraduate studies, that they both became formally employed as a research associates in Mexican orchids by the Instituto Chinon, A.C., in particular in the AMO Herbarium as a base. The objective was to further the knowledge of Mexican orchids from every viewpoint. Through the years Miguel was able to travel to the most important herbaria in the Americas and Europe, spending sometimes weeks or months at a time, studying and photographing Mexican orchid material. The information contained in the images was transferred to the electronic database at AMO, now covering some 130 thousand records. Most of the slides have been digitalized for easier access. Miguel thus obtained an encyclopedic knowledge of the Mexican orchids. One of his goals was to Orchids of Mexico , a treatise covering all the taxonomic and ecological information of all the known Mexican orchid species. This work was done in collaboration with Hgsater, who focused on the genus Epidendrum , Gerardo A. Salazar, on terrestrial orchids, and Rodolfo Solano in relation to the Pleurothallidinae; Rolando Jmnez worked on Oncidium . Unfortunately, this work, now covering From the early beginning he stood out as someone willing to learn from others, doing team work and sharing his information with others. This can be seen through the number and variety of thesis he directed, not only with students in Mexican universities, but also at Oxford, England and Riverside, California, as well as in the many papers he co-authored. Miguel tackled many groups of mainly epiphytic genera, and had a special interest in the Mexican Laelia , where he worked together with Federico the species and lower taxa, having personally visited with Federico practically each and every locality, to understand each entity. During several years he worked with Mariana Hernndez in the state of Michoacn studying the population dynamics of Laelia speciosa , studing how market in ways that could be sustainable. They also followed seedlings on oak trees for several years, recording how many survived each successive year, studies with terrestrial orchid species in Europe and the United States, but very few with epiphytic species in tropical countries. Mariana’s thesis was recognized as the best thesis of the year by the Mexican Botanical Society. Once, while studying the material of Phragmipedium exstaminodium he exclaimed “you collected this specimen the day I was born!” Indeed

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that June day in 1963 I was visiting the lakes of Monte Bello, a two day jeep-ride from Comitn, through cattle ranches and virgin forests in the highlands of Chiapas. The clear-water, coloredlakes where surrounded by Pine-Oak forests covered with epiphytes, including many orchids, and the amazing lady-slipper with two-foot long petals in full bloom! Miguel and I were deeply aware of the destruction of many natural habitats, mainly due to their transformation into farm-land. That forest very hot spring of 1998, after a deep-cold winter the previous year had killed many of the epiphytes in evaluate the state of conservation of Phragmipedium exstaminodium for the IUCN. A couple score mature plants were found, but no young seedlings, there appeared to be no natural reproduction. Miguel, with his interest in ecology, participated in numerous conservation conferences, was a member of the IUCN Orchid Specialist Group, and collaborated with numerous governmental agencies regarding orchid conservation. He prepared the revision of the list of endangered orchids for the Mexican government, prepared monographs of each species, the illustrations prepared by several other botanists at AMO. during these 25 years of working together. We often discussed plans for the future and worked on many projects, one of them being a new system of Epidendrum , combining over 300 species of the genus throughout most of its range, most of the samples from plants which I had Colombia and Ecuador. We came to have a good understanding of the variation. His sudden departure leaves this project in chaos. One of our dreams, since we began working together, was to eventually publish an illustrated book on the Orchids of Mexico. We agreed that it should rather a narration of the many diverse habitats where orchids can be found, a walk through these forests and savannas. Through the years we gathered material, and he became intimately acquainted with each and edition. I discussed the issue with several prominent late 2004, at the pharmaceutical company of which Instituto Chinoin is a part, I found myself discussing how we could commemorate the upcoming 80th anniversary of the introduction of its products to the Mexican market. All things came together; we could publish a book of interest to a cultivated public, which was the result of the accumulated knowledge long cherished dream into action. Miguel was at the help of Gerardo Salazar. The photographs were to be made mainly by Rolando Jmnez Machorro and Marco Lpez Rosas, the illustrators and photographers at AMO. We contracted one of the it was evident that we could not include photographs of all the species known to Mexico, we decided to add a CD with as many as we could get. We asked for and got collaboration from all quarters of the world for suitable slides, three dozen photographers participated. Miguel dropped everything else he was working on, including his doctoral thesis on Vanilla . He got involved in everything, including the balance and quality of the color selections, and the lay-out of revision by Robert Dressler. The book was printed in Japan and arrived in time for the anniversary of the pharmaceutical company. Miguel had put a full year’s work into it, often working until mid-night. It was an immediate success, within a year the 25,000 copies printed in Spanish were distributd and the edition was sold out; those who received the book did not let go of it. It had, however, a surprising unintended effect; it became the standard show-book for sellers of wild-collected orchid species in the local markets in Mexico City and elsewhere. We were dismayed by this. After years of efforts by Miguel Angel to curb this illegal market which depleted populations of mostly desirable horticultural species, growing and collecting species orchids became popular again, with very few sources of propagated plants.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.274 LANKESTERIANA

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Probably one of Miguel’s most cherished projects was Vanilla . Aside from being the subject of his doctoral thesis, for which he sequenced some 500 samples from around the world, he discovered that single clone probably selected by the Totonaca people of the State of Veracruz. He had special interest in working with local communities in recuperating this valuable crop. He searched for the few individual plants with diverse genetic structures. Working in collaboration with several specialists throughout the world he prepared a proposal for a new generic new to science. Seven papers were prepared for publication, unfortunately only one was sent to the by the coauthors, and will hopefully be published sometime soon, some in this issue dedicated to Miguel ngel. Miguel was working at his home in Torren, Coahuila, late at night on August 27th. His sister had been helping him till eight o’clock that evening. Suddenly, an assailant entered his house, and after unknown. His body was discovered the following day. He was 47 years old. His sisters and parents, the whole orchid community, is shocked, as we have all lost a great friend and collaborator, and one of the greatest contributors to the knowledge of orchids in Mexico, not only their diversity, but their ecology and conservation. May he rest in peace.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. HGSATER — In memoriam: Miguel ngel Soto Arenas275

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Lorenzo S.A., L., A. Ramrez, M.A. Soto Arenas, A. Breceda, M. del C. Caldern, H. Cortz, C. Puchet, M. Sierra Madre del Sur, Mxico. Bol. Soc. Bot. Mx. 44: 97-102. Soto Arenas, M.A. 1982. El Pedregal de San Angel: un refugio natural de orqudeas. Ocelote 1(1): 9-11. Soto Arenas, M.A. 1986. Orqudeas de Bonampak, Chiapas. Orqudea (Mx.) 10(1): 113-132. Soto Arenas, M.A. 1986. El gnero Elleanthus en Mxico y una nueva especie de Guerrero, Elleanthus teotepecensis Orqudea (Mx.) 10(1): 161-190. 1986. Soto Arenas, M.A. 1987. Una revisin de las especies mexicanas de Trichosalpinx subgen Trichosalpinx. Orqudea (Mx.) 10(2): 247-296. M.A. Soto Arenas. 1998. Orchid conservation in Mexico. Selbyana 19(1): 15-19. M.A. Soto Arenas. 1989. Lepanthes hagsateri, una nueva especie de Guerrero, Mxico. Orqudea (Mx.) 11: 15-22. 1989. Soto Arenas, M.A. Orch. Research Newsletter 13: 8-9. Soto Arenas, M.A. 155-172 in J.L. Camarillo y F. Rivera (eds.) Areas Naturales Protegidas y Especies en Extincin. UNAM. Mxico, D.F. Soto Arenas, M.A. Phragmipedium xerophyticum, una nueva especie del sureste de Mxico. Orqudea (Mx.) 12(1): 1-10. (German translation: Neue Frauenschuhorchideen aus Mexiko, Phragmipedium xerophyticumDie Orchidee 42(5): 253-262. 1991). M.A. Soto Arenas. 1990. Una nueva especie de Malaxis de Chiapas. Acta Botnica Mexicana 10: 45-49. Soto Arenas, M.A. 1990. Una nueva orqudea de Morelos, Mxico: Ponera dressleriana. Orqudea (Mx.) 12: 117-126. A. Soto Arenas. 1990. Oncidium leleui Orqudea (Mx.) 12(1): 57-64. (French translation: Oncidium leleui du Mexique. Orchidees. Culture et Protection 10:3-9. 1992). Soto Arenas, M.A. 1991. Studien ber die Population und die Moeglichkeit der Erhaltung des Phragmipedium exstaminodium (Orchidaceae), ein gefehrdeter Arten. pp 132-134. In E. Hgsater (ed.) Phragmipedium exstaminodium bedroth. Die Orchidee 42(3): 127-134. M.A. Soto Arenas. 1992. Lepanthes williamsii (Orchidaceae), una nueva especie de Mxico y Guatemala. Orqudea (Mx.) 12(2): 139-143. M.A. Soto Arenas. 1992. El complejo Oncidium maculatum. Orqudea (Mx.) 12(2): 297-316. Meave, J., M.A. Soto Arenas Bol. Soc. Bot, Mxico 52: 31-77. Soto Arenas, M.A. 1992. Una nueva especie de Chiapas: Ornithocephalus obergiae. Orqudea (Mx.) 12(2): 193-198. Soto Arenas, M.A.1992. Maxillaria histrionica y M. tonsoniae. Orqudea (Mx.) 12(2): 244-250. Soto Arenas, M.A. 1992. Maxillaria hagsateriana, una nueva especie similar a Maxillaria densa. Orqudea (Mx.) 12(2): 251-260.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PEER REVIEWED PUBLICATIONS (P APERS AND CHAPTERS IN BOOKS) MIGUEL NGEL SOTO ARENAS: PUBLICATIONS AND MAIN CONFERENCES, 1983-2009 276 2010. LANKESTERIANA 9(3):

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.Miguel ngel Soto Arenas: publications and conferences (1983) 277Soto Arenas, M.A. 1992. Draculla pusilla Orqudea (Mx.) 12(2): 277280. Soto Arenas, M.A. Maxillaria lexarzana, un nuevo nombre para Psitacoglossum atratum Orqudea (Mx.) 12(2): 237-243. Soto Arenas, M.A. (1993). Population studies in Mexican orchids. pp. 153-160 in A.M. Pridgeon. [ed.] Proceedings of the 14 th World Orchid Conference, Glasgow. HMSO Publ., Edinburgo. Soto Arenas, M.A. (1994). Orchidaceae pp. 170-176 en Martnez, E., C.H. Ramos y F. Chiang. Lista Florstica de la Lacandona, Chiapas. Bol. Soc. Bot. Mxico 54: 99-177. Soto Arenas, M.A. (1994). Barkeria melanocaulon y Barkeria whartoniana. Orqudea (Mx.) 13(1-2): 233-244. Soto Arenas, M.A. (1994). Barkeria fritz-halbingeriana. Orqudea (Mx.) 13(1-2): 245-248. Soto Arenas, M.A. (1994). Eurystyles Orqudea (Mx.) 13(1-2): 269-274. Soto Arenas, M.A. (1994). Vanilla odorata. Orqudea (Mx.) 13(1-2): 290-294. Soto Arenas, M.A., G.A. Salazar y A. Rojas. (1994). Nomenclatural changes in Rhynchostele and Lemboglossum (Orchidaceae, Oncidiinae). Orqudea (Mx.). 13(1-2): 145-152. Jimnez, R. y M. Soto Arenas. (1994). Oncidium hagsaterianum, una nueva especie de Mxico y Guatemala. Orqudea (Mx.) 13(1-2): 159-164. Cribb, P. y M.A. Soto Arenas. (1994). The genus Cypripedium in Mexico and Central America. Orqudea (Mx.) 13(1-2): 125-144. (Cribb, P. und M. Soto Arenas. 1994. Die Gattung Cypripedium America. Die Orchidee: 1995). Solano, R. y M.A. Soto Arenas. (1994). Stelis aeolica y S. vespertina en R. Solano, El gnero Stelis en Mxico. Orqudea (Mx.) 13(1-2): 18-21; 106-108. Salazar, G. y M.A. Soto Arenas. (1996). A new species of Ornithocephalus (Orchidaceae) from Oaxaca, Mexico. Brittonia 48(2): 209-212. Soto Arenas, M.A. (1996). The importance of research. pp. 33-38. en IUCN/SSC Orchid Specialist Group. Orchids Status Survey and Conservation Action Plan. IUCN, Gland, Suiza. Soto Arenas, M.A. (1996). Mexico (Regional Account). pp. 53-58 en IUCN/SSC Orchid Specialist Group. Orchids Status Survey and Conservation Action Plan. IUCN, Gland, Suiza. Soto Arenas, M.A. (1997). Introduction aux orchides mexicaines. Orchidees. Culture et Protection. 31: 18-21. Hgsater, E. y M.A. Soto Arenas. (en prensa). Notes for a of Epidendrum, the largest Neotropical orchid genus. Proceedings of the 18th World Orchid Conference, Miami, Florida. Estados Unidos. van der Berg, C., W.E. Higgins, R.L. Dressler, W.M. Whitten, M.A. Soto Arenas, A. Culham, y M.W. Chase. (2000). A phylogenetic analysis of Laeliinae (Orchidaceae) based on sequence data from Internal Transcribed Spacers (ITS) of nuclear ribosomal DNA. Lindleyana 15(2): 96-114. Salazar, G.A. y M.A. Soto Arenas. (2001). A new species of Stanhopea (Orchidaceae) from Mexico. Lindleyana 16(3): 144-148. Salazar, G.A. y M.A. Soto Arenas. (2001). Nomenclatural changes in Mexican Orchidaceae. Lindleyana 16(3): 149-150. Salazar, G.A., M.W. Chase y M.A. Soto Arenas. (2002). Galeottiellinae, a new subtribe and other nomenclatural changes in Spiranthinae (Orchidaceae: Cranichideae). Lindleyana 17(3): 172-176. Salazar, G.A., M.W. Chase, M.A. Soto Arenas y M. Ingrouille. (2003). Phylogenetics of Cranichideae with emphasis on Spiranthinae (Orchidaceae, Orchidoideae): Evidence from plastid and nuclear DNA sequences. Am. J. Bot. 90(5): 777-795. Light, M., J. Reddoch, E. Hgsater y M. Soto Arenas. (2003). Edward W. Greenwood. In memorian. Bol. Soc. Bot. Mx. 71: 85-84. Soto Arenas, M.A. 2003. Vanilla. Pp. 321-334. In: Pridgeon, A.M., P.J. Cribb, M.W. Chase y F.N. Rasmussen.

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(eds.). Genera Orchidacearum vol. 3. Orchidoideae (Part two), Vanilloideae. Oxford University Press. Oxford. Soto Arenas, M.A. Vanilla. Pp. 383-387. In B.E. Hammel, M.H. Grayum, C. Herrera y Manual de Plantas de Costa Rica Missouri Botanical Garden,-INBio-Museo Nacional de Costa Rica. Soto Arenas, M.A. Domingoa. In: Pridgeon, A.M., P.J. Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. in A.J. Garca-Mendoza y M.J. Biodiversidad de Oaxaca. Instituto de Biologa, UNAM-Fondo M.A. Soto Arenas. 2005. Epidendrum In: Pridgeon, A.M., P.J. Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. M.A. Soto Arenas. 2005. Transfers to Epidendrum from Oerstedella Rchb.f. Lankesteriana 5(1): 73-75. Soto Arenas, M.A. 2005. Alamania In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Arpophyllum. In: Pridgeon, A.M., P.J. Cribb, M.W. Chase Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Artorima. In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Barkeria. In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Subtribe Chysinae. Chysis In: Pridgeon, A.M., P.J. Cribb, Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Hagsatera. In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. HelleriellaIn F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Homalopetalum. In: Pridgeon, A.M., P.J. Cribb, M.W. Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Laelia In Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Meiracyllium. In: Pridgeon, A.M., P.J. Cribb, M.W. Chase Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Microepidendrum In: Pridgeon, A.M., P.J. Cribb, LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.278 LANKESTERIANA

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Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Nemaconia In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Ponera In F.N. Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Subtribe Ponerinae. Pp. 422-424. In Rasmussen. (eds.). Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2005. Arpophyllum (pp. 22-25), Barkeria (pp. 46-51; en colaboracin colaboracin con E. Hgsater), Isochilus (pp. 382-385) y Jacquiniella (386-389) In: Pupulin, F. (ed.) y colaboradores. Vanishing Beauty: Native Costa Rican Orchids. vol. 1. Editorial de la Universidad de Costa Rica. San Jos. M.A. Soto Arenas In Genera Orchidacearum vol. 4. Epidendroideae (Part one). Oxford University Press. Oxford. Soto Arenas, M.A. 2006. La vainilla: Retos y perspectivas de su cultivo. Biodiversitas 66: 1-9. Soto Arenas, M.A. Domingoa, Homalopetalum (Orchidaceae: Laeliinae) and Nemaconia (Orchidaceae: Ponerinae). Neodiversity 2: 7-9. Schlter, P.M., M.A. Soto Arenas Vanilla planifolia (Orchidaceae). Economic Botany 61(4): 328-336. Soto Arenas, M.A. Mexican orchids. Lankesteriana 6: 123-132. van den Berg, C., W.E. Higgins, R.L. Dressler, W. Mark Whitten, M.A. Soto-Arenas phylogenetic study of Laeliinae (Orchidaceae) based on combined nuclear and plastid DNA sequences. Annals of Botany, 104 (3): 417-430. BOOKS Soto Arenas, M.A. 1990. (30/100 texts of plates, second author) in Mxico, part 1. Icones Orchidacearum. Asociacin Mexicana de Orquideologa. Mxico D.F. M.A. Soto Arenas. 1996. El gnero Lepanthes en Mxico. Asociacin Mexicana de Orquideologa, A.C. Mxico D.F. 231 pp. M.A. Soto Arenas. 1997. Laelias of Mexico. Herbario AMO. Mxico D.F. 160 pp. Soto Arenas, M.A In: Orchids of Mexico, parts 2 and 3. Icones Orchidacearum fasc. 5-6. Herbario AMO. Mxico D.F. Hgsater, E., M.A. Soto Arenas 2005. Las Orqudeas de Mxico. Instituto Chinon, Mxico D.F. (simultaneous edition in English, Orchids of Mexico). Soto Arenas, M.A. In: of Mexico, part 4. Icones Orchidacearum fasc. 10. Herbario AMO. Mxico D.F. OTHER PUBLICATIONS Preparation and edition of the “Timbre de la Tuberculosis”. Comit Nacional de la Lucha contra la Tuberculosis y Enfermedades del Aparato Respiratorio”. (1997). Soto Arenas, M.A., E. Hgsater, R. Jimnez Machorro, G.A. Salazar Chvez, R. Solano Gmez, R. Flores LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.Miguel ngel Soto Arenas: publications and conferences (1983) 279

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Win-Mac. Herbario AMO. Instituto Chinon, A.C. Mxico. Soto Arenas, M.A G.A. Salazar). 2007. Fichas tcnicas de las especies de orqudeas includas en la NOM-059-ECOL-2007 (SEMARNAT). SPECIAL INVITED AND KEYNOTE CONFERENCES “La Flora y la Vegetacin de Mxico”. Museo Nacional de Historia, Castillo de Chapultepec, Cd. de Mxico. January 1993. “Population Studies in Mexican Orchids”. 14th World Orchid Conference, Glasgow, Scotland, April 1993. “Orqudeas Mexicanas en Peligro de Extincin”. V Congreso Nacional de Horticultura Ornamental, 1995, FES Cuautitln, UNAMAsociacin Mexicana de Horticultura Ornamental. Keynote Conference. “Laelias, Barkerias and Rhynchosteles. Jewels of the Mexican Orchids. “ Iowa Orchid Society. Des Moines, IA., 1997. “Systematics of Vanilla (Orchidaceae)”. Royal Botanic Gardens, Kew, Jodrell Laboratory, U.K. April 1999. “Genetic Resources of Vanilla”, 16th World Orchid Conference, Vancouver, May 2000, “Orqudeas Mesoamericanas: Evolucin, experiencias y perspectivas de conservacion”. Chairman conference. II Seminario Latinoamericano de Orquideologa, San Jos, Costa Rica (in collaboration with G. Salazar and E. Hgsater), May 2001. “La Conservacin de las Orqudeas de Mxico”. Simposium de Conservacin, XII Congreso Mexicano de Botnica, Quertaro (in collaboration with G. Salazar and E. Hgsater), October 2001. “CITES, conservation and orchid taxonomy in the Neotropics; personal notes and a proposal of criteria for noncommercial collections” (in collaboration with E. Hgsater), 17th World Orchid Conference, Shah Alam, Malaysia, April 2002. “Phylogeny of Epidendrum” (in collaboration with E. Hgsater), 17th World Orchid Conference, Shah Alam, Malaysia, April 2002. “Evolucin y recursos genticos en Vanilla (Orchidaceae)”. Seminarios de Botnica, Instituto de Biologa, UNAM. May 2002. “Evolucin de Epidendrum” y “Biogeografa de las orqudeas del Neotrpico” (Keynote Conferences), VI Taller Internacional de Orqudeas, Jardn Botnico Orquideario Soroa, Universidad de Pinar del Ro, Cuba. November 2002. “Evolucin en Vanilla (Orchidaceae)”. Seminario, Colegio de Posgraduados, Montecillos, Edo. de Mxico. “Las Orqudeas de Oaxaca” . Jardn Histrico Etnobotnico. Centro Cultural Santo Domingo, Oaxaca, Oax. “Vanilla The challenges of a crop based on an endangered species with a complex life history”. Vanilla 2005. Third International Congress, Boca del Ro, Ver., November 2005. “La Vainilla Los retos de un cultivo basado en una especie amenazada con una historia de vida compleja”. Congreso internacional de productores de Vainilla. Papantla, Ver. May 2006. Epidendrum, the largest Neotropical orchid genus” (in collaboration with E. Hgsater), 18th World Orchid Conference, Miami, Florida. U.S.A. January 2008. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.280 LANKESTERIANA

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A NEW SPECIES OF VANILLA FROM SOUTH AMERICA1 MIGUEL A. SOTO ARENAS 281. 2010. LANKESTERIANA 9(3): During my work with the phylogeny of Vanilla it became evident that some specimens represent undescribed species in the genus. The species here described is among the most showy in the genus, suggesting that additional, undescribed vanillas may be remain to be found*. The taxon is native to South America, where Vanilla reaches its highest diversity, and from where much more material is needed. Its relationships are discussed. Vanilla espondae Soto Arenas, sp. nov. HOLOTYPE: [N.] Colombia: Tributary of R. Magdalena, cult. F. Prez-Vera 563, K-L! (illustration voucher). A Vanillae trigonocarpae Hoehne disco labelli papillis verrucisque ornato recedit. Hemi-epiphytic herb . Stems apparently thin, terete, smooth, olive green, ca. 6.3 mm thick, the internodes as long as the leaves. Leaf shortly petiolate, the petiole ca. 8 mm long, blade oblanceolate, abruptly acuminate, the apex incurved, base obtuse, the basal x 3.6 cm. axillary, a shortly pedunculate bracts unknown. Ovary sub-trigonous conspicuously white. Flowers : buds whitish at base, apex green, with the midrib of petals protruding between the diameter when spread out; tepals ivory white, outer, basal surface of the lip ivory-white, mid lobe and throat ivory-white lined with yellow-ochre, papillae of the midlobe yellow; the segments spreading. Dorsal sepal recurved, apparently long oblanceolate, acute, perhaps as long as the lateral ones. Lateral sepal s directed downwards, somewhat falcate, obliquely oblanceolate, obtuse-subacute, base long attenuate, with a prominent axial vein on the abaxial surface and corresponding to the axial groove on the adaxial surface; very smooth, 74-78 x 20 mm. Petals spreading, somewhat arcuate, slightly recurved at the apex, narrowly elliptic-oblanceolate, acuteacuminate, convex, apically conduplicate, broader than the sepals, longitudinally keeled on the outer surface, the keel broad and conspicuous, at least 76 long, more than 16 mm wide (not well preserved). Lip very showy, forming a long tube, marginally fused to the column at least 2.7-2.9 cm; when spread rounded shoulders, oblong-triangular, tapering at apex, ca. 28 x 12 mm; midlobe oblong, rounded, the base somewhat narrowed and forming an isthmus, ca. 16 x 15 mm; the disc covered by 7 showy rows of papillae, longer towards the apex, the papillae digitiform, up to 3.5 mm high, continuous with warty veins at the apex of the midlobe. Column unknown. ETYMOLOGY : This species is dedicated to my dear friend Mrs. Nora Esponda, administrative assistant at ABSTRACT . A new species from the basin of the Rio Magdalena in northern Colombia is proposed, Vanilla espondae. KEY WORDS: Orchidaceae, Vanilla , V. espondae, Rio Magdalena, Colombia 1 Editor’s note: Miguel Angel Soto Arenas passed away August 27h, 2009. We wish to thank Dr. Phillip Cribb for preparing the camera lucida drawing of the lip of the holotype, and Rolando Jimnez Machorro for preparing the attached illustration, prepared from a digital image of the holotype and the rendering of the lip by Dr. Cribb. * Editor’s note: Additional new species from Mexico and Central America are being described elsewhere in this issue: 285.

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FIGURE 1. Holotype of Vanilla espondae Soto Arenas, by permission of the Keeper, Herbarium, Royal Botanic Gardens, Kew. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.282 LANKESTERIANA

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the Herbario AMO. Nora’s work is evident in every product of our team and she has been of indispensable assistance during our research work. DISTRIBUTION : Known only from the type; from the basin of the Ro Magdalena in northern Colombia. FIGURE 2. Vanilla espondae Soto, prepared from the holotype by Rolando Jimnez M.; the lip based on a camera lucida drawing by P. Cribb.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS — A new Vanilla from South America283

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.284 LANKESTERIANA This is one of the most beautiful Vanilla species. Although it is not closely related to any other Vanilla , it could be allied to V. trigonocarpa , with which it shares the huge, whitish flowers, and bright yellow disc; V. trigonocarpa lacks the papillae and warts on the disc. Other species with similarly adorned lips, like V. helleri A.D.Hawkes or V. insignis Ames, have either smaller flowers or green tepals. The type specimen consists only of a single leaf, petal, and the lip, yet the species is so different from any other described species of Vanilla, that I have little hesitation in proposing it.

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A REVISION OF THE MEXICAN AND CENTRAL AMERICAN SPECIES OF VANILLA PLUMIER EX MILLER WITH A CHARACTERIZATION OF THEIR ITS REGION OF THE NUCLEAR RIBOSOMAL DNA 1 MIGUEL A. SOTO ARENAS 2,3 & ROBERT L. DRESSLER 4,52 Instituto de Ecologa, UNAM, Circuito Exterior s.n. Ciudad Universitaria, Coyoacn 04510 Mxico D.F., MEXICO3 4 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Costa Rica5 Corresponding author : kerry@bio-photo.com 285. 2010. LANKESTERIANA 9(3): ABSTRACT . We present a revision of the Mexican and Central American species of Vanilla . There are 15 different species in the area; Vanilla costaricensis, V. cribbiana, V. dressleri, V. martinezii and V. sarapiquensis are here proposed as new taxa, and V. pompona subsp. pittieri and V. pompona subsp. are recognized at Vanilla calyculata, V. hartii, V. helleri , V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia , V. pompona and V. trigonocarpa distribution, and other aspects of interest are discussed. Additionally, we include a key to the species. Several Mexican and Central American species of Vanilla are closely related to V. planifolia, V. x tahitensis, and V. pompona, the cultivated species of the genus, and these are thus important in plant breeding. We also include a cladistic analysis of nucleotidic sequences of the internal transcribed spacer region of the nuclear ribosomal permitting, with some exceptions that are discussed, the recognition of sterile samples and indicating that it is a good molecular marker to infer the phylogeny of this group. The similarity and relationship between V. x tahitensis * and V. odorata is discussed. Se presenta una revisin de las especies mexicanas y centroamericanas de Vanilla . Existen 15 especies reconocidas en el rea. Vanilla costaricensis, V. cribbiana, V. dressleri, V. martinezii y V. sarapiquensis se proponen aqu como nuevas especies, V. pompona subsp. pittieri y V. pompona subsp. tambin se Vanilla calyculata, V. hartii, V. helleri , V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia , V. pompona y V. trigonocarpa tambin se describen, ilustran y se discuten aspectos Vanilla estn cercanamente relacionadas a V. planifolia y V. pompona, las dos especies ms frecuentemente cultivadas, y por lo tanto representan un germoplasma importante para este cultivo. Adicionalmente se presenta un anlisis 1 Editor’s note: Miguel ngel Soto Arenas passed away August 27th, 2009, before this manuscript was submitted for review. Vanilla remains one of the most poorly studied of all large orchid genera, and both the editors and reviewers considered that this publication is an important addition to the orchid and general botanical literature. Ken Cameron, Phillip Cribb, Eric Hgsater, Gerardo Salazar, and Rodolfo Solano did their best to offer corrections and comments to the original draft manuscript. In some instances, however, Soto Arenas quoted preliminary analyses and refers to new taxonomic concepts we were unable to recover. We have not changed the letter of the original concerns about some of the data. * Editor’s note: A paper on Vanilla tahitiensis has been published by Lubinsky et al., Neotropical roots of a Polynesian Spice: the hybrid origin of Tahitian vanilla, Vanilla tahitensis (Orchidaceae). Amer. J. Bot. 95(8): 1040 . 2008. In the article, the senior author acknowledges Soto Arenas for suggesting the topic of his PhD thesis.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.286 LANKESTERIANA cladstico de secuencias nucleotdicas de la regin de los espaciadores internos transcritos (ITS) de los genes nucleares ribosomales, donde se muestra que las secuencias de este marcador molecular ampliamente Se comenta la similitud y relacin entre V. tahitensis * y V. odorata, y el origen hbrido de la primera . KEY WORDS: Orchidaceae, Vanilla , Vanilla calyculata, V. hartii, V. helleri, V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia , V. pompona, V. trigonocarpa, Mexico, central Americas, new species The Pantropical genus Vanilla Plumier ex Miller is a group of orchid vines with about 107 species (Soto Vanilla is an ancient genus (Chase, 2001; Cameron, 2000, 2003) and the largest in the subfamily Vanilloideae (Soto Arenas, 2003). Some Vanilla species are perfumery, especially Vanilla planifolia Jacks., with V. x tahitensis J.W.Moore and V. pompona Schiede being much less cultivated (Correll, 1944; Purseglove, 1975). Vanilla exports generate US $60-80 million in foreign exchange for producing countries (Smith et al., 1992), perhaps more at present, and it is the most The culture of V. planifolia and V. pompona originated in Mexico, probably in northern Veracruz (Bruman, 1948). Although frequently cited as native of Mexico and elsewhere, there were until recently very few records of V. planifolia from wild populations, and its precise original distribution is uncertain (Soto Arenas, 1999). Many Vanilla species are similar to V. planifolia V. planifolia . Even some Mexican plantations planted with wild Oaxacan Vanilla vines include a mixture of V. cribbiana, V. insignis, V. odorata , and V. pompona , besides the true V. planifolia . Similar “mixed” plantations occur in Ecuador (P. Lubinsky, pers. com.) and Guatemala (Beza, pers. com.). It is well-known that Vanilla has peculiarities that at the top of the forest canopy), the huge vegetative variation and phenotypic plasticity characteristic of shoots and differently shaped leaves on the same individual plant as in many hemiepiphytic aroids; Putz of the membranaceous, strongly three-dimensional to the pollination by Euglossine bees (which permits the maintenance of species with little morphological other genera of orchids); Williams, 1982), and the problems derived from the less than perfect quality of type specimens. Additionally, some species are very rare, with sparse populations, and the vines are attained considerable size. Small pieces of these plants may be transplanted to greenhouses or gardens, but There are abundant data that indicate that V. planifolia is in danger of extinction in Mexico (Soto Arenas et al., 2004). Scarcity of material can be illustrated by the of V. hartii, which was found in anthesis only after 7 years of observations. Vanilla helleri and V. martinezii while V. costaricensis and V. sarapiquensis are each known from a single pressed specimen. Vanilla phaeantha and V. helleri time from Mexico, based on sterile material, whose of these taxa. Most cultivated specimens of Vanilla in botanic gardens and living collections never produce Vanilla plantations face several agricultural problems, the most important being the root rot disease caused by Fusarium batatis f . vanillae (Childers et al., 1959). It has been suggested that susceptibility to root rot is perhaps due to a narrow * Vase la Nota del Editor en la pgina anterior.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla287genetic variation, and that is to be expected in this crop due to its vegetative propagation (Purseglove, 1975; Purseglove et al., 1981; Smith, et al., 1992; Soto Arenas, 1999; Cibrin, 1999). A study of the circumscription of the cultivated species, their distribution, variation, and information on their related taxa, or those with which they may be likely confused is a necessary step toward the establishment of a germplasm bank and a breeding program in this orchid in order to enlarge the genetic foundation of the crop. In the absence of a thorough taxonomic revision, and phylogenetic framework, the breeding programs with Vanilla have used very distantly related species that are unlikely to produce fruits with commercially interesting aromatic properties (e.g. with V. aphylla , or V. barbellata ). The taxonomy of the Central American Vanilla species has been previously recognized as confusing (Dressler, 1993), and the available regional treatments are not accurate. Bouriquet (1954) revised the entire genus Vanilla, including the Central American species, but his study is out of date and he worked only with herbarium material. In “The Orchidaceae of Mexico”, Williams (1951) listed four species as native of the country, one of them, V. pfaviana Rchb.f., is actually a synonym of V. inodora Schiede. In recent years, Soto Arenas (1989, 1994) reported additionally V. mexicana Miller and V. odorata. Later, Castillo and morphology, phenology, and compatibility behavior in the cultivars of V. planifolia (which suggested more than one taxon or a strong genetic structure within the species), and additional species have been recognized in Mexico in recent years (Soto Arenas, 2003; Hgsater et al. 2005). On the other hand, V. insignis Ames (1934) was described from Honduras, V. helleri Nicaragua, and Dressler (Dressler, 1979) from Panama. Vanilla hartii Rolfe was reported from several countries of the area (Correll, 1965), and V. phaeantha Lindl. cited from El Salvador (Hamer, 1974). McLeish et al. (1995) listed three species from Belize; Hamer (1974) listed two taxa from El Salvador, Dressler (1993) included three species in his ‘Field Guide to the Orchids of Costa Rica and Panama’. Dix and Dix (2000) reported four species for Guatemala. The brief treatment recently published for Costa Rica this country, three of then unnamed, and it is largely based on the present revision. Special mention is necessary for the enigmatic V. x tahitensis J.W.Moore, described from Tahiti, but apparently introduced to the botanic garden at Papeete from Manila, Philippines in 1848, together with the true V. planifolia (Ptard 1986: 123; Hermann et al. 1989: 20). Vanilla tahitensis is very similar in morphology to some Central American taxa, and very records of it, either wild or cultivated exist from Central America, or in any other American country, but evidence has been presented of its hybrid origin from V. planifolia and V. odorata . A study of the historical records, literature, and has proven that some of the reports for Mesoamerica be reduced to the synonymy of previously described taxa, and that some other species of the area remain undescribed. This revision of the taxonomy of Mexican and Central American vanillas, together with a molecular of this area, even if it is sterile. This is an important issue, in view of the necessity of identifying vegetative material of rare, economically important For most Central American taxa, sequences of the Internal Transcribed Spacers of the nuclear ribosomal DNA (ITS) were obtained and proved to be speciesor juvenile living material. The ITS analysis is compared with sequences gathered from two larger, rbc L and mat K) in order to corroborate its utility. Material and methods Herbarium and spirit preserved material was studied in most institutions that house Central American orchids (AMO, AMES, BM, BR, CHAPA, CR, ENCB, F, G, GOEL, HEPF, K, INB, LL, MA,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.288 LANKESTERIANA W, WU, and XAL.). Efforts were made to study living material and wild specimens were observed in Mexico, Guatemala, Costa Rica, and Panama. Pollination observations and analysis of fragrances by gas chromatography (conducted by Dr. Neil da Acosta, Bush Boake Allen, London) were an additional element in circumscribing some taxa and are presented in detail elsewhere*. Due to the phenotypic variation and plasticity seen in the vanillas and mentioned above, the descriptions of vegetative features were prepared only from conditions. Stems and leaves of descending (shaded, ground-oriented shoots) are etiolated, often thinner, Vanilla after being pressed; for this reason the measurements of V. cribbiana , V. hartii , V. inodora , V. insignis , V. martinezii , V. odorata , V. planifolia , V. pompona, and V. sarapiquensis were taken from living and/or the dimensions were taken from pressed specimens, and those of V. calyculata and V. trigonocarpa also include measurements of living specimens published elsewhere. specimens (perhaps not necessarily different clones in the case of V. planifolia ) were available, either in fresh condition or in spirit, the dimensions of standard deviation, in order to determine the possible all herbarium material examined were boiled or reconstituted in a solution of ammonium hydroxide. Internal Transcribed Spacer (ITS) sequences of nuclear ribosomal DNA were collected in order to determine if this widely used molecular marker could be a useful tool to identify material. The sampling (see Table 1) does not include V. costaricensis nor V. sarapiquensis from which adequate tissue samples were not available. A special effort was made to confirm the specific identity of the specimens, even of some sterile ones. Due to difficulties in preparing vouchers, some samples remain unvouchered; however, several are housed at AMO or MEXU; pictures of other plants are kept in M. Soto’s files (which eventually will be deposited in AMO), but no voucher specimens are known to exist for most plants in living collections (e.g. Royal Botanic Gardens Kew, Nancy Botanic Garden, Jardn Nacional de Cuba, Marie Selby Botanical Gardens, Missouri Botanical Garden, Finca La Gavilana), although they were confidently identified from the living material, and locality data may help to eventually confirm the identity of the populations. Sequences of South American specimens of the V. pompona complex were also included. Also a sequence of V. claviculata , from Puerto Rico, another of V. barbellata , from Florida, and another from V. mexicana Miller, species reported for our area from unconfirmed records, were also included. The sequences of Epistephium parviflorum and Lecanorchis multiflora were defined as outgroups. It is unknown which is the sister genus of Vanilla , although morphological data suggest that it is likely Dictiophyllaria , a poorly known genus which is known only from the type specimen (Soto Arenas, 2003). Epistephium, Clematepistephium, Eriaxis and Vanilla form a polytomy in a cladistic analysis of sequences of the plastid gene rbcL . On the other hand, Lecanorchis (achlorophyllous, and in which amplification of rbcL gene has been unsuccessful) was found to be sister to Vanilla from analysis of the nrDNA gene 18S, although a more complete sampling of this region places Epistephium sister to Vanilla (Soto Arenas, unpublished data)**. DNA was extracted mostly from fresh leaves (1980), or from leaves preserved in silica gel (Chase using liquid nitrogen and adding 2% of PVP to the extraction buffer. Vanilla tissues oxidize badly when they are pressed, especially in the Membranaceous species; all attempts to extract DNA from herbarium specimens failed, although bad quality DNA has * Editor’s note pollination systems in the genus” has been found among Miguel Angel’s unpublished manuscripts. ** Editor’s note: It is unclear to which data the senior author refers here.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla289 TABLE 1 . Specimens surveyed for ITS scrutiny. K. Cameron 794 Br or TR: without precise locality? [??] Vanilla calyculata J.Linares 3386 V. calyculata J. Linares s.n. V. calyculata J. Linares s.n. HD: Moroceli. V. calyculata J. Linares s.n. V. calyculata J. Linares 8531 HD: Lizapa. V. calyculata J. Linares s.n. HD: Moroceli. V. barbellata RBG Kew s.n. without locality data V. claviculata L. Rostgaard s.n. PR: Maricao V. cribbiana M. Soto 7940 (AMO) MX: Chiapas, Chajul V. cribbiana M. Soto 8439 MX: Chiapas, Lacanj-Chanzayab V. cribbiana M. Soto 8370 MX: Chiapas, Chajul V. dressleri N. Byrd I-A-6 V. dressleri N. Byrd I-D-3 V. dressleri N. Byrd II-C-2 CR: San Jos, Ro Blanco, Pie de la Tijerilla V. dressleri N. Byrd II-C-3 CR: San Jos, Ro Blanco, Pie de la Tijerilla BR: Baha, Cat BR: Par, Xing GYF: Sinnamary, St. Elie V. cf. GD: Guadeloupe, without precise locality GY: cultivated at Kew, without precise locality V. cf. without locality data V. hartii M. Soto 8348 MX: Chiapas, Chajul V. hartii E. Salas 1 (INB) CR: San Jos, Ro Savegre, F. Chaca V. hartii N. Byrd II-F-1 CR: Puntarenas, Sndalo, Quebrada Terrones V. helleri N. Byrd II-F-4 CR: Punta Arenas, Pir V. helleri N. Byrd II-D-5 V. cf. helleri M. Soto 8818 MX: Oaxaca, Sta. Mara Chimalapa V. inodora M. Soto 7954a (AMO) MX: Chiapas, Chajul V. inodora M. Soto 8626 (AMO) MX: Jalisco, El Tuito V. insignis M. Soto 7668 MX: Campeche, Calakmul V. insignis M. Soto 8611 GT: Izabal, Punta Palma V. insignis G. Carnevali s.n. MX: Quintana Roo, La Pantera V. insignis J. Linares s.n. HD: Ro Funes (type locality). V. martinezii M. Soto 8601a (AMO) GT: Izabal, El Estor V. martinezii M. Soto 8602a (AMO) GT: Izabal, El Estor V. mexicana A. Vale s.n. CB: cultivated at La Habana, without precise locality

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.290 LANKESTERIANA TABLE 1 . Specimens surveyed for ITS scrutiny ( continues ). V. odorata M. Soto 8356 MX: Chiapas, Chajul V. odorata M. Soto 8797 MX: Oaxaca, Sta. Ma. Chimalapa V. odorata M. Soto 8822 MX: Oaxaca, Valle Nacional V. odorata M. Soto 7955 MX: Chiapas, Chajul V. odorata M. Soto 8365 MX: Chiapas, Chajul V. aff. SU: without precise locality V. phaeantha G. Carnevali 4825 MX: Yucatn, Sotuta V. phaeantha M. Soto 9920 PMA: Veraguas, El Higo V. phaeantha J. Bot. Nal. 1197-6a CB: without precise locality V. phaeantha RBG Kew s.n. UK: cultivated at RBG Kew V. planifolia M. Soto 7648 MX: Veracruz, Papantla V. planifolia M. Soto 8355 MX: Chiapas, Chajul V. planifolia M. Soto 8526 MX: Oaxaca, Usila V. planifolia M. Soto 8808 MX: Oaxaca, Sta. Mara Chimalapa V. planifolia J. Linares s.n. V. planifolia F. Pupulin 1966 CR: Heredia , Sarapiqu V. planifolia N. Byrd II-A-1 CR: Limn, Guapiles, Los Diamantes V. planifolia Dressler s.n. USA: Florida, cultivated at FLAS, paralogous and orthologous sequences V. planifolia RBG Kew cf. 1956-25802 UK: cultivated at Prince of Wales Conservatory, without precise locality V. pompona M. Soto 8614 V. pompona M. Soto 7632 (AMO) MX: Oaxaca, San Gabriel Mixtepec V. pompona M. Soto 7747 V. pompona S. Cruz sub M. Soto 8777 MX: Veracruz, Los Tuxtlas, Meyacapan V. pompona M. Soto 8119 (AMO) MX: Oaxaca, Ayozintepec V. pompona J. Linares s.n. (EAP) HD: Moroceli. V. pompona N. Byrd I-A-2 CR: Punta Arenas: Pto. Jimnez, F. La Pajuita V. pompona N. Byrd I-A-4 CR: Punta Arenas: F. La Lucrecia V. pompona Selby 1985-0211A PMA: without precise locality V. pompona M. Soto 9921 PMA: Panam, Cerro Azul V. pompona J. Bot. Nal. Cuba 86-4746 PMA: without precise locality V. tahitensis Colin s.n. FR: originally from French Polynesia V. tahitensis Gen Bank AF 391785 PNG: appartently the type of V. hirsuta V. trigonocarpa N. Byrd I-E-1 CR: Puntarenas, Pir V. trigonocarpa N. Byrd I-E-5 CR: Puntarenas, Pir V. trigonocarpa W. Stern s.n. PMA: Collected by R.L. Dressler in PanamaAbbreviations: BR = Brazil, CB = Cuba, CR = Costa Rica, FR = France, GD = Guadeloupe; GT = Guatemala, GY: Guyana, GYF: French Guyana HD = Honduras, MX = Mexico, PMA = Panama, PNG = Papua New Guinea, PR = Puerto Rico, SU = Suriname, UK = United Kingdom, USA = United States; F. = Finca; J. Bot. = Botanic Garden, RBG Kew = Royal Botanic Gardens, Kew.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla291been extracted from a couple of South American and Asian herbarium specimens of other species. The ITS region, including the 5.8S gene was then amplified with the primers ITS2, ITS3, ITS4, and ITS5 (White et al. 1990; Baldwin, 1992) or 17SE and 26SE of Sun et al. (1994). PCR reactions were better when 3% of DMSO was added to the cocktail. PCR products were run in low melting point agarose gel and the band cut to be extracted with QIAGEN Gel Extraction Kit (QIAGEN, Ltd.) or QIAquick silica columns (QIAGEN, Ltd.) adding guanidin chloride (35%) to remove primer dimers. Bi-directional sequencing was performed using cycle-sequencing (ABI Prism dye terminator cycle sequencing ready reaction kit, PE Applied Biosystems, Inc.) with the same primers with which they were amplified using different automated sequencers following manufacturer’s protocols. Electropherograms were edited using EditView, and the resulted sequences were initially aligned using Clustal x (Thompson, 1995) and adjusted by eye. Although alignment of Vanilla species is easy, the alignment with the outgroups is largely ambiguous and alignment is based mostly on the lengths of the regions. Phylogenetic analysis was performed with PAUP* 4.0 (Swofford, 1998) with the following specifications: Epistephium parviflorum and Lecanorchis multiflora were defined as the outgroups. An heuristic search with equally weighted parsimony analysis. The analysis consisted of 1,000 replicates with SPR swapping and 20 trees saved per replicate, to save time swapping on islands with large numbers of trees and allowing the detection of multiple islands of equally parsimonious trees (Madison 1991). Then another equally weighted parsimony analysis was performed, swapping to completion of all the trees from the previous analysis. This was followed by a bootstrap analysis, holding 10 trees per replicate for 1000 replicates, using SPR swapping and MULPARS. Bootstrap values were obtained from 100 replicates. Results We recognize 15 Vanilla species in Mexico and Central America. A key and a taxonomic treatment are presented in the following pages. Five species, Vanilla costaricensis , V. cribbiana , V. dressleri , V. martinezii , and V. sarapiquensis are proposed as new species and three subspecies are recognized within V. pompona (the typical subspecies, subspp. pittieri and ). primers 17SE and 26SE of Sun et al. (1994) and with them only a single band was obtained. Two bands, one probably from and endophytic fungus were The analysis of nucleotidic sequences resulted in 669 characters, from which 246 were constant, and 271 sites were parsimony-informative. The heuristic search yield a total of 39,090 most parsimonious trees (tree length = 1070). The strict consensus of this tree is presented in Fig. 1. This tree is largely congruent with phylogenetic analysis based on morphological and molecular characters, either of plastid ( rbcL gene, Soto 2003; matK of nuclear sequences (18S and ITS sequences from species from all the world). In all the analysis two main groups of Vanilla are recognized. One of them includes the Membranaceous vanillas ( V. inodora and V. martinezii in the tree; V. costaricensis and V. sarapiquensis , from which DNA was not available belong to this clade). The other lineage of Vanilla membranaceous species. Two very dissimilar sequences of V. planifolia were obtained from two different PCR reactions of the specimen Dressler s.n. (cultivated at FLAS). One of these sequences is practically identical to other eight sequences of the same species included in the sample. The anomalous V. planifolia sequence goes sister to a V. mexicana sequence, forming a clade which occupies a basal position to the rest of Vanilla . When Clematepistephium smilacifolium and Eriaxis rigida are included into the analysis, this clade occupies an intermediate position between ClematepistephiumEriaxis-Epistephium and Lecanorchis (data not presented). Vanilla mexicana is the type species of separate from, other membranaceous vanillas (e.g. V. martinezii and V. inodora ). These anomalous sequences make both Vanilla outgroup non-monophyletic . The rbcL (Soto Arenas,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.292 LANKESTERIANA FIGURE 1. Strict consensus from 30,090 minimal trees obtained from ITS sequences of the nuclear ribosomal DNA.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla2932003), matK, and 18S topologies (Soto Arenas et al., in prep) are incongruent with the position of these two particular sequences of Vanilla and we regard them as paralogous ITS copies. Cloning in 10 PGEM a PCR product of V . cf. kaniensis Schltr. (a species from New Guinea) yield three different ITS clones which differ only in 1 and 2 DNA bases and are clustered together in the ITS phylogeny, therefore ITS is a very useful identifying tool. Other paralogous ITS sequences of Vanilla (from species of other regions) have occasionally resulted, especially when DNA has been extracted from herbarium specimens or less often from tissues dried in silica gel. No more paralogous copies of American vanillas have been detected, and both included represent the two main lineages in the genus. If they Vanilla become monophyletic. The paralogous sequences This is probably because the duplication event that produced them seems to have been previous to the separation of Vanilla from their related groups. The leafless group is not well-documented from Mesoamerica (see discussion in ‘Excluded taxa’); a Puerto Rican specimen of V. calyculata and another of V. barbellata were included. There are no confirmed reports of these species from Mexico or Costa Rica, respectively; in any case, it is rather surprising that leafless taxa are absent in Yucatn, very close, and with similar habitats to places in Cuba where leafless vanillas are present. The rest of the Mesoamerican vanillas belong to the group recognized by Soto (2003) as “American fragrant Vanilla spp.” Phylogeny of Vanilla is discussed in more detail in Soto (2003). the species form monophyletic groups with very high bootstrap support. The only exceptions are V. odorata which proved to be paraphyletic, since V. helleri and V. tahitensis are nested in it. Two samples from east of the Tehuantepec Isthmus share a different ITS sequence. Vanilla tahitensis sequences are not different from most of V. odorata , a result which is not incongruent with a the hypothesis of a recent hybrid origin of this taxon (between V. planifolia and V. odorata ), as its morphology suggests. The Vanilla pompona complex has a bootstrap value of 99%. All the Mexican samples of V. pompona plus two specimens (SEL 1985-0211A and a sequence in the Gen Bank U66819) form a clade with 99% of bootstrap support. Several South American specimens of form a weakly supported (63% bootstrap value) branch, and the rest of the South American specimens of V. pompona and the Costa Rican, Honduran, and Panamanian specimens of the V. pompona complex form a polytomy at the base. This structure in the V. pompona complex is congruent with the distributional In conclusion, ITS sequences are a very useful tool to separate sympatric, vegetatively indistinguishable vines like V. planifolia, V. helleri, and V. cribbiana, or to identify specimens with juvenile or etiolated vegetative characters. We suspect that one of the reasons for the lack of an important breeding effort in Vanilla material. The origin of V. tahitensis has been much argued. These data indicate clearly that V. tahitensis is very closely related to V. odorata . Vanilla tahitensis is clearly a member of the American fragrant clade of vanilla, characterized by leafy plants and penicillate calluses, and which is absent in the Old World Tropics (as in New Guinea) where the vanillas geographically nearest to Tahiti occur), therefore the hypothesis that it was introduced recently to Tahiti is strongly supported. Since vegetatively it has much broader leaves than V. odorata , it is almost certain that it is than in V. planifolia , and it is rather intermediate in all traits between V. planifolia and V. odorata . These molecular data, and other unpublished data including many more species from other areas, suggest that V. phaeantha, V. insignis, V. helleri, V. odorata , and V. tahitensis are the closest relatives of the vanilla of the commerce, V. planifolia , even much more closely related than V. pompona with which it has formed hybrids that are at present cultivated. The close relatives all have fragrant fruits and have different habitat preferences; thus they represent a pool of interesting traits with potential to be incorporated into this crop.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.294 LANKESTERIANA KEY TO THE SPECIES AND SUBSPECIES remote bracts; lip united only basally to the column (< 4 mm), column smooth or basally keeled, but without substigmatic hairs; lip without a penicillate callus formed by a tuft of laciniate scales, instead, the callus may 3. Lip distinctly trilobed, midlobe emarginate-bilobed, callus massive, cushion-like V. inodora 3. Lip subentire, truncate at apex, callus made up of 2 prominent axial keels and several warty lateral keels V. costaricensis 4. Lip ca. 42 mm long, with low callus made up of several axial, longitudinal, warty keels V. martinezii keels V. sarapiquensis 1. bracts; lip attached to the column up to the stigmatic region, usually > 2.5 cm, column with trichomes on the ventral surface, lip with a penicillate callus formed by a tuft of laciniate, retrorse scales 5 5. Lip without a clear claw, rather cuneate, strongly trilobed, the midlobe short, transversely oblong and covered by conspicuous, complanate papillae along nerves V. helleri 5. Lip conspicuously long-clawed, claw usually very distinct from the blade, the latter entire to trilobed, the midlobe variously shaped, smooth or variously covered with papillae, trichomes or warts 6 (2-3) V. trigonocarpa 6. 7 7. Leaves conspicuously shorter than the internodes, lip almost smooth, without conspicuous appendages, except by minute warts at the apical part of the axial veins 8 8. Leaves acuminate; lip < 41 mm long, subentire V. hartii 8. Leaves obtuse to subacute, sometimes mucronate; lip > 60 mm long, trilobate at apex V. phaeantha 7. Leaves usually as long or longer than the internodes, lip apex covered with papillae, retrorse trichomes or the veins thickened, wavy and complanate-warty 9 9. Lip fringed or denticulate at margins, with an apical thickening with warts, papillae, or retrorse V. insignis 10. erose or denticulate, midlobe with few (less than 5) retrorse papillae or warty 11 V. odorata 11. Leaves oblong to elliptic, acute or abruptly acuminate; lip margin undulate-denticulate, V. planifolia 9. Lip entire or undulate at margins, without apical thickenings or if present without warts, papillae, 12 V. calyculata

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla295 V. dressleri 13. 14 14. Stems 3-5 mm thick, leaves 10-22 x 2.3-7.5 cm; lip apex truncate, entire, ovary and sepals fairly papillose-verrucose V. cribbiana 14. Stems 10-24 mm thick, leaves 22-29 x 8-14 cm; lip margin undulate, ovary and sepals smooth V. pompona TAXONOMIC TREATMENT 1. Vanilla calyculata Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 42-43. 1920. TYPE: Colombia, Cauca: 1000 m M. Madero, not located; (here designated): VALLE: Municipio de Tula, Corregimiento Mateguadua, Jardn Botnico, laderas en va de repoblacin natural. Altura 1100 m. Enredadera; spalos verde claro, ptalos amarillo claro, labelo amarillo intenso, frutos maduros color marrn; muy fragantes. 29 sept. 1984. W. Devia 815 holo. MO (3245054)! COMMON NAMES : “Vanilla” Hemiepiphytic vine, branching, leafy, up to ca. 4 m high. Stems internodes 7-15.5 cm long. Aerial, free roots apparently subterete, pale brownish, up to 8.2 cm long, 2-3 mm thick. Leaves sessile to subpetiolate, the petiole up to 7-10 mm long; the blade oblong-lanceolate, rather narrow, the base rounded, the apex acute, coriaceousslightly revolute, 7.5-21 x 1.7-4 cm. terete, green, ca. 4-6 mm thick. Bracts widely ovate, obtuse, green, concave, progressively smaller, up to 10 x 6 mm. Flowers opening successively, 2-3 open at once, pendant, bell-shaped, very showy, sepals pale green to yellowish green, petals pale yellow, lip deep yellow, ca. 9 cm long; strongly fragrant. terete, smooth, sulcate, the grroves twisted, ca. 4-4.5 mm long, 4 mm thick. Dorsal sepal oblanceolate to narrowly elliptic, apex subacute-obtuse, somewhat thickened, subcalyptrate, recurved, base attenuatesubunguiculate, basally canaliculate, convex above, smooth, ca. 12-13-veined, 66-80 x 8-17 mm. Lateral sepals oblong to narrowly elliptic, oblique, the lower margin more curved, apex subacute-obtuse, somewhat thickened, subcalyptrate, recurved, basally attenuatesubunguiculate, smooth, ca. 12-13-veined, 67-80 x 12-17 mm. Petals obliquely oblanceolate, lower margin more arcuate, apex attenuate, rounded, very abaxial surface, 1 mm wide, ending in a free, conic, blunt, short, 1 mm long process; ca. 11-12-veined, 65-80 x 9-16 mm. Lip attached to the column along the margins of the basal half (ca. 41-46 mm), tubular, cymbiform, deeper near the middle; axially grooved on the abaxial surface, and deep, when spread out 78-90 x 36-45 mm; long unguiculate, the claw apically rugose-papillose on the inner surface, 26 x 3-3.8 mm; the blade trilobed, ca. 36-veined, veins branched, the lateral lobes forming an triangular-semiovate, margins entire, undulate near the midlobe, ca. 37-39 x 14 mm; midlobe dilated and recurved, approximately transversely oblong, deeply emarginate-bilobed, margins crenulate-undulate, veins conspicuously thickened, 14-16 x 21-26 mm; penicillate crest at ca. 41 mm from the base, 4.5 x laciniate, retrorse, scales, some of them united to each other along the lateral margins; disc with progressively more thickened veins from the center to the apex of the lip, forming an apical, swollen, low cushion, obclavate in outline, rugose-papillose at the apex, 25-27 x 6 mm, 2-3 mm high; with a group of transversal, yelloworange hairs at the basal part. Column very elongate and slender, subtrigonous-subclavate, slightly sigmoid, and lanuginose at the distal half; apex dilated (6 mm undulate, ca. 4 x 1 mm. Stigma trilobed, the lobes x 4.5 mm; x 1.8 mm. Anther versatile, attached to the wide clinandrium emarginate, 5 x 5 mm. Fruit fragrant, 8-15 cm long, apparently thick and trigon ous. Fig. 2, 17A.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.296 LANKESTERIANA FIGURE 2. Vanilla calyculata Schltr. Based on published pictures (Hamer, 1981) and Devia 815. Drawing by R. Jimnez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla297 DISTRIBUTION Honduras, Costa Rica, and Colombia; a fruiting specimen from Michoacn, Mexico, may belong here. ECOLOGY : Hemiepiphyte at relatively high altitudes (800-1300 m), often in rather dry places. Flowers recorded in April (El Salvador and Honduras) and September-February (Colombia). The identity of V. calyculata has been obscure and no original specimens are known to us. However, Schlechter’s description can be applied with in Fig. 1. No other species known from Colombia (V. columbiana, V. dressleri, V. espondae, V. hostmanii, V. methonica, V. odorata, V. pompona, V. ribeiroi, and V. sprucei) agrees with the description of V. calyculata. a minute, cup-like structure sometimes distinguished in some vanillas and more developed in the species of the related genera Epistephium and Lecanorchis . The calyculus in V. calyculata is rather conspicuous when compared with other vanillas, and is more evident in pollinated ovaries that start to swell. Schlechter’s description differs from our material of V. calyculata in the column described as glabrous instead of pubescent below the stigma (a trait found in all American penicillate Vanillas) and its smaller measurements of the perianth segments. We believe that Schlechter’s description could had been based on a bud. However, the calyculus, the sessile, oblong, rather small leaves, the deeply excised midlobe of lip, the pubescent lip claw, the 3-5 thickened midveins on the disc, the unusually high altitude for a Vanilla and its strong resemblance with the Brazilan V. chamissonis (its sister species, Soto unpublished) suggest that the name V. calyculata may Vanilla calyculata is distinguished from V. chamissonis by its larger, bell-shaped, pendant, longer midlobe of the lip. Despite its unique characters, specimens of V. calyculata have been regarded by Reichenbach and Rolfe (1896; e.g. Lehmann 2263) as V. pompona. Later it was reported from El Salvador (Hamer, 1974) as V. phaeantha Rchb.f. Jos Linares, from Escuela Agrcola Panamericana has told us that this species is abundant, well-known, and usually confused with V. pompona in the dry valleys of central Honduras. The specimen O. Pank sub F. Hamer 203 bears two very odd stigmas. OTHER RECORDS : EL SALVADOR: La Palma-Finca El Refugio, ro Nanuapa, 1000 m alt., terr. climbing oaks or manzana rosa, collected 3.4.1969. April 11 1969, O. Pank y F. Hamer 203 AMES(*112862; *113837)! HONDURAS : COMAYAGUA : Fruit triangular, up to 6 in. long. Climbing in shrubs, thicket along river, plain near Sihuatepeque. 1050 m altitude. 7/23/36, *AMES(46667)! G! K! *MO(1115382)! NY! abajo de la Quebrada de Sta. Clara. Alt. 850 m, Abril 28, 1948, A. Molina 808 *F(1676153)! Bejuco, creciendo (por la tarde). Quebrada Santa Clara, ca. 2 km al norte del de junio de 1996. J. L. Linares 3386 MEXU! El PARAISO : Mpio. Morocel, Quebrada El Cajocote conocida tambin como El Terrero, 8.7 km al N de Morocel, por el camino a Mata de Pltano. Veg. riparia, veg. circundante selva baja caducifolia y bosque de pino-encino, 14 10’’ N, 86’’W, 680 m. Bejuco creciendo sobre Guettarda oro, olor suave y dulce, 29 abril 2004, J. L. Linares 7313 MEXU! [cf., sterile] “Vainilla”, trepadora sobre rboles y matorrales, de Barranco de las Mesas, Alt. 900 m, Agosto 1, 1957, F(1590341)! Open savannah, Las Mesas region near Yuscarn. August, 1960, H.W. Pfeifer 1454 *US(2563382)! COLOMBIA: TOLIMA dunkelgrn. Blt. grnlich meist mit heller Lippe. Frchte Plata. 800-1500 m. 3.12.1882. F.C. Lehmann 2263 BM! G! Vanilla , white and red inside, cultivated E. Dryander 2379 BM! “Columbien” Lehmann and probably the right shoot)!The following sterile specimens may belong here: MEXICO: MICHOACAN : Hacienda de Coahuayula, Feb. 1901, G.M. Emrick 16 F(95476)! COLOMBIA: [cf., leaves atypically broad] TOLIMA: La Plata, 1000-1300, *Lehmann 6278 AMES(14875)!2. Vanilla costaricensis Soto Arenas, sp. nov. TYPE: COSTA RICA : ALAJUELA: Llanura de San Carlos. Alt. 200 m., Feb. 20, 1966. Lip white, tepals same green as leaves; vine. Lowland rain forest between Los Chiles and Venecia. A. Molina R., L.O. Williams, W.C. Burger and B. Wallenta 17565, holo. CR(062310)! iso. MO(2367689)!

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.298 LANKESTERIANA labello integro breviore differt. Hemiepiphytic vine, leafy. Stems keeled, 8 mm thick (in dried condition); internodes ca. 12.5 cm long. Aerial, free roots dorsiventraly compressed, pale brownish, ca. 2.5 mm wide. Leaves petiolate, the petiole canaliculate, ca. 17 mm long, 9 mm wide; blade elliptic, abruptly mucronate, base acute to attenuate, green, membranaceous (in dried condition), 21-22 x 8.7-9.5 cm. similar to the vegetative developed), 28-32 cm long, internodes up to 6 cm long; peduncle 15-23 cm, rachis ca. 3 mm thick. Bracts foliaceous, subpetiolate, the petiole ca. 3 mm long; blade elliptic, acute-acuminate, base obtuse-rounded, membranaceous, 2.9-7.3 x 1.6-3.4 cm. Flowers successive, 2 open at once, with spreading segments, ca. 3.5 cm in diameter; tepals green, lip white. rather sigmoid, 34-40 x 3.5 mm. Dorsal sepal strongly twisted and the margins contorted-widely undulate, elliptic, apex acute, rounded, base cuneate, ca. 10 veined, smooth, 41 x 11 mm. Lateral sepals narrowly elliptic, acute-subacute, base obtuse, margins undulate, x 10.5 mm. Petals narrowly elliptic, apex truncate, notched, base widely cuneate, strongly twisted, widely undulate-contorted, without distortion, ca. 10 veined, 35.5 x 9 mm. Lip attached to the column less than 2 mm, slightly arcuate, almost straight, with the lateral margins erect forming apex truncate, obcurely trilobed, the lateral lobes as long as the midlobe, or slightly longer, base roundedtruncate, the apical margin entire to dentate, ca. 26 veined; 24-27 x 20-21 mm; callus made up of a pair of the begining of the apical third, then separated in 3 low, erect, congested keels ending at the apex; additional lateral keels with complanate warts adorn the basal half. Column relatively short, straight to very slightly arcuate, semiterete, with a dilated apex, 16 mm long; smooth, except by the callus of verrucose keels at base, x 3 mm. Stigma lobed, the midlobe, convex, very prominent, almost perpendicular to the column axis, ca. 2 x 1.5 mm; lateral lobes much smaller, fused in an ovatesubquadrate, emergent blade, ca. 0.5 mm long. Anther galeate, protruding at apex, laterally compressed, 4 mm DISTRIBUTION : Known only from northern Costa Rica. ECOLOGY : In lowland rain forest at 200 m altitude; This species is known only from the type locality V. mexicana Miller, a different, closely related species from the West Indies and northeastern South America. Vanilla mexicana has a distinctly trilobed lip, with the apex of the lateral lobes slightly above the middle of the lip; in V. costaricensis truncate and just obscurely trilobed, but the lateral lobes of the apex are subequal or slightly longer than the midlobe; additionally, the keels at the apex of the lip in V. mexicana are much more conspicuous. Vanilla costaricensis has large bracts similar to those of V. inodora Schiede, easily distinguished by the cushionlike, axial callus on the lip. Also closely related are V. guianensis and V. martinezii with much larger illustrated by Dr. Rafael Lucas Rodrguez (Rodrguez C. et al., 1986) based on a specimen that has not been located. The closest relative of Vanilla costaricensis is V. oroana Dodson of Ecuador. They have similar column almost identical. However, V. oroana has a longer, relatively narrower, trilobed non-truncate lip, and the surface of the lateral, basal part is minute, but conspicuously more sculptured than in V. costaricensis. Vanilla oroana In the examined material of V. costaricensis (and also V. oroana ) the lateral lobes of the stigma are fused to each other to form an ovate-subquadrate, emergent blade. This is notoriously different from the concave stigma found in other membranaceous (e.g. V. inodora ). of the poor preservation of the column, and the large amount of sticky substance in the stigmatic area, visible in fresh material but absent in pressed specimens. OTHER RECORDS : COSTA RICA : ALAJUELA: Llanura de San Carlos. Alt. 200 m., Feb. 20, 1966. Lip white, tepals same green as leaves; vine. Lowland rain forest between Los Chiles and Venecia. A. Molina R., L.O. Williams,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla299 FIGURE 3. Vanilla costaricensis Soto Arenas. Based on A. Molina et al. 17565 based on A. Molina et al. 17567. Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.300 LANKESTERIANAW.C. Burger and B. Wallenta 17567 *SEL(016793)! *F(1771938)!3. Vanilla cribbiana Soto Arenas, sp. nov. TYPE: MEXICO: CHIAPAS: Mpio. Ocosingo: Estacin de Biologa Chajul, en el borde del Ro Lacatm; a 680 m del Puente Hamaca, por la vereda a Arroyo Miranda, ca. 180 m s.n.m., selva con tpalos blanco-cremosos, spalos verdosos en intenso con rayas anaranjadas en las venas, labelo y conspicua, a mentol y ctrico. Ovario verdeblanquecino con papilas. 20 junio 1996, M. Soto 7945 y R. Solano, holo. *AMO!; iso. K !, AMO (in spirit)! Vanillae hostmanii primo adspectu similis, sed foliis labello obscure 3-lobato, lobo medio distincto, subquadrato, quinque nervis papillosis incrassatis in centro apice ornato versus circa decem nervos in lobos laterales extensos COMMON NAMES : “Vainilla”. Hemiepiphytic vine, somewhat branching, leafy, up to 12 m high. Stems terete, smooth, green, 3-5 mm thick; internodes 3-9 cm long. Aerial, free roots pale subterete, brownish, ca. 2 mm thick; attaching, aerial roots conspicuously dorsiventrally compressed, 2-5 mm wide. Leaves subpetiolate, petiole 1-1.5 cm long; blade obliquely elliptic to elliptic-oblong, abruptely acuminate-apiculate, stiff, brittle, 10-22 x 2.3-7.5 cm. mm long, 4 mm thick. Bracts ovate-triangular, obtuse, very concave, progressively smaller towards the apex, up to 8 x 4 mm. Flowers successive, 2-3 open at once, with spreading segments, apparently ephemeral, very showy, white-cream tepals, sepals externally cream-greenish, lip deep yellow with orange stripes, ca. 8 cm diameter; fragrance strong, mentholate and citric, similar to that of V. pompona. arcuate, dorsiventrally slightly compressed, thickened at the very base, whitish-green, conspicuously papillose, more densely towards the perianth, sulcate, the grooves twisted, 44-46 mm long, 4-5.5 mm thick. Dorsal sepal narrowly elliptic, apex subacute-rounded, apically concave, 9-veined, minutely papillose on the abaxial surface, the papillae in longitudinal rows and stiff; 63-65 x 12.5-13.5 mm. Lateral sepals obliquely oblong-elliptic (upper margin curved, lower margin more straight), apex subacute-rounded, subcalyptrate, base attenuate, canaliculate basally, slightly convex, apically concave, apex somewhat recurved, margins conspicuously involute, ca. 12-14-veined, minutely papillose in the abaxial surface, the papillae in longitudinal rows and forming transverse, undulate x 12.5-14 mm. Petals oblanceolate, very slightly oblique (the lower margin more straight), slightly arcuate, apex obtuse, widely rounded, somewhat thickened at apex, base attenuate, basally canaliculate, slightly concave free (ca. 2 mm) process; dorsal surface conspicuously canaliculate apically, surface colliculate, the cells in longitudinal rows, 14-veined; with granular, threadlike, somewhat branched inclusions; 64 x 12.5-13.5 mm. Lip attached to the column along the margins of the basal half (ca. 28 mm), tubular, trumpedshaped, cymbiform, deepest near the middle; axially grooved on the abaxial surface; when spread out 5052 x 38 mm; unguiculate, claw with 4 obscure bands of unicelluar, minute, elongate, yellow-brownish, margin conspicuously entire, subtruncate, lip with inclusions similar to those found in the petals; lateral lobes widely and obliquely triangular, overlapping above the column, 33 x 16 mm; midlobe small, x 14 mm; penicillate callus made up by ca. 10 congested, retrorse, trapezoidal, laciniate scales, the scales sometimes united to each other along the lateral margins, ca. 5 x 4 mm; the region just after the penicillate callus, smooth, with denser inclusions; with 5 low, rugose, rounded, densely papillose keels secondary, more inconspicuous keels, shorter, not reaching the apex. Column elongate, 34 mm long,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla301 FIGURE 4. Vanilla cribbiana Soto Arenas. Based on M. Soto 7945. Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.302 LANKESTERIANA3 mm wide; ventral surface pubescent-lanuginose at the middle, hairs yellowish; apex dilated, 4 mm wide; vertical wings trapezoidal, the lower margin projected and acute, ca. 2 x 4.3 mm. Stigma trilobed, the lobes emergent, with a membranaceous, convex rostellum, 4 mm wide; lateral lobes subquadrate, rounded, ca. 1.2 x 1.2 mm. Anther versatile, articulate to the wide, convex, clinandrium, ovoid-subquadrate, 3.5 x 3 mm. Fruit short, thick, trigonous in cross section, 10-16 cm long, 1-1.4 cm thick; fragrant when ripe, aroma similar to common vanilla, but weaker. Fig. 4, 16A. DISTRIBUTION : Known from Mexico (Oaxaca and Chiapas), Guatemala, Belize and Honduras. ECOLOGY : Hemiepiphyte in rain forest on soils with variable drainage in areas with 2500-4000 mm of rainfall, at 150-350 m altitude. It is the most common Vanilla in many areas with tropical rainforest of the Selva Lacandona (Chiapas) and the Petn (Guatemala). It is the only penicillate Vanilla in N Central America blooming during the rainy season, in July and August. Vanilla cribbiana is a member of the V. hostmanii group. Like the other members of this complex, it has a slightly trilobed to subentire lip with a disc adorned by some thickened veins. It is different from V. hostmanii Lindl. from Amazonia, probably its closest relative, the lip, and only ca. 5 thickened, papillose veins in the center of lip apex, not ca. 10 and spreading to the lateral lobes. From Vanilla dressleri Soto Arenas from Costa Rica to Colombia, it is distinguished by lip, less conspicuous thickened veins on the disc, papillose-granulose outer surface of the sepals and broader, thicker leaves. Vanilla ruiziana Klotzch (synonym V. weberbaueriana Kraenzl.) from Peru and Bolivia has a larger, ovate midlobe of the lip which bears 1-3 elevated, axial keels near the apex and the tepals are acute and the sepals neatly calyptrate. The the rainy season, anf it is also distinct. Vegetatively it is easily confused with V. planifolia but the leaves of V. cribbiana are usually more elliptic, basally more attenuate, and acuminate at the apex. However, the available, and especially in sterile shoots grown in shade. Therefore, Vanilla cribbiana has been confused with V. planifolia, and also with V. pompona and even V. inodora, in the herbaria and in the plantations. The fruits of this species are fragrant, and probably of some potential commercial interest, but the vine is in the plantations of northern Oaxaca, where it was introduced when confused with V. planifolia (Prez Eulaema bee. Beza (com. pers.) reports that the fruits of this species have been cured in Guatemala. OTHER RECORDS : MEXICO: OAXACA: cultivated by J. Prez Mesa at Instituto Tecnolgico Agropecuario # 3, San Bartolo Tuxtepec, from plantation in Jaltepec de Condoyoc, near Ma. Lombardo, 24-IV-1997, J. Prez sub M. Soto 8504 AMO(in spirit)! CHIAPAS: Mpio. Ocosingo, Estacin de Biologa Chajul, ca. 200 m s.n.m., selva alta perennifolia despus de la sabana II, 19 junio 1996, M AMO(sterile)! Chajul, Camino a Arroyo Miranda. Fruto trgono, ca. 16 cm de largo, 14 S. Sinaca sub M. Soto 7953A MEXU! Mpio.Ocosingo, Estacin de Biologa de Chajul, camino a la Sabana I, selva alta perennifolia en loma, ca. 230 m s.n.m. 13-IV-1997, M. Soto 8387 *AMO(sterile)! Mpio Ocosingo, 1.5-2 km al SW de la Colonia Benito Jurez Miramar, sobre la desviacin a Tierra y Libertad. Acahual Bursera simaruba, Vochysia hondurensis, 360 m s.n.m., 20N, A. Reyes-Garca y M. Sousa 2029 MEXU(584270, 584313)! Mpio. Ocosingo, entre Bonampak y el Ro Lacanj, selva alta perennifolia con Dialium guianense, Brosimun spp., Ficus glabrata y Terminalia amazonia, 300-350 m s.n.m., abril 2000, M. Soto 9617, S. Maldonado, P. Schltter, L. Lpez AMO(sterile)! [cf., fruit apparently sulcate] a 2 km del Crucero Corozal, camino Palenque-Boca de Lacantm, selva alta subperennifolia, 13 feb 1985, E. Martnez 10299 MEXU! Mpio de Ocosingo, carretera Palenque-Marquz de Comillas, Crucero San Javier; selva alta-mediana subperennifolia, inundable, perturbada con Chrysophylla, Vochysia M. Soto 8438-8440 AMO(sterile)! GUATEMALA: PETEN: Fleshy epiphytic vine. Canchacn, in high rain forest of southeastern Petn. July 14 1959. C.L. Lundell 16457 *LL(x2, buds)! MO(3832548)! “Vianilla”, vine, fruits black, fragrant, Dolores, in low forest of pinal about 800 m south of the village on the Machaquila Road, May 18, 1961, E. Contreras 2333 MEXU(511605)! NY! LL(fruit)! LL(buds)!

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla303 “Vainilla” Fleshy vine, fruit green; Dolores, on Ro Mopan trail, in high forest, October 17, 1961, E. Contreras 3063 LL(fruit)! BELIZE: STANN CREEK yellow. In high ridge on hill top. Middlesex, 2 July 1939. P.H. Gentle 2894 *AMES(58082)! K! LL! NY! TOLEDO: or fruiting, stiff, thick leaves, 1973, C. Whitefoord 1816 BM! HONDURAS: ATLANTIDA: Near Tela. Guaymas. Clambering over tree. March 17, 1923 O. Ames II 211 AMES(36945, fruit)!4. Vanilla dressleri Soto Arenas, sp. nov. TYPE: PANAMA. COLON: End of Pipeline road on Ro Agua Salud, 9-10 mi N of Gamboa. Tropical wet forest. Elev. 20-50 m. Vine. Flowers greenishyellow, lower lip white. Column yellow. Sweet smelling, 15 April 1982, (holo. MO 3032052!). longioribus, labello longiore subintegro anguste obtuso differt. Hemiepiphytic vine, leafy. Stems apparently terete, grooved, 3-4 mm thick (in dried specimens); internodes 7.5-11.5 cm long. Aerial, free roots terete, 2 mm thick; attaching, aerial roots strongly Leaves petiolate, the petiole canaliculate, 11-18 mm long, ca. 5 mm wide; the blade oblanceolate, obovate to narrowly elliptic, much larger than the internodes, sometimes slightly oblique, basally rounded to attenuate, apex acuminatemesophytic, conspicuously pendant in living condition, 10.5-21.5 x 3.0-6.5 cm. raceme, rather lax, contiguous bracts distant up to 11 mm; peduncle 15-35 mm long, rachis 15-43 mm long, at least 4 mm thick. Floral bracts patent, spreading, concave, widely ovate to elliptic, obtuse, up to 14 x 8 mm. Flowers successive, 1 open at a time, big and showy, tepals white to greenish-yellow, lip orangeyellow in the inner surface, fading to white, with brownish veins, column yellow; fragrance clove-like. subterete, straight, thickened towards the apex, bisulcate, the grooves twisted, very inconspicuously and minutely papillose, 42 mm long, 4-5 mm thick; sometimes slightly calyculate. Dorsal sepal linear to narrowly elliptic, apex obtuse, rounded, subcalyptrate, thickened, with 2-3 warts at the apex of the outer surface; base narrowed, canaliculate; conspicuously keeled on the outer surface, blade apparently x 11.5 mm. Lateral sepals obliquely linear to narrowly elliptic, apex obtuse, rounded, calyptrate, thickened, inconspicuously warty at the apex on the outer surface, base narrowed, concave, blade ca. 11-veined; obscurely x 12 mm. Petals linear to narrowly elliptic, slightly oblique, apex widely rounded, base narrowed, basally concave, the keel ending in an acuminate, laterally compressed process, ca. 2 mm long, apex strongly grooved, ca. 10 veined; 68 x 13.5 mm. Lip attached to the column along the margins of the basal half (ca. 28 mm), tubular, apparently concave, axially grooved on the abaxial surface; when spread out 65 x 39 mm; unguiculate, the claw canaliculate, lanuginose becoming pubescent towards the blade, 15 mm long, 5 mm wide near the blade; blade subrhombic, margin undulate, pleated, apex obtuse-rounded, very slightly notched, ca. 24-veined; the ca. 11 veins of the apical part thickened ones rather elevated; the region corresponding to the lateral lobes ca. 36 x 15 mm; the wanting midlobe 10 x 15 mm; penicillate callus at 28 mm from the lip base, made up by ca. 12, obtriangular, basally long attenuate, x 2.5 mm. Column elongate, rather straight, ca. 34 mm long, 3 mm wide; densely pilose on apical 2/3 of the ventral surface, below the stigma; apex dilated, vertical wings oblong, lower apex acute, ca. 1.5 x 3 mm. Stigma trilobed, the lobes emergent, rostellum strongly convex, lateral ones apparently very narrow. Fig. 5, 16B. DISTRIBUTION : Known from Costa Rica, Panama and Colombia. ECOLOGY : In lowland to submontane (20-1000 m), wet forest. Flowering from March to early June. The species seem to be fairly common, at least in the Ro Savagre area of W Costa Rica. The clove-scented ETYMOLOGY : Dedicated to Dr. Robert L. Dressler, authority in orchid evolutionary biology and in orchid V. hostmanii

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.304 LANKESTERIANA FIGURE 5. Vanilla dressleri Soto Arenas. Based on G. McPherson 9196. Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla305 sc arcely trilobed lips, granulose ovaries and sepals, and the dist al veins of the disc thickened. Among the group, lip, with the blade subrhombic in outline, and very scarcely pappilose sepals; the other members of the group, V. cribbiana, V. hostmanii and V. ruiziana have conspicuously granulose sepals. From V. cribbiana it is additionally distinguished by the narrower, thinner leaves which are more long acuminate-cuspidate, more lax in March-May, not in July-August. Vanilla hostmanii thicker, but not as strongly acuminate-cuspidate. Vanilla ruiziana , from Peru and Bolivia has larger, broader, less acuminate-cuspidate, and thicker leaves, but is similar to V. dressleri in having rather elevated axial keels of the lip. Vanilla gardneri from Brazil has a subacute lip apex and oblong leaves, not cuspidate. Central American specimens of V. dressleri have V. planifolia and V. insignis, distantly related species, easily separated by the distinctly trilobed lips with well developed retrorse warts.O THER RECORDS : PANAMA : Flowers yellow green; lip yellow with green patch at apex. Collector, N.H. 56-76-11[accession num.], *A. Pridgeon s.n. SEL(019244; V. phaeantha)! COLON: Santa Rita Ridge, southeast of Coln, c. 9’N, 79’W. Along ridge road, 10-12 miles from Transisthmian Highway, c. 550 m. Vine; perianth white, the lip orangeG. McPherson 9196, holo. MO(3432756)! COCLE: Huge plant purchased in the El Valle market. Flowers pale yellowgreen. 6 March 1976. Taylor 768 *SEL(009187)! COSTA RICA : ALAJUELA: amarillo. 7 June 1988. CR INB! 10’N 84’W, 820 m. Bejuco. Flores blancas. 10 June 1987. INB! Upala, San Jos, Villa Nueva, 11 km al noreste de San Jos. 10’N, 85’W, 40 m. Terrestre, trepadora, escandente. Flores en botn verde amarillento. 18 April 1988, *G. Herrera 1840 CR MO(3864177)! CARTAGO: 12 km S of Turrialba by air, 4 km SE of Pejibaye along Ro Gato. Disturbed primary forest along river. Vine. 700 m alt. 9’N, 83’W. Vine.16-17 April 1983, *R. Liesner 14435 MO(3102177)! SAN JOSE: Sepals and petals greenish yellow. Lip cream-colored with greenish-brown raised lines. Callus in middle of lip raised and with rows of attached hairs or brushes. Vine. Parque Nacional Braulio Carrillo; Carrillo station, 300-600 m, 31 May-5 June 1980, *C. Todzia 1291 LL! HEREDIA: Sep column loosly, interior of lip and base yellow, apical 3rd fading to white, leaves suculent, stems single, not branched. Sarapiqu: (April) .... The lip has been apressed .., do not anthesis, and they have no perceptible odour at 10:30 a.m. Leaves drooping apparently from a rather slender stem, April 1959, *C.H. Lankester 1746 SEL (011365)!. LIMON: Reserva Biolgica Hitoy Cerere, Valle de la Estrella. Bosque con blanco, 4 December 1990, G. Carballo 325 INB! PUNTARENAS de Coto Colorado. Estacin Esquinas: Seccin Esquinas. 08N 83W, 100 m. Bejuco, frutos verdes. 17 July 1993, INB! COLOMBIA: VALLE: Ro Calima (regin del Choc); blanquecinas, 19 febr.-10 mar. 1944, *J. Cuatrecasas 16550 AMES(71363)! Cordillera Occidental; vertiente occidental: Hoya del ro Anchicay, lado derecho, bajando a La planta, bosques, 200-350 m alt.. Bejuco herbceo, crasisculo; tpalos ocrceo blanquecinos; labelo amarillo claro. 27 sept. 1943, *J. Cuatrecasas 15220 AMES(71364,71365)! CHOCO: Parque Nacional de Utra: Colecciones realizadas en la serrana ubicada al este de la ensenada de Utra, en un Elida (Mesica) siguiendo la trocha que conduce al acueducto cruzando por la quebrada Aguada. 6N, 77W, 0-100 F. Garca C. MO(3878341)! CHOCO(not seen). WITHOUT LOCALITY: No collection data: Climbing ridge along center; lip white with crenulate, entire margins, inner throat with yellow and brown lines; column yellow; sweet, spicy fragrance. 6 May 1992, *S.W. Ingram 1421 SEL(066924)! same data *S.W. Ingram 1422 SEL(066925)! 5. Vanilla hartii Rolfe, Bull. Misc. Inform. Kew: 133. 1899. TYPE: Trinidad, Cabasterre Arima, Hart 6355, holo. K!, iso. AMES(67785)! V. leprieurii R. Port., Bull. Soc. Bot. France, 98: 94. 1951.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.306 LANKESTERIANAType: “Guyane Franaise: Cayenne, dans les forts humides, Leprieur (s.n.), en 1846” (not seen, drawing!) COMMON NAMES : “Vanilla”, “vainilla”. Hemiepiphytic vine, branching, leafy, relatively weak, up to 3 m high; perhaps up to 8-10 m long. Stems terete and smooth, green, 3-6 mm thick; internodes 7.012.4 cm long. Aerial, free roots terete, pale brownish, 8-12 mm long, 1-2 mm thick; attaching roots up Leaves usually slightly shorter than the internodes; conspicuously petiolate, the petiole straight or twisted, canaliculate, with a clear junction line with the blade, ca. elliptic, acuminate, rounded at base, coriaceous, green, 9-11 nerved (and a similar number of smaller veins intercalated); 6.0-11.5 x 1.4-3.5 cm. a 3-3 mm thick; peduncle of 4-7 internodes; rachis 5-15 mm long. Bracts sessile, variable, triangular-ovate, acuminate to obtuse (the upper ones), concave (the upper ones) to cymbiform, not very stiff, progressively smaller, 3.5-9 x 2.5-5 mm. Flowers successive, 1-2 open at a time, segments spreading, ephemeral, but at least some of them remaining for more than one day, relatively small, tepals white, the sepals tinged with green, lip white with grayish or brown faint lines on the throat and an inconspicuous green cushion at lip apex; ca. 3.8 cm high; fragrance imperceptible. terete, green, smooth, 2.7 cm long, 1.2 mm thick. Sepals fused at base ca. 2 mm, subunguiculate and canaliculate at base, smooth. Dorsal sepal oblanceolate, apex acute, rounded, slightly incurved, 12-veined, 39-49 x 6-8 mm. Lateral sepals obliquely oblanceolate, the lower margin more arcuate, apex obliquely subacute, rounded, thickened, slightly subcalyptrate, and obscurely warty, 12-veined, 39-46 x 7-8.6 mm. Petals obliquely oblanceolate, somewhat arcuate, the lower margin more straight, long attenuate at base, apex subacute to widely rounded, ca. 12-veined, the abaxial surface, ending in a cylindric, acute, attached process, 39-50 x 6-8 mm. Lip attached to the column margins ca. 24-26 mm, long tubular, trumpet-shaped, basally gibbous, with an abaxial longitudinal groove; when spread out 37-43 x 14-25 mm; the claw slightly sigmoid, ca. 16 mm long, minutely papillose-pubescent; blade cymbiform, ca. 7 mm depth, subentire or entire, constricted near the apex and appearing slightly trilobed, basal half with inconspicuous rows of papillae up to the penicillate callus, distal part with thickened veins, inconspicuously papillose, the axial ones more prominent and forming an apical inconspicuous cushion, margins denticulate-undulate, plicate, apex truncate; penicillate callus at 26-30 mm from the lip base; ca. 4 x 3 mm, laciniate scales. Column very elongate and slender, semicylindric, 32-33 x 2-2.5 mm, apex dilated ca. 3-4 granulouse inclusions, ca. 3 x thick, ca. 1 mm long, 2 mm wide. Anther versatile, obovoid, ca. 3 x 2 mm. Stigma trilobed, the lobes broad; lateral lobes emergent, ovate-quadrate, tongueshaped, ca. 1 x 1 mm. Fruit linear-fusiform, elongate, along 2 lines; fragrant, characteristic vanilla aroma, very sweet; 92-140 x 4-5 mm. Fig. 6, 16C. DISTRIBUTION : Mexico (Chiapas, perhaps reaching eastern Oaxaca), Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Trinidad, Guyana, French Guiana, and probably Brazil. ECOLO G Y : Rare in wet forests at low altitudes, usually growing on understory treelets, in deep shade. Flowering time January to April. The Mexican population of V. hartii is sympatric with V. cribbiana, V. insignis, V. odorata, V. planifolia , and V. inodora , but V. hartii is open understory with tree ferns and large sedges; the other vanillas are absent from this habitat although they may be found a few meters away. Female Euglossa bees know if they are effective pollinators. Vanilla hartii is a spring of 2000, and then in spring of 2002. by Correll (1965). The Central American material of V.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla307 FIGURE 6. Vanilla hartii Rolfe, based on M. Soto 9729-9731. Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.308 LANKESTERIANAhartii vs. 32 mm long) and the lip apex was described originally as acute to subobtuse, not obtuse to widely rounded. pictures from Trinidad and Mexico show that they are due to the herborization process. Three collections of V. hartii vegetatively stouter than Mexican and Trinidadian We do not know if these differences may indicate that they should be regarded as a distinct taxon; although molecular data suggest that they are closely related (see ITS analysis). Vanilla leprieurii is here considered as a synonym of V. hartii. Portres (1954) stressed that his V. leprieurii was distinct because the nervation in the center of the lip was denser, with the zone before the nervation thicker, more imbricate scales in the penicillate callus, and the apex of the lip more emarginate, apiculate and pleated. Furthermore, the lip looks constricted, almost trilobed in the published drawing. All these features are variable in the material examined and again, they were pressed. Vanilla hartii is a relatively small vine similar to the V. planifolia group. It is somewhat similar to V. odorata but has shorter racemes, much smaller like thickening at the lip apex without retrorse papillae (vs. the 3-4 longitudinal rows of apical, retrorse, big papillae) and broader and shorter elliptic leaves (vs. long triangular-ensiform). The elliptic leaves, shorter than the internodes, and the slender stems make it vegetatively similar to V. bicolor from the Caribbean and Guyanas. However, V. bicolor has acute to Most Nicaraguan and several of the Central American specimens previously assigned to V. planifolia are actually V. hartii. Also the reports of wild V. planifolia from Rio Palenque Center in Ecuador are based apparently in V. hartii (P. Lubinsky, com. pers.). Vanilla hartii is morphologically similar to V. planifolia, and even confused with it in many treatments. However, all the molecular data place it in a rather basal position among the American penicillate vanillas. Vanilla hartii tolerates damper, more shady conditions, but the fragrant fruits are much smaller, O THER RECORDS : MEXICO: CHIAPAS: Estacin de Biologa de Chajul, selva alta perennifolia, selva de loma con Calophyllum y sotobosque de cyperceas altas y helechos arborescentes, sobre el camino a Arroyo Miranda, 16’’N, 90’’W; 200 m s.n.m., 12-IV-1997, M. Soto 8347 *AMO(sterile)! 8350 *AMO(fruit)! same data 14 abril 2000, M. Soto *9727(x2), *9729(x2), S. Maldonado, L. Lpez y P. Schltter AMO!; same data: el engrosamiento apical, con lneas inconspicuas, cafs, tenues en la garganta, sin fragancia aparente. Rondada por Euglossa hembra, pero no capturada ni vista polinizando la dulce M. Soto 9730, S. Maldonado, L. Lpez y P. Schltter *AMO! Sobre la Vereda La Granja, Estacin de Biologa de Chajul, selva alta perennifolia en zona de loma con muchos arroyos, algo perturbada, ca. 16’N, 90’W, M. Soto 9731 y P. Schltter *AMO! BELIZE: TOLEDO: between Orange Point and Moho River, April 28 1952, P.H. Gentle 7673 MEXU(511492)! Southern Maya Mountains, Bladen Nature Reserve, mountain, 1.7 airline north of Ex Xux archeological site, 16’’N, 881’’W, 1996, G. Davidse 36251 BM! : [cf.] “vanilla”, vine, P.H. Gentle 328 F(713628)! GUATEMALA: : Leaves subcoriaceous, dark dull green above, slightly paler dull green below. Stem terete, dull green. Petals and sepals pale greenish-white. Lip white. Leaves somewhat narrower than in typical V. fragrans. Swamps of Salomn Creek, 1/2-1 mi. south of Bananera, alt. 50 m. April 6, 1940. J.A. Steyermark 38944 *F(1043051)! [cf.] Quebradas, 19-22, May 1919, H. Pittier 8589A NY(sterile)! *US(1013493; sterile)! HONDURAS : ATLANTIDA: [cf.], sterile Lancetilla Valley, near Tela, altitude 20 to 600 m; “vainilla”, creeping on tree in wet forest; frequent, Dec. 6, 1927-Mar. 20, 1928, P.C. Standley 52824 *AMES(36946, fruit)! F(582560)! *US(1407340; sterile)! NICARAGUA : : Monkey Point; ca. 11’N, 83’W, elev 0-20 m; beach and bluff near village; vine on understory tree, W.D. Stevens, B.A. Krukoff 20021 Sung, N of road between Puerto Cabezas and Ro Wawa; approximately 14-11’N, 83-31’W, elev. less than 10 m; gallery forest and adjacent savanna. Vine on tree trunk, sterile. SEL(036668)!

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla309 elev. ca. 10 plain. Epiphytic vine, sterile. W.D. Stevens, B.A. Krukoff 8835 leaf, comments and analytical drawing on envelope perhaps belong to V. planifolia ] A.H. Heller s.n. *SEL(03849)! COSTA RICA : PUNTARENAS: Hilly slopes west of Villa Nueva and Ro Naranjo. Evergreen rainforest formations on 84’ W. Vine growing in partial shade 1 m up on stump. W. Burger, K. F(2009056)! Cantn de Osa, Aguabuena. Cuenca superior de Quebrada Aguabuena. 8”N 83”W, 200-400 m. Bejuco trepador verde, labelo blanco con mancha puntual, apical, verde, columna blanca, polinios amarillo blanco. 18 January 1991, G. Herrera 4846 INB! MO SAN JOSE: Ro Savegre, aproximadamente 1 km antes de la entrada a la Finca La Gavilana, cerca de los lmites del Parque Nacional Tapant, selva lluviosa con Anacardium, ca. 200 n.s.n.m., trepadora, con cpsulas, relativamente muy robusta, M. Soto INB! HEREDIA: Sarapiqu, Chilamate. Finca El Bejuco, S end of Cerros Sardinal (N of Ro Sarapiqu). 10’N, 84’W, 60 m. Vine climbing in understory tree at edge of forest to 3-4 m (transplanted here from swampy area of primary forest nearby). Sepals and wing petals cream-white. Labellum pure white, gibbous at very base, constricted 1/4-1/3 towards apex, gibbous again (more prominently) in apical 3/4, slightly narrowed before rotate margin. Labellum with grayish lines within. 24 January 1987. *MO(3593651)! PANAMA: PANAMA:cf.] Ro Tecumen. Moist forest; herbaceous vine; scarce, January 3, 1924, P.C. Standley 29353 *AMES(31441; sterile)! 6. Vanilla helleri A.D.Hawkes, Phytologia 14(1): 34, 19-20. t. 13.1966. TYPE: NICARAGUA: Dept. Chontales: 2 miles south of La Libertad on the road to Sto. Toms, growing as a vine on a wild avocado (Persea sp.) tree, alt. 1900 feet, April 1962, A.H. Heller 7946, holo. AMES? (not located), iso., SEL(fragments and drawing)! Hemiepiphytic vine, branching, leafy. Stems olive green, minutely and inconspicuously papilloserugose in living condition; 5-10 mm thick; internodes up to 15 cm long, the apical stems conspicuously covered 1-3 mm wide. Leaves subpetiolate, the petiole 9-12 mm long, the blade oblong-elliptic to oblanceolate towards the acuminate apex (acumen 10-15 mm), 1215 x 2.5-4.5(-6) cm. raceme, candelabrum-shaped, 4-10 cm long; peduncle 1-2 cm long. Bracts triangular-ovate, acute to obtuse, dark green, up to 9 x 8 mm. Flowers successive, 1 open at once, with partially spreading segments, ephemeral, showy, medium-size, sepals whitish, greenish at the apical half on the abaxial surface, petals whitish, lip bright yellow with orange appendages on the midlobe, column white; ca. 6 cm diameter. slender, subterete, white, apex green, 39-50 mm long, 3 mm thick. Dorsal sepal oblanceolate-elliptic, subcalyptrate at apex, 11-veined, 40-42 x 11-14 mm. Lateral sepals obliquely elliptic, obtuse, slightly x 13-14 mm. Petals oblong-elliptic, oblique, subacute to obtuse, rounded, 11-13-veined, dorsally keeled, 4047 x 11-13 mm. Lip attached to the column margins cuneate, without a claw, the blade clearly trilobed, when spread out 35-40 x 30-33 mm; lateral lobes repand with branched veins, ca. 5 mm wide at apex; midlobe subquadrate to transversely oblong, truncate, covered with retrorse, complanate, appendages up to 3 mm long, along the veins; disc with inconspicuous axial rows of minute warts, with abundant trichomes on the sides of the basal half; penicillate callus at 20-23 mm from the base, almost continuous with the axial warts and the apical appendages, a tuft of long laciniate, scales, much divided, ca. 5 x 5 mm. Column subclavate, rather short and thick, semicylindric, 28 x 4 mm, apex dilated ca. 5 mm wide; ventral surface ca. 1 mm long, 2 mm wide. Stigma trilobed, with a mm broad; lateral lobes emergent, quadrate, tongueshaped, ca. 1 x 1 mm. Anther versatile, transversely ellipsoid, ca. 3 x 3 mm. Fig. 7. DISTRIBUTION : Costa Rica and Nicaragua, and also apparently in Oaxaca, Mexico. Our knowledge of the distribution of V. helleri is very fragmentary. It is known from the type locality on the slopes of the

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.310 LANKESTERIANA FIGURE 7. Vanilla helleri Hawkes. Based on . Drawing by M. Lpez and M. Soto.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla311 Costa Rica, especially in the Osa Peninsula. ECOLOGY : In rainforest, with a short, marked dry April. This species has a lip with elongate, complanate appendages, similar to that found in V. insignis. The outline of the lip, with a cuneate base lacking a claw is very anomalous among American penicillate vanillas and is reminiscent of some Old World species. In the original description Hawkes stressed that this species has no close allies in Central America. However, morphological and molecular data suggest that it is closely related to V. insignis and V. odorata, but the have conspicuously long-clawed lip. Vanilla helleri, described 30 years ago is one of the rarest vanilla in herbarium specimens. Despite its rarity in the herbaria, V. helleri could be more common than usually thought. Several specimens of it are growing at Finca La Gavilana, Ro Savegre, Costa Rica. Additionally, have been located around Sta. Mara Chimalapa, in Oaxaca, Mexico. Most Mexican specimens have rather elliptic, broader leaves than Costa Rican material, but they share the sulcate stems, papillose stem surface, abundant wax on stem apex, and the nucleotidic sequences are very similar to those of Costa Rica. Originally, the Mexican specimens were thought to be a hybrid between V. planifolia and V. insignis, since V. helleri has vegetative traits intermediate between these O THER RECORDS : COSTA RICA : PUNTARENAS: 2 km SW of the intersection of the Ro Ceibo and Carretera Interamericana; elev. 200 m. Large vine climbing in a tree. 18 March 1969, F(1731785)! MEXICO: OAXACA : [cf.] Planta jven, silvestre, en selva alta perennifolia con Sebastiania, Quercus oleides, Terminalia, Acacia , Ampelocera, ca. km 42 del camino MezquiteSta. Mara Chimalapa, 390 m s.n.m., 20 Marzo 1997, (not preserved). [cf.] Tallos speros al tacto y subpapilosos, longitudinalmente sulcados; al sureste de Sta. Mara Chimalapa, vereda hacia el Ro Milagro, paso Quetz Tug, 15/XI/98 a 6/XII/98, H. Hernndez V7 sub M. Soto 8806 AMO(sterile, photo)! [cf.] Hojas subcuneadas en la base, algo oblanceoladas y algo oblicuas; 3 km al este de Sta. Mara Chimalapa, cerca de la cascada Arroyo Sangre. 15/XI/98 a 6/XII/98 H. Hernndez V3 sub M. Soto 8811 AMO(sterile, photo)! [cf.] Tallos muy delgados, sulcados, speros, algo glaucos, hojas elpticas, de Sta. Mara Chimalapa, a la orilla del camino Sta. MaraH. Hernndez V8 sub M. Soto 8817 AMO(sterile, photo)! Tallos muy delgados, sulcados, speros, hojas elpticas, Sta Mara Chimalapa, a 5 m de la vereda de Paso Lagarto. H. Hernndez V2 sub M. Soto 8818 AMO (sterile, photo, ITS sequence)! REFERENCES : Hamer, Orch. Nicaragua, Ic. Pl. Trop. pl. 1192. 1984; Hamer, Selbyana 11. Orch. Centr. A. p. 847. 1990. 7. Vanilla inodora Schiede, Linnaea 4(4): 574-575. 1829. TYPE (?): “Baynilla de puerco Misantlensium”; Mexico, Misantla. Schiede 1044, K; “fruto inodori, Vanilla inodora nob. interin Vaynilla de puerco, Misantlensis .... Misantla, Mart 29 BM! W(s.n.; x2)! Vanilla pfaviana Rchb.f., Gard. Chron. 2, 20. 230. 1883. Holotype: “Mexico” actually from Chiriqu, Costa Rica, W(19347)! Vanilla preussii Krnzl., Notizbl. Bot. Gart. BerlinDahlem. 7: 320. 1919. Type: Bei der Planzung El Bal im Buschwald, Heisst bei den Eingeborenen “Vainilla silvestre” (Preuss n. 1445), not seen. C OMMON NAMES : “Vainilla de puerco” (Ver.); “vainilla”, “vainilla cerro amarillo” (Oaxaca). Hemiepiphytic herb, vigorous,up to 12 m high. Stems terete, ca. 4-5 mm thick; internodes ca. 7.5 long. Leaves usually hanging and with bending blades; petiolate, the petiole ca. 1-2 cm long, twisted; blades elliptic to broadly elliptic, abruptely acuminate, thin, membranaceous when dry; 12-21 x 4.5-7 cm. similar to the vegetative shoots; ca. progressively smaller foliaceous bracts. Bracts similar to the leaves, although smaller or much smaller, 3.3-12 x 1.0-5.2 cm; the internodes 3.5-7 cm long, ca. 2 mm thick. Flowers resupinate, with spreading segments, very showy, tepals apple-green or

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.312 LANKESTERIANA FIGURE 8. Vanilla inodora Schiede, based on M. Hernandez s.n. sub M. Soto 7864. Drawing by R. Jimnez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla313yellowish green, shiny, lip white with yellow-orange tinge in the throat; ca. 4.5-5 cm diameter; fragrance sweet strong or faint medicinal, to weak solanaceous straight to slightly arcuate, terete, smooth, 3-sulcate, the grooves twisted, 47-55 (48.67.25) mm long, 3-4 (3.33.47) mm thick. Tepals contorted, with very undulate, somewhat revolute margins, and the apices recurved or rolled backwards. Dorsal sepal erect, ellipticlanceolate, broadly obtuse to acute, ca. 11-veined, 44-50 (47.19) x 12-18 (13.4.8) mm. Lateral sepals descending, elliptic-lanceolate, broadly obtuse to acute, ca. 11-veined, 41-44 (42.4.02) x 13-17 (15_1.79) mm. Petals spreading, elliptic-lanceolate, attenuate towards the obtuse apex, axially canaliculate on the abaxial surface, ca. 9-veined, 43-47 (45.67) x 9-11 (10.26) mm. Lip attached to the column ca. 2.5 mm at base; arcuate, trilobed, the lateral lobes covering (33.16) mm long, 29-34 (31.6.62) mm wide when spread out; lateral lobes erect, semiobcordate, broadly rounded, reduplicated and covering the distal half of the column; 20-24.5 (22.7.23) x 9-13.5 (12.2.69) mm; midlobe subcuadrate, spreading, deeply emarginate, the margins undulate, 12-17 (14.81.94) x 13-19.5 (15.8.38) mm; the midlobe and part of callus, 16-21 (18.79) mm long, 5-8 (6.8.16) mm broad, ca. 3-5 mm high. Column strongly arcuate, the apical part resting on the callus surface, 21-24 (22.6.16) mm long, 5-6.3 (5.76.56) mm wide at the apex. Stigma panduriform cavity with thickened, yellow borders; without a rostellum. Anther hinge-like, broad, attached to the clinandrium, and forming together with it, a couple of lateral auricles. Pollen soft, somewhat sticky. Fruit cylindric with attenuate base and apex, not aromatic, 18 cm long, ca. 8 mm thick. Fig. 8. DISTRIBUTION : Mexico (Puebla, Veracruz, Jalisco, Guerrero, Oaxaca, Chiapas), Belize, Guatemala, Panama. ECOLOGY : Vanilla inodora grows in wet forests from 150 to 1600 m altitude; it is the only Vanilla that inhabits the cloud forests of the area, and is found in the lowlands only in sites with more than 2500 mm period. It is sometimes found in savanas with slow drainage. Vanilla growers believe that this species can be self-pollinated; the large fruit set in some populations supports this idea; however, other populations have a condition for 2-3 days, and are therefore long-lived compared with other vanillas; often older and younger species of Schomburgkia group of Laelia structure; the tepals are contorted, strongly twisted, with a varnished appearance and the column lies on the lip surface; pollination must be carried out by large, strong bees that try to enter to the throat separating the lip from the column, as in Schomburgkia or Barkeria; in the latter this work is done by carpenter bees (Xylocopa), and they might be good candidates to pollinate this species. We have observed carpenter bees approaching V. inodora in Chiapas, but they were V. inodora was collected by Schiede near Misantla; the original collections are housed at K, BM and although probably some specimens have had fruits (see Rolfe 1896); so its identity has been somewhat obscure. The type specimen is evidently a member of Portres subsect. Membranacea, and as only a single species of this group is found in Mexico, and it is common in Veracruz, we discard the possibility of other species, from elsewhere, to which the name V. inodora has been applied. There is no evidence that the Vanilla mexicana, as suggested by many authors, despite the of V. inodora are larger than in other Membranaceous species and are clearly visible in the type. Vanilla inodora has been known in recent years in the region as V. pfaviana Rchb.f. In their Orchids of Guatemala, Ames and Correll (1952) mentioned that they had not examined Guatemalan material of V. inodora with V. pfaviana. It seems, however, that the sterile specimens (or those very badly preserved) of this V. mexicana or V. inodora to be V. pfaviana. Vanilla inodora and V. mexicana

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.314 LANKESTERIANA are V. inodora is very different from the 3-keeled callus found in V. mexicana; furthermore, V. mexicana can have much larger leaves. It should be noticed that the callus of V. inodora may appear to be keeled in old, The type of V. preussii Kraenzl. was destroyed in the Berlin bombing, but the description matches V. inodora very well. Furthermore, Schlechter, who must Trckheim II 1764 as V. preussii. The allied species in Central America are V. martinezii, from Guatemala, that is easily distinguished V. costaricensis with entire lip lacking a massive callus, and V. sarapiquensis, with short midlobe and a callus formed of the lip. However, its closest relative may be the Andean V. methonica emarginate lip, but having 3 thickened and somewhat OTHER RECORDS : MEXICO: JALISCO: Steep mountainsides 3-10 km generally east on the road to Mina Cuale, from the junction 5 km northwest of El Tuito, Mpio. Cabo Corrientes; pine-oak forest on decomposed granitic soils, with Podocarpus, oaks, and other deciduous trees in rocky stream valleys, elev. 850-1150 m. Seen once, in rocky or more in trees, 16-19 February 1975, R. McVaugh 26397 MEXU! Mpio. Cabo Corrientes, km 2.6 de la brecha hacia a la izquierda que sale del km 9.8 del camino El TuitoCuale, bosque de galera de Hedyosmum, Inga, Podocarpus, Magnolia, Clusia, en medio del bosque sabanoide de pinos y encinos, sobre granitos intemperizados, 900 m s.n.m., 20.4’’ y 105.9’’. Banco con suelo profundo con vegetacin densa, similar a acahual de selva montana lluviosa con Heliconias , Philodendrum tripartitum, Lindenia muchos frutos, cerca 400 m de extensin. Muy rara, slo un espcimen visto. 27/junio/1998 M. Soto 8626 y E. Huerta *AMO(x6)! IBUG! GUERRERO: System of Teotepec, near San Vicente, N.E. of Atoyac, in mixed oak-pine forest on tree trunks. 100’W, 17’N, 850 m, January 10, 1933. O. Nagel sub E. Oestlund 1984 AMES (with sketch by B. Ames; 41396)! *AMES(51815)! *MO(11411617; sterile)!; San Vicente, north-east of Atoyac, towards Mt. 17’N, ca. 100-1000 m, 10 Jan 1933. O. Nagel sub E. Oestlund 1985 *AMES(41476, sterile)! BM(sterile)! *MO(1145095; sterile)! PUEBLA: Vicinity of Puebla, Dc. 8. 1907. B.G. Arsne 1682 *AMES(71359, sterile)! *MO(843101, sterile)! NY! *US(1032010; fruit)! : Miradores, 4/42 Liebmann 297 K! Mirador, Liebmann W(11762)! Hacienda de Java, 5/1841, Liebmann 295/296 C.A. Purpus 8481 *AMES(71356)! NY! Region of 700 m, 10 Jul 1935, C.A. Purpus sub E. Oestlund 4876 On trees and shrubs, 12 Feb 1932, 96’ W, 19’ N, ca. 900 m, O. Nagel sub E. Oestlund 2683 *AMES(sterile, trunks, 8 Oct 1935, ca. 800 m, C.A. Purpus sub Oestlund 5045 *AMES(fruit, 51838)! Falda del Volcn de San Martn Tuxtla, San Andrs Tuxtla, 18’, 95’W, alt. 800 m s.n.m., selva alta perennifolia primaria, suelo negro arenoso, blanca, 29-10-1973, J.I. Calzada 01031 BM! CHAPA! MEXU! El Mirador, Mpio. Totutla, bosque de encino en blanco, fruto verde, escasa, 18-VI-1973, F. Ventura 8417 *AMO(436)! CHAPA! MEXU! Lote 67, Estacin de Biologa Tropical Los Tuxtlas, 95’ y 95’ O, 18’ y 18’ N. Mpio. San Andrs Tuxtla, borde selva alta perennifolia, 300 m s.n.m, bejuco herbceo, fruto verdegrisceo, “vainilla”, julio 14 de 1986, S. Sinaca C. 835 *AMO(7066)! OAXACA: Cerro Martn, cerca de Usila, 400 m selva alta perennifolia, predio de Sebastin Arista, 22 mayo 1993, M. Hernndez s.n. AMO(in spirit)! Usila, M. Hernndez s.n. AMO(in spirit)! Usila, 2 mayo 1994, M. Hernndez s.n. AMO(in spirit)! Cerro Martn, ca. Usila, 1992, M. Hernndez s.n. AMO(in spirit)! La Escalera, Chinantla. “Vainilla Cerro Amarillo”. abril-1990. M. Hernndez Apolinar sub M. Soto 10707 AMO! 1844, Mexico, Karwinski M W(19347, sketch)! Mpio. Sta. Ma. Chimalapa: Arroyo Sangre ca. 2 km E de Sta. Ma. Chimalapa, selva perturbada con Calophyllum , Tapirira, Brosimum, etc., suelo caf parduzco con mucha hojarasca, en aceite para el cabello, H. Hernndez 1125 CHAPA! selva alta perennifolia, loc.: Cuaje, Mpio. Ixtln de Jurez, Dto. Ixtln, Sierra Norte, 17/05/94, E. Torres 367 AMO(16696; young fruit)! TABASCO pukt en vegetacin riparia, Teapa, 31-05-1990, V. Ramn MEXU [sterile]! CHIAPAS: “Local name: “vainilla”, in wet forest, Libertad Acacoyagua, June 1 1948, E. Matuda 17912 AMES(66534; young fruit)! F(1616949)!

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla315 MEXU(85052)! En Crucero Corozal, camino PalenqueBoca Lacantm, Mpio. Ocosingo, 180 m s.n.m., bejuco con fruto, selva alta subperennifolia, 8 enero 1986, E. Martnez 15730 *LL! MEXU(436979)! MO(4272282)! *XAL! Triunfo, Dec. 1936, E. Matuda 360 *US(1689405)! TriunfoJurez, Escuintla, Dic. 1936, E. Matuda 369 MEXU(85064)! Arroyo Miranda, entre los 6 y 20 km del arroyo partiendo del Ro Chajul, Mpio. Ocosingo, 150 m s.n.m., primaria, “vainilla”, trepador, 20-02-1985, G. Castillo et al. 4011 *XAL! 2 km al NW de Lacanj-Chansayab; Mpio. de Ocosingo, 400 m, enredaderas; vainas verdes, lisas, negras cuando maduras; numerosas semillas negras cubiertas con una secrecin pegajosa de las paredes internas de las vainas; abundante en remanente de selva alta perennifoia relativamente madura, abril 16, 1991, M. Gonzlez Espinosa et al. 1413 CHIP! MEXU(563979)! Bonampak, Mun. Ocosingo, 350 m s.n.m. selva alta perennifolia, suelo hasta de 7 m de largo. abril 1982. M. Soto 1020 AMO! Estacin de Biologa Chajul, Mpio. Ocosingo, ca. 200 m s.n.m., 16’ N, 90’ O. Cerca del Puente Hamaca en la vereda a la Sabana I. Loma con selva mediana-alta perennifolia, en la base de la loma. Muy escasa. 23 junio 1996. M. Soto 7954-A y R. Solano *AMO! km 223.6 de la carretera Palenque-Marquz de Comillas, ca. de Benemrito de las Amricas, selva mediana subperennifolia con Cocoloba y Roupala, 220 m s.n.m., 11-IV-1997, M. Soto 8342 et al. AMO(also in spirit)! Mpio de Ocosingo, ca. de Chajul, selva alta perennifolia inundable con Bactris, 230 m, 16’ N, 90’ O. Sobre Guarea en sombra densa, con frutos. 12-IV-1997. M. Soto 8343 *AMO! Estacin de Biologa de Chajul, junto al Ro Lacantun, vereda La Granja, en la interseccin con la vereda a La Sabana, selva alta perennifolia con Ficus glabrata en suelos profundos, verde con labelo blanco, fragancia dulce, intensa, 15 de abril de 2000, AMO! BELIZE: TOLEDO. Gracie Rock, Sibun River, 1 May 1935, P.H. Gentle 1672 *AMES(42338)! *LL! “Vainilla”. ridge, near river beyond Columbia. January 31, 1947, P.H. Gentle 6152 *F(1599314)! G! *LL(x3)! NY! *US(2572753)! Ocotal, Pine Ridge, 3 miles in trail from 7 Miles, Punta Gorda-San Antonio Road, January 20, 1950, P. Gentle 6957 *LL(x3)! MEXU(511462)! *MO(3832518)! “Vianilla”, vine, in acahual, Feeders Road leading to Big Fall, April 13, 1950, P.H. Gentle 7023 *LL(fruit, x2)! “Vianilla”, vine, in cohune ridge, one mile from 7 Miles, San Antonio-Punta Gorda Road, July 1, 1950, P.H. Gentle 7078 *LL(sterile x 2)! “Vianilla”, vine, in cohune ridge, near Columbia, August 15, 1950, P.H. Gentle 7108 *LL(fruit)! GUATEMALA: : [cf., sterile] wet forest,. “Vainilla”. Creeping on tree trunk. Near Entre Ros, alt. about 18 m., April 30, 1939, P.C. Standley 72709 F(991636)! ESCUINTLA Ficus, April 9, 1937, W.C. Muescher 12480 F(905455)! SAN MARCOS coriaceous, rich green above, practically same color but slightly paler beneath, above Finca El Porvenir on “Todos Santos Chiquitos”, lower south facing slopes of Volcn Tajumulco, alt. 1300-1500 m, March 7, 1940, J.A. Steyermark 37076 F( 1041850)! HUEHUETENANGO : [cf.] Epiphyte, alt., At 3000 ft. alt. Cerro Chiblac, between San Rafael and Ixcn, Sierra de los Cuchumatanes, July 22, 1942, J.A. Steyermark 49171 *AMES(63277, sterile)! F(1495682)! : [cf., sterile] Epiphyte on above, dull green beneath, stem pale green, in cafetal on opposite side of Finca, southern lower slopes of Volcn east of Puebo Nuevo, alt. 500-800 m, Feb 2, 1940, J.A. Steyermark 35412 F(1041244)! : Wald in Paujal, 1000 met April 1907, Bl. grn, Lippe weiss, H. von Trckheim II 1764 *US(825825)! HONDURAS : COLON: 100 ft., tropical rain forest, March 1938, von Hagen 1352 F(942976)! NY! ATLANTIDA: [cf.] banks of the Salado River, above the village of Salado ... on the mountain slopes and coastal plains, vicinity of La Ceiba, July 10, 1938, T.G. *AMES(fruit, 50661)! NY! COSTA RICA : ALAJUELA: Reserva Biolgica Monteverde. Ro Aguas Gatas, Laguna de Arenal, El dentro del bosque. Flor con caliz verde, corola blanca en forma de tubo. 11 August 1989, E. Bello 1146 INB! [cf., entre el Ro San Lorenzo y la estacin. 10’’N, G. Herrera Ch., I. Chacn, D. Hernndez, A. Sols y H. Gmez 386 SEL(062313)! CARTAGO: vainilla, “Chitera” = Chitara, alt. 750 m, 15/4/36, F. Sols F(833816)! [cf., sterile] LIMON: Epiphyte, 7 km al SW de Bribrs, 100-250 m, May 4 1983 [sterile], L.D. Gmez, R. Liesner, E. Judziewicz 20437 MEXU! MO [cf., sterile] San Clemente, Apr. 1920. Lankester (k331) K! SAN JOSE: Herbaceous vine, attached by roots. Fls green with white labellum, in forest. Vicinity of El General, alt. 1130 m, Feb. 1936, A. Skutch 2592 AMES(*44231, *44232, *44233)! K! *MO(1105371)! NY![ Cataratas de San Ramn, marzo de 1931, A.M. Brenes 13679 F(906350)! PUNTARENAS: la casa. Cerro Anguciana. Fila Cruces, 08N, 831 W, 900 m. Bejuco trepador. Caliz verde, labelo

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.316 LANKESTERIANA blanco. 10 December 1993. R. Aguilar 2736 INB! PANAMA: BOCAS DEL TORO: [cf.] Vanilla, epiphyte; Oct., 27, 1940, H. von Wedel 2880 *AMES(61513, sterile)! REFERENCES 4(3): 26-29. . 1936.8. Vanilla insignis 102. . 1934. TYPE: REPUBLIC OF HONDURAS, Dpt. Comayagua, Esquas, El Ro Funes. Epiphyte in river-valley forest at 2,500 feet altitude.Sepals and petals green, lip white at base. April 27, 1933.J.B. Edwards 407, holo. AMES(40085)!; isotype AMES(40086)! COMMON NAMES : “Sisbic” (Maya), “vainilla”, “vainilla cimarrona” (Veracruz). Hemiepiphytic, branching, leafy vine, usually very vigorous, up to 30 m high; up to a hundred meters long. Stems groove along the entire internode, surface fairly rugose-papillose, dark to olive green, ca. 6-9 mm thick; internodes, 10-17 cm long. Terrestrial roots pubescent, brown-whitish, ca. 2-3 mm thick; aerial free roots, terete, greenish gray, ca. 1 mm thick; aerial in contact with the substrate, ca. 3-4 mm wide. Leaves subpetiolate, petiole canaliculate, up to 11 mm long, 4 mm wide;blade oblong-elliptic, abruptly acuminate rather stiff, 4.2-19.5 x 1.5-3.4 cm. a compressed, 15-25 mm long, 6.5-8 mm thick; rachis 28-90 mm long, progressively slender towards the apex. Bracts sessile, small, broadly ovate, obtuse, apex, commonly less than 9 x 6 mm, rarely up to 10 x 10 mm. Flowers successive, 1-3 open at once, ephemeral (from 8:00 to 14:00 hrs; starting to close at noon), very showy, segments completely spreading, tepals pale apple-green, shiny, lip cream-white with orange to dull yellow appendages on the midlobe, extreme apex green, penicillate callus cream, column white; ca. 11-12.4 cm wide, 9-11 cm high; fragrance weak, spicy. subterete, very slightly dorsiventrally compressed, smooth, arcuate, 2-sulcate, the grooves almost straight, 45-54 (48.753.27) mm long, 4.5-5.5 (50.41) mm thick. Dorsal sepal long oblanceolate, apex acute, disatally rounded, slightly thickened, subcalyptrate, base attenuate-subunguiculate, claw ca. 22 x 11 mm, basally canaliculate, concave at middle, fairly incurved at apex, ca. 12-13-veined; 69-76.5 (73.16 _ 2.56) x 10.5-14 mm (11.61.24). Lateral sepals obliquely oblanceolate, the lower margin more arcuate, apex subacute, thickened, apiculate, subcalyptrate, smooth, the abaxial surface minutely warty, especially at the minute apicule, base attenuatesubunguiculate, basally canaliculate; slightly concave towards the apex, margins somewhat involute, ca. 11-14 veined, 64-74 (69.673.19) x 12.5-13.5 (12.92 0.35) mm. Petals obliquely linear to oblanceolate, somewhat arcuate, apex obtuse, distally rounded, oblique, subcalyptrate, base long attenuate; concave, the abaxial surface, ending in a subtrigonous, free, adpressed process, ca. 2 mm long; ca. 12-14-veined, 68-74 (70.92.11) x 9-10 (9.760.39) mm. Lip fused to the column along the margins of the basal half (37-44 mm), long tubular, slightly concave, the abaxial surface, the groove deep; when spread out 65-73 (69.332.49) x 30-35 (32.61.62) mm; long unguiculate, adaxial surface of claw hairy on the distal half, the trichomes dense, more or less in rows; ca. 26 x in outline, trilobed, ca. 30-veined, the veins branched keels near the base, disappearing near the basal third of the blade; lateral lobes subelliptic, oblique, ca. 33 x near the joint with the midlobe; cilia up to 7 mm long; midlobe ovate-suborbicular to oblong, obtuse, the margin undulate-crenulate to lacerate-dentate towards the lateral lobes, 11-15 (131.41) x 12.5-14 (13.37 0.65) mm; penicillate callus at ca. 34 mm from the base, 4 x 5 mm; made up by ca. 13 imbricated, laciniate scales, sometimes united to each other along the lateral margins, scales almost without adpressed processes on the surface, continuous towards the lip apex with 7 rows of tubercles, near of the base of the

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla317 FIGURE 9. Vanilla insignis Ames. Based on M. Soto 7684. Drawing by M. Soto.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.318 LANKESTERIANAmidlobe becoming enlarged to form conspicuous, retrorse, subtriangular, thick, complanate, sometimes conical, obtuse appendages, that cover almost entirely the surface of the midlobe, distributed approximately in 10 rows, the central appendages bigger, up to 4 mm long, 1 mm wide; extreme apex thickened, mound-shaped. Column very elongate and slender, semicylindric-trigonous, 49-52 (50.37.08) mm minute trichomes below stigma; apex dilated, with x 2.3 mm; clinandrium galeate, ovate, convex, 2 x 2 mm. Stigma trilobed, the lobes emergent, midlobe (rostellum) a transversely oblong, convex blade, axially grooved, ca. 3 x 4 mm, covering and parallel to the lateral lobes, these quadrate, rounded, convex, slightly divergent from each other, ca. 1.5 x 1.5 mm. Anther versatile, saddle-shaped, axially canaliculate, 3 mm wide, 1.8 mm thick, attached to the clinandrium Pollen in monads, not forming a clear pollinarium Fruit short, thick, subclaviform-semifusiform, rounded, swollen towards the apex, green turning yellow when ripe, dehiscent along a single line, fragrant (less than V. planifolia), like a blend of common vanilla with coconut; ca. 7-14 cm long, 0.7-1.75 cm thick (n = 6). Fig. 9, 17C. DISTRIBUTION : The Caribbean watershed of N Central America, in Honduras, Belize, Guatemala, and Mexico (Yucatn, Quintana Roo, Campeche, Chiapas, Tabasco, Oaxaca, and Veracruz; perhaps also in Puebla). Reported from Panama (Dressler, 1993) but the report seems to be based on specimens of V. dressleri. ECOLOGY : From the level to ca. 900 m elevation. Vanilla insignis is probably the most common vanilla in Mexico (Soto Arenas, 2003), where it is widely distributed and forms large populations. It grows in dry and wet areas (1000 to 4000 mm of annual especial edaphic conditions. It has been recorded only from calcareous substrates. In the Yucatan Peninsula it grows in the subdeciduous forests of Bucida buceras, Brosimum alicastrum, and Manilkara sapota, often with the understory dominated by the palm Cryosophila argentea; these areas have slow drainage during the rainy season, and are frequently associated Haematoxylon campechianum.In the much moister areas of Chiapas, it is found in savannas with Coccoloba belizensis, Quercus oleoides, and Roupala borealis.In Central Veracruz V. insignis grows in tropical deciduous forest or in warm oak forest. Some specimens in Campeche and Chiapas seem to occupy areas up to 4,000 m2, and undoubtedly they are the largest plants of any Vanilla of the area. Some of these specimens may prove to be the most massive orchidaceous specimens in the world, with weights of bees of Eulaema polychroma. This is another member of the V. planifolia complex and it h V. planifolia by some authors (e.g. Williams, 1956). However, it is are rather similar to those of V. planifolia in pressed V. planifolia (1-2-dimethil-ciclopentane, ethyl acetate, and 1-8-cineol as principal constituents, although ocimene-trans is notoriously absent. It is easily recognized because the stems have internodes conspicuously sulcate and their surface is fairly rugose-papillose. Similar vegetative traits are found in some very distantly related Asian species, namely V. yersiniana V. moonii Thwaites , and it allies. The sulcate stems are thought to be an adaptation that permits the stem to store more water during the rainy period in seasonal areas, since the groove expands and becomes inconspicuous when the stem tissues are swollen. The characteristic tepals, cream-white lip, the midlobe adorned with thick, triangular, retrorse projections, up to 4 mm high. The fruit (ca. 10-12 cm long) is thick and fragrant when ripe, but the aroma is similar to a mixture of vanilla with coconut. The ITS tree (Fig. 1) and a survey of additional genomic regions, including also non-Mesoamerican Vanilla species show that the closest relatives of V. insignis are V. odorata Presl, V. helleri A.D.Hawkes, V. uncinata Huber ex Hoehne, and V. tahitensis J.W.Moore. Vanilla odorata, V. uncinata, and V. tahitensis have much narrower leaves, non-sulcate d less developed retrorse appendages on the lip. From V. helleri, from Oaxaca,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla319 Nicaragua and Costa Rica, with which it shares the papillose stem surface, sulcate internodes, and long, retrorse, orange-coloured papillae on the distal part of the lip, it can be easily distinguished by the long claw of the lip found in V. insignis vs. the cuneate lip blade of V. helleri. Vanilla insignis was described in 1934, and it is surprising that it has not been reported until recently from Mexico (Carnevali et al., 2001; Soto Arenas, 2003) although it was collected in Veracruz by C.A. Purpus in 1919. Neither has it been reported so far from Guatemala or Belize, although a picture of V. insignis V. pfaviana in “Native Orchids of Belize” (McLeish et al., 1995). However, this species has been known and used by the Mayas for a long time, who call it “sisbic”. Almost all the specimens V. planifolia or V. fragrans from the Yucatan Peninsula and Veracruz belong to V. insignis . The report of V. odorata from Quintana Roo, based on Cabrera 4611 (Soto Arenas, 1989) is based in a specimen with buds that is actually V. insignis . Vanilla insignis, could add desirable features to the commercial vanillas; it also has fragrant fruits, it is a more xerophytic, stouter species, and apparently We have cultivated this species and it is also the Mesoamerican species most tolerant to low and high temperatures, and much more resistant to the attack of pathogens than V. planifolia.O THER RECORDS : MEXICO: June 1919, C.A. Purpus 8482 *AMES(71362)! NY! Near 1932, 96’ W, 19’ N, ca. 900 m, O. Nagel sub E. Oestlund 2682 AMES(*41478, *51844, sterile)! Region 800 m, C.A. Purpus sub E. Oestlund 5961 *AMES(51845)! 800 m, C.A. Purpus 4866 seen, *51848, sterile)! Carretera Xalapa-Veracruz, km 16 SE of Xalapa, 1 km SE of main Jalapa-Huatusco highway, 5 km SW of bridge over Ro Los Pescados, 5 km (by air) SE of Tuzamapan, Mpio. Coatepec, 19’N, 96 50”W, 680 m alt., “selva baja caducifolia”, thorn scrub along now dry canyon, now very dry and most trees without leaves. Vine to 3 m, fruits green, hanging, March 19, 1983, M. Nee F(1985066)! *XAL! Mpio. Emiliano m s.n.m., selva baja, primaria, suelo arcilloso, pedregoso, color negro, muy seco, clido, bejuco perenne, 6 m, escaso, fruto verde; trepadora, 15-04-1979, 531 desviacin de la carretera Xalapa-Veracruz, 16 km al SE y amarillo, escasa, “vainilla”, 23/V/1976, C.H. Ramos 402 MEXU! [cf., fruits] Mpio. Soteapan, San Fernando, 18’, 600 m, acahual, selva alta perennifolia, 2-3 m, “vainilla”, 19-IX-86, (usos) aromatizante de aceite, M.C. Gonzlez R. 303 *XAL! OAXACA : [cf., sterile] Forests ca. 25 km east long aerial roots in leafmould. Isthmus of Tehuantepec, ca. 94’ W, 17’ N, alt. ca. 100 m, 20 feb 1935, O. Nagel sub E. Oestlund 4584 AMES(51847)! *US(1805098)! Plan Juan Martnez, camino Reforma-Ayozintepec, 80 m. Comprada al Sr. Eugenio Hilario Justo, quien la colect expresamente para nosotros. 19-III-1997, M. Soto 8120 y A. Cibrin *AMO(sterile)! TABASCO: [cf., sterile] Balancn, carretera no. 25, km 45 del entronque con la carretera E.W.O. hacia la carretera W-10, 10 m s.n.m. selva mediana subperennifolia primaria, asoc. Manilkara sapota vul. “vainilla”, 06-12-1975, P.E. Valdivia 2063 XAL! CHIAPAS: [cf., fruits] La Cueva, al NW del Rancho Corocito, Reserva del Ocote, Mpio. Ocozocuautla. Alt. 770 m s.n.m., selva mediana perennifolia, primaria, suelo negro delgado con rocas calizas, ruderal, hierba, perenne, 3 m, escasa, fruto verde, nom. vul. vainilla, 29-04-1983, *XAL! Mpio. Ocosingo, Estacin de Biologa de Chajul, Sabana I, a unos 3 km del Ro Lacantm, bosque sabanoide con Roupala, Byrsonima, Scleria , Pteridium , ca. 200 m s.n.m. 16’ N, 90’ O. 13-IV-1997, M. Soto 8361 *AMO(buds)! CAMPECHE C.L. Lundell 1070 F(700398)! Selvas medianas subperennifolias (ManilkaraChryosophila) y bajos inundables, entre el Ejido 20 de Noviembre y las ruinas de Ro Bec, Reserva de Calakmul, Vida, slo una vista con botones. Los mayas del ejido no la conocen. 11-IV-1995. M. Soto, E. Martnez, G. Tavera, et al. 7656 Laguna, selva mediana subperennifolia con Cryosophila argentea, ca. 200 m altitud, 14 abril 1995, tpalos verdes, labelo crema con apndices anaranjados, frutos aromticos, fragancia a vainilla y coco, M. Soto 7667 AMO(x2, also in spirit)! same data, [fruits], M. Soto 7670 *AMO! Reserva de Calakmul, ca. de Nuevo Becar, en bajo, 13-IV-1995. M. Soto 7681 selva mediana subcaducifolia-subperennifolia de Brosimum alicastrum, Protium copal, Platymiscium sp., con mucha Cryosophila argentea, 230 m s.n.m.; planta vigorosa a la cerraron cerca de las 12:30; 3 Eulaema se aproximaron a

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.320 LANKESTERIANA *AMO! AMO(in spirit)! Ejido El Refugio, selva inundable de Bucida buceras a la orilla de la laguna, no la hay en el bajo de Haematoxylon contiguo. 230 m s.n.m., 17-IV1995, AMO(x2)! 200 m, 18’N, 89’W, 1 Feb 1076, 173 MO. QUINTANA ROO: Dense forest near shore of lagoon Chichankan, on shrubs. N part of lagoon, 88’N, 19’ W, ca. 50 m, 16 Aug 1935, O. Nagel sub E. Oestlund 4973 *AMES(51850, sterile)! *MO(1145555; sterile) *US(1805110; sterile). Mpio. F. Carrillo Puerto, camino al Ejido X’konha’, 4 m s.n.m., lat. 19’N, long. 88’W, selva mediana perennifolia primaria, clido hmedo, suelo pedregoso de color negro con mucha materia orgnica, asoc. J.S. Flores, E. Ucn 8236 CICY *XAL! En C. Vallarta, a 17 km al oeste Selva mediana con Manilkara, Vitex y Thrinax, 17 de abril de 1983, E. Cabrera 4611 y H. de Cabrera *AMO! MEXU! [cf., sterile] A 16 km al S de la terminal del Ferry, cerca de la entrada a Palancar, selva baja a mediana con abundante Lonchocarpus, Dalbergia sobre tronco, 22/Nov/1982, E. Cabrera 9768, O. Tllez, y E. Linares MEXU(421555)! La Pantera, 1997, Carnevali s.n. AMO(sterile)! YUCATAN: San Antonio, Rancho al sur de Pixoy, lat. 20’N, long. 88’W, alt. 22 m s.n.m., selva baja caducifolia, secundaria, en la orilla de una mensura, suelo moreno, pedregoso; abundancia regular, hierba, 6 m, perenne, fruto verde; obs. provoca comezn en la piel, 12-081983, E. Ucan 2761 CICY *XAL(fruto)! Mpio. Valladolid, Ebtn cabecera rumbo a Pixoy, lat. 20’N, long. 88’W, alt. 22 m s.n.m., selva baja caducifolia, secundaria, en la orilla del camino, suelo moreno, abundancia regular, hierba, E. Ucan 2463 *XAL! GUATEMALA: : [sterile] climbing on dry pine slope; leaves coriaceous, dull green above, paler dull green beneath, stems dull olive-green, warty rugulose, with a sulcation on each side, between Milla 42.5 and ridge, April 1, 1949 J.A. Steyermark 38539 F(1043863)![cf., fruit] A 8 km al NO de El Estor, 210 m s.n.m., hierba trepadora con fruto, sabana,30 agosto 1988, Stevens MEXU(480867)!MO(3656561)!Punta Palma, Sto. Toms, 100 m de la entrada de la playa por el lado norte, 3 4 plantas en la playa, 22 febrero 1998, M. Dix sub M. Soto 8611 AMO! : [cf., sterile] Climbing, stems terete, deep green, savanna north of Concepcin, 3-5 miles southeast of Finca Yalpemech, near Alta VerapazPetn boundary line, alt. 100-110 m, March 23, 1942, J.A. Steyermark 45233 *AMES(sterile, 63988)! F(1195510)! : [cf.] Jocol, climbing up trees, wild species of Vanilla 100 ft, Jan 30 1921, H. Johnson 1178 AMES(22753, sterile, perhaps V. cribbiana)! PETEN Tikal National Park, Bajo de Santa F, salida de Arroyo Corriental, in tintal on Aguada Trminos road, March-June, 1959, C.L. Lundell 15940 *LL(fruit)! same data C.L. Lundell 15818 *LL(fruit)! BELIZE: : Maskall, Dec. 1933, P. Gentle 1063 *AMES(40496, sterile)! NY(fruit)! TOLEDO Alta Vista Road, November 2, 1953, P.H. Gentle 8054 10 km south of Siuna; elev. 250 m. Vine; scrambling over D. Neill 4219 SE(049333)!9. Vanilla martinezii Soto Arenas, sp. nov. TYPE: GUATEMALA Livingston, El Golfete, a 20.4 km al NE de Ro Dulce por lancha camino a Calix, bejuco herbceo, mediana perennifolia ‘swampo’, 15’’N, 88’’W; E. Martnez S. 36410 y D. Alvarez, holo. MEXU!, iso. AMO! BIGUA! MO! Vanillae inodorae similis sed foliis membranaceissepalis rectis, petalis undulatis, labello subintegro vel quinque lateribus, fere longo quam lato, carinis humilibus inconspicuis ornato. Hemiepiphytic vine, leafy, up to 10 m high. Stems terete-subquadrate, somewhat keeled (in dried condition), ca. 2-4.5 mm thick; internodes 3.9-7 cm long. Aerial, free roots pale brownish, dorsiventrally compressed, 2-3.3 cm long, ca. 1 mm wide. Leaves petiolate, the petiole up to 16 mm long, canaliculate; blade elliptic, acuminate, base obtuse, somewhat conduplicate; membranaceous-chartaceous (in dried condition), 7.5-21 x 3.5-7 cm. strongly disimilar to the vegetative shoots, 26-50 mm long, a base), subsessile, the peduncle up to 13 mm long, the by 9-14 mm; with 1-2 peduncle bracts, clasping, ovate, concave, up to 10 x 7 mm. Floral bracts ovate, concave, obtuse to acute, apparently thin, 4-17 x 2-6 mm. Flowers successive, apparently 2 open at once, very showy, with rather spreading segments, tepals yellow green, lip white, ca. 5.5 cm high, 6.5 cm wide.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla321 FIGURE 10. Vanilla martinezii Soto Arenas. Based on the type, E. Martnez 22790, and M. Soto 8602a. Drawing by M. Lpez and M. Soto.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.322 LANKESTERIANA terete, smooth, arcuate to straight, 2-sulcate, the grooves almost straight, 52-53 mm long, 4 mm thick. Dorsal sepal lanceolate elliptic, apex acute, base obtuse, basally and apically grooved on the abaxial surface, obscurely and broadly keeled at the middle, apex, concave at middle, ca. 11-veined; 45 x 14.5 mm. Lateral sepals very slightly oblique, elliptic, the upper margin may be or not almost straight, acute to obtuse at apex, almost subcalyptrate, slightly recurved and curved upwards; basally obtuse, slightly canaliculateconcave, broadly keeled on the abaxial surface, the keel narrower and higher at apex, smooth, margins thinner, undulate, little pleated to straight, ca. 11-veined, 4647 x 17-18 mm. Petals arcuate, obliquely elliptic to lanceolate, somewhat arcuate upwards, apex long acuteacuminate, recurved-rolled, base acute, subtruncate, conspicually grooved all length of the abaxial surface, ca. 10 veined, margins undulate and pleated, especially near the apex, 43-47 x 11-13 mm. Lip fused basally to the column ca. 7 mm, arcuate, funnel-shaped, the lateral lobes-margins erect and forming a gullet obscurely trilobed, trapezoidal-subpentagonal to suborbicular-subpentagonal, base truncate with widely rounded shoulders, apex truncate-rounded, minutely mucronate, appearing triangular in natural position, axially grooved on the abaxial surface; 42-44 x 44-47 mm, ca. 27-34-veined, veins branched above the middle; callus lip length, made up by a broad, 7 mm wide, the base to near the middle, progressively becoming ovate-triangular in cross section and then divided in ca. 11 low, sinuous, rugose-warty, inconspicuous keels reaching the apex as a narrow high, almost smooth keel; the lateral ca. 10 veins (on each side) slightly raised and minutely rugose-warty; lateral lobes-margins almost margins slightly undulate-pleated. Column relatively short, strongly arcuate, the basal part forming a rather abrupt 60 angle with the apical part, semiclaviform, apex dilated, smooth, ca. 28 mm long (across de arch); vertical wings oblong, inconspicuous, 4.5 x 1 mm. Anther strongly attached to the clinandrium by a broad, Stigma a cavity, the midlobe concave, oblong, perpendicular to the column axis. Fruit clearly dehiscent, opening by 2 sutures, leaving unequal valves, 2-3 and 7-10 mm wide respectively, non-aromatic except by a resinous faint smell, dark brown, blackish inside, with ellipsoid seeds with slightly warty surface; 9.5-15 cm long, 6-8 mm thick before dehiscence. Fig. 10. DISTRIBUTION : Known only from eastern Guatemala, but also to be expected from adjacent wet areas of Belize and Honduras. It may also be native to Mexico. DNA from a sterile specimen of a membranaceous Vanilla collected in Crucero Corozal, in the Selva Lacandona, Chiapas, Mexico, was sequenced (ITS, matK), and it shows a strong relationship to V. martinezii, yet it seems different from the sympatric V. inodora; we suppose that it belongs to V. martinezii, although its sequences are somewhat divergent from the Guatemalan material. This locality is known to have populations of very thermophilous plants, that are very rare in other rain forest areas of Mexico (e.g. Lacandonia schismatica, Chysis limminghei, Warrea costaricensis, Ligeophila clavigera, Specklinia haberi, Maxillaria alba); furthermore the substrates in this ECOLOGY : In lowland, wet, swampy areas of high rainfall. Locally abundant. Flowering in February and July. As far as we know, this vine grows only on islets in the delta of the Polochic River into Lake Izabal and similar habitats near the coast in the area known as that we have examined, the midlobe of the stigma is perpendicular to the column body, and the anther is also protruding. The fruit set in Vanilla martinezii is very high (up to 53% in a clone) which suggests that it could be self-pollinated. Vanilla martinezii is known only from two or three nearby localities. Vanilla martinezii is a species of the membranaceous group; which, together with V. costaricensis, V. inodora, and V. sarapiquensis, are the only members of this clade in Mesoamerica. It is different from other species by the following combination of characters: membranaceouschartaceous leaves, short, lax, almost subsessile

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla323 with straight sepals, undulate petals, and by the huge subentire to subpentagonal, free lip, about as long as wide, and with very low and inconspicuous axial keels. Both V. costaricensis and V. inodora have elongate like bracts. Vanilla inodora is also easily distinguished by its cushion-like, axial callus and its emarginate lip; V. costaricensis is more similar to V. martinezii, the lip surface is adorned with a rugose sculpturing, especially at the sides of the lip. The most similar species is V. guianensis Splitgerber, from Guyanas and Amazonia (better known by its synonyms V. acuta Rolfe and V. latisegmenta Schweinf.). Both have membranaceous-chartaceous non-foliaceous bracts, subentire lip which is strongly veined, and similar column morphology with basal wings. However, in Vanilla martinezii the axial keels at the apex, some of them extending to the basal, lateral sides of the lip and the latter is not as trilobed at apex; in V. guianensis the callus is formed by 5 basal keels and 3 rather prominent ones, but it is not or only scarcely verrucose, and the lip apex is trilobed. Vanilla martinezii seems to be another endemism of the wet lowlands around Lake Izabal, a site which has been proposed as a primary refuge for a diverse V. guianensis make its distribution even more interesting. OTHER RECORDS : GUATEMALA: : Creek Lagarto, Ensenada de los Lagartos, El Estor, 2 m s.n.m., bejuco, 1988, MEXU(480869)! Lago Izabal, desembocadura del Ro Polochic, Creek Lagarto, al SW de El Estor, selva mediana inundable con Pachira aquatica , sobre suelos con mucha materia orgnica (peat), con Epidendrum stamfordianum, sphacelatum, O. luridum, Pleurothallis marginata, P. sertularioides, Maxillaria crassifolia, M. elatior, Gongora aff. quinquinervis, Coryanthes picturata, Myrmecophylla brysiana, Sobralia decora; cerca del nivel del mar, ca. 15’N, 89’W; comn, hasta de 10 m de alto; 24 febrero 1998, M. Soto 8601a AMO! mismos datos, 23 cpsulas de M. Soto 8602a AMO! The next specimen from a nearby locality is sterile, but it matches V. martinezii in vegetative aspect; however it could be V. inodora Schiede, usually with broader leaves: 15-31, 1922, P.C. Standley 24554 10. Vanilla odorata C.Presl, Reliq. Haenk. 1: 101. 1827 [1830]. TYPE: [ECUADOR:] Hab. in Guayaquil, Haenke, holo. PR(305753)! iso. (x2; 305751,305752) [all sterile and mounted with fragments of a Dimerandra species]. V. ensifolia Rolfe, Kew Bull. 1892, p. 141. Aout, Goudot, K! [with a drawing of the other -Patia, presented by Mr. J. Hanbury 1884” preserved)! Epidendrum vermifugum Mex. ed. 2: 201. 1894. Lectotype: (Soto Arenas, 1994): “Epidendrum vermifugum, Maldonado (4358) MA [sterile]!; isolectotype “Epidendrum vermifugum de Mxico absque “Epidendrum uni C OMMON NAMES : “vainilla Tlatepusco” (Usila, Oax.); “Vainilla de T eutila”, Humboldt. Hemiepiphytic vine, branching, leafy, up to 6 m high. Stems whitish dots, 4-6 mm thick; internodes 7-10.5 cm long. Terrestrial roots conspicuously pubescent; aerial, free roots terete, green to pale green, 1 mm thick; attaching, Leaves subsessile; the blade lanceolate to ensiform, very narrow, sometimes oblique, acuminate to long acuminate at apex, very dark green on both surfaces, 8-13 x 1.5-2.7 cm; ca. 1.8 mm thick in fresh. rachis 30 mm long, 6 mm thick. Bracts subsessile, concave, membranaceous, subsessile, progressively smaller, ca. 8 x 5 mm. Flowers successive, 1-2 open at once, with spreading segments, ephemeral (from 7:00 to 16:00 hrs), showy, tepals whitish green, translucent, lip greenish white, the throat striped with pale yellow lines, callus white, column white; ca. 8 cm high, 7 cm wide;

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.324 LANKESTERIANA FIGURE 11. Vanilla odorata Presl. Based on M. Soto 6617. Drawing by R. Jimnez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla325fragrance weak, green, fresh. terete, smooth, ca. 3 mm thick. Dorsal sepal linear to oblanceolate, apex acute, rounded, subcalyptrate, base attenuatesubunguiculate, concave, 9-veined, 47-54 x 8.0-8.3 mm. Lateral sepals narrowly elliptic to oblanceolate, oblique, apex subacute, rounded, subcalyptrate, base attenuatesubunguiculate, concave, ca. 12-veined, 44.5-52 x 9.5 mm. Petals obliquely linear to oblanceolate, somewhat arcuate, apex obtuse, rounded, sometimes emarginate, keel at the abaxial surface, ending in a cylindric, free, adpressed process, ca. 1 mm long; ca. 11-veined, 46-53 x 7 mm. Lip attached to the column along the margins of the basal half (ca. 23-29 mm), tubular, cymbiform, spread out 42-49 x 19-26 mm; long unguiculate, the claw dense papillose at base and apex on the the inner surface, the papillae digitiform, unicellular, 13-14 x 2.7trilobed, ca. 20-veined, the veins branched in the distal third and slightly thickened; the lateral lobes narrow (cilia 1-3.5 mm long), ca. 25 x 8-9 mm; midlobe approximately subquadrate-semiorbicular, the margin undulate-crispate, laciniate, obscurely emarginate, 7-9 x 7-11 mm; penicillate callus at 29 mm from the base, 3-4.5 x praemorse, retrorse, laciniate scales, sometimes united each to other along the lateral margins, with some adpressed processes on the surfaces; continuous towards the lip apex with 3 inconspicuous keels, progressively x 2 mm, with 3-4 conic, prominent, retrorse papillae and many small, elongate warts; other warts on the veins of the lateral lobes. Column very elongate and slender, semicylindric, 33-38.3 mm long, 2 mm wide; ventral bigger towards the apex, absent at the basal third; apex dilated, with vertical wings, inconspicuously trilobed, ca. 2 mm long, 3 mm wide; clinandrium prominent, transversely oblong, concave, ca. 2 x 3 mm. Stigma somewhat convex blade, ca. 2 x 3 mm, covering and parallel to the lateral lobes, quadrate, slightly divergent each to other, ca. 1 x 1 mm. Anther versatile, attached body urceolate in outline, with 2 upper, ovoid, small, divergent lobes; 3 x 2.5 mm. Fruit narrowly cylindrical, slightly compressed, attenuate at apex, dark green, 1717.5 cm long, 8-10 mm thick; strongly fragrant, aroma similar to that of V. planifolia . Fig. 11. DISTRIBUTION : Mexico (Veracruz, Oaxaca, Tabasco, and Chiapas), Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, and probably Brazil. ECOLOGY : This species can be common in secondary vegetation derived from tall, evergreen, tropical forest and rainforest, in gaps or in slopes with cleared canopy in primary forests. It is known from the sea level to 650 m altitude. Flowering time is mainly in April and May, but the vanilla growers in the Oaxacan Chinantla The narrow, lanceolate to ensiform, long acuminate leaves, thin stems, translucent whitish-green tepals, white-greenish lip with the throat striped with pale yellow and the laciniate margins of the lip are characteristic. Vanilla insignis is very similar, but it has thicker, sulcate, rugose-papillose stems, broader, xeromorphic leaves, and the retrorse appendages of the callus are much more numerous, bigger and orange-yellow. Mesoamerican specimens of V. odorata are very similar to that illustrated by Blanche Ames (Ames, Sched. Orch. 9: 1-6. 1925) based on Ecuadorian material and supposedly close to the type; perhaps the segments are slightly broader and the wings of the can be shorter in the Ecuadorian and Colombian (e.g. Fonnegra et al. 1784) material. With these bases, the small differences do not seen to warrant the recognition of separate taxa. Some specimens of Vanilla odorata from Belize V. hartii Rolfe, and a picture of V. odorata was labeled as V. hartii in “Native Orchids of Belize” (McLeish et al., 1995). Soto (1989) reported V. odorata from Mexico based on a specimen with buds from Quintana Roo (), but that specimen actually belongs to V. insignis. Humboldt cited a “vanilla de Usila”. From his

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.326 LANKESTERIANA description it seems that the involved species was V. odorata, which is well-known by the Chinantecan Indians of the region, and grown on a small scale at the present time. At the beginning of the 19th century some vanillas were used as vermicide in Mexico and Cuba, and that In the same way, the name “lombricera” was applied to Vanilla odorata produces aromatic fruits, similar to those of V. planifolia, with a strong, pleasant fragrance; they are very much appreciated in the regions where the species is wild. In Chiapas they are occasionally more common than V. planifolia, and actually it is the only vanilla collected for its beans in the Selva Lacandona, as in many other areas of Tropical America. In northern Oaxaca the species is grown as a curiosity in the plantations, intermingled with V. planifolia; the growers mention different cultural requirements from those given to V. planifolia, since it needs stronger sunlight. Although the fragrance is appreciated, the by fungi and because they dehisce if treated the same way as the fruits of V. planifolia; open fruits of vanilla usually bring a lower price. Vanilla odorata is found in dry to damp sites, usually in areas with higher rainfall than those of V. planifolia, and seems to be tolerant to a wide range of light conditions. Vanilla odorata is undoubtedly a species to be considered in any breeding program with commercial vanillas (see comments under Vanilla tahitensis, in “Excluded species”). OTHER RECORDS : MEXICO: : [cf., sterile] “Vainilla”. Vine on “Jimba” shrubs and on trunks of trees. Leaves leathery, glabrous. Common in some areas in dense 30-60 m, February 1937, L. Williams 8919 F(897099)! OAXACA : In sylvis umbrosis prope Lobani, Chinantla, Oajaca, Liebmann 6142 W(11759)! Tuxtepec, Ejido Chiltepec, “vainilla”, domina Lonchocarpus, M. Sousa 947 MEXU [sterile]! Dto. Tuxtepec, Usila, Arroyo Iguana, vainillal de Silvano Bautista, ca. 250 m s.n.m., 24 abril 1992, *AMO(x3; illustration voucher)! Dto. Tuxtepec, Mpio. San Jos Independencia, Cerro Clarn, en el extremo SW de la Presa Temascal, ca. 120 m s.n.m., 29 abril 1994, AMO! AMO(in spirit)! same data, abril 1994, U. Snchez s.n. y 6 AMO(in spirit)! Mpio.Valle Nacional, Arroyo de Banco, vainillal del Sr.Lzaro Prez Justo, 250 m s.n.m. plagas y patgenos que V. planifolia, 23-IV-1997, M.Soto Usila, en cafetal, “vainilla tlatepusco”, 23, abril 1995, M. Hernndez s.n. Sochiapa-San Juan Lalana, 1.2 km antes de San Juan Lalana, 210 m s.n.m. Cafetal derivado de selva alta perennifolia, en de Galino Tllez, 18-III-1997, AMO(fruit)! Dto. Tuxtepec, Mpio. San Jos Independencia, Cerro Clarn, en el extremo SW de la Presa Temascal, ca. 120 m s.n.m., 29 abril 1994, AMO! AMO(in spirit)! CHIAPAS: alrededores del sitio arqueolgico de Bonampak, vegetacin secundaria derivada de selva alta perennifolia, 350 m s.n.m., abril 1981, M. Soto 1001 AMO! Mpio. Ocosingo, km 5 del camino del Crucero San Javier a Bonampak, vegetacin secundaria derivada de selva alta perennifolia, sobre suelos rojos arcillosos, ca. 350 m s.n.m. 26 junio 1996 AMO(fruit)! Mpio. Ocosingo: Estacin de Biologa Chajul, en el borde del Ro Lacatm; camino a Arroyo Miranda, selva mediana subperennifolia inundable sobre terrenos planos con Scheelea y Sabal, ca. 180 m s.n.m., muy escasa, 21 junio 1996, M. Soto 7950 y R. Solano AMO(sterile)! BELIZE: Trail through light jungle, near Camp 2. Alt. 2000 ft. Wiss (vine) with glossy green succulent leaves and C. Whiteford 1316 MO(2584013)! GUATEMALA: : [cf., sterile] Twining vine; leaves deep green, succulent; fruit green, thick and succulent, twisted, exuding clear thick liquid when crushed; faint vanilla odor. Quebradas, 19-22 May 1919, H. Pittier 8589A NY! US(1013492); : “Vainillita” Chirujija Oxec.; near the Finca Sepacuit, April 23, 1902, *US(408445)! PETEN: “Vainilla”. Fleshy vine, Tikal National Park, Tikal, in botanal north of hotel, January 20, 1961 E. Contreras 1841 *LL(fruit)! La Libertad and vicinity, Aug.-Nov. 1933, M. Aguilar H. 164 *AMES(40519; steril)! HONDURAS : COLON: Capuchin site east, mangrove forest. 1.8 mi strip on the north bank of rio Guaimoreto between old bridge and opening of Laguna Guaimoreto 4.5 mi NE of Trujillo on old road to Castilla. Lat. 15’’N; Long. 85’’W. 2 Feb 1981, J. Saunders 1008 *LL(sterile)! *SEL(038496, fruit)! Lancetilla, 150 ft., NY! NICARAGUA : SEGOVIA : E of Jalapa, elev. 1600 ft., May-June, A.H. Heller 6106 SEL(013289; 003851, drawing, fragments)! “Segovia Prov.”, A.H. Heller s.n. F(1598348)! without data, A.H. Heller s.n. SEL(003851)! : [sterile]” Vainilla”, bejuco, sobre rboles, Guamil de segunda clase. Area de Ocotal, Ro Grande, Ro Grande, Alt. 0-15 m, Abril 23, 1949, A. Molina 2312

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla327 F(1364505)! Cerro Waylawas, ca. 13’N, 84-49’W, elev. ca. 100-268 m; sheer dog tooth limestone peak and plain on E side of peak. Pendant epiphyte, sterile. 16 March 1978. W.D. Stevens, B.A. Krukoff7385 SEL(049332; cf., steril)! COSTA RICA : ALAJUELA: Upala, San Jos, Alrededores de Laguna Las Camelias, 10’N, 85’’W 100 m, bejuco trepador, 19 Nov 1987, G. Herrera 1346 INB(sterile)! MO(3709189)! LIMON: Hamburg Finca, on the Ro Reventazn below Cairo, altitude about 55 meters. Large epiphytic vine; laeaves and fruit dark green. Vanilla nevermanii, “Vainilla”. Febr. 19, 1926, Valerio No. 48917 *AMES(32665; fruits)! *US(1309433)! [cf.] Hamburg Finca, on the Ro Reventazn, F. Nevermann s.n.PANAMA: PANAMA : [cf.] Barro Colorado Island, 10 Jan. 1940, F.W. Hunnewell 16433 *AMES(sterile, 87907)! COLOMBIA: ANTIOQUIA : Hierba trepadora, tpalos externos verde amarillosos, internos amarillosos. Carretera MutatPavarand, entre Haciendas La Esperanza y Mocar, 150 m s.n.m. Marzo 6 1987. R. Fonnegra, F.J. Roldn, J. Betancourt, B. Echeverry, O. Escobar 1784 K! *MO(3592235)! PERU: LORETO: Leticia on the Amazon River, September 1929, L. Williams 3161 *AMES(43483; sterile)! Herbaceous vine. Yurimaguas, lower Ro Huallaga; alt. about 135 m; dense forest, August 23-September 7, 1929, *AMES(43484; old BOLIVIA: IXIAMAS : Best kind here, fruit fragrant, vine, common, climbing over small trees in damp forest, 15-20 ft, 1000-1500 ft alt, Dec 13 1921, O.E. White 1115 *AMES(28024; fruits)! BENI: Rurrenabaque. Rank growing species; swamp woods, common. Fls. said to be white, fruits make good vanilla. 900-1000 ft alt., Dec 3, 1921, O.E. White 1821 Schweinfurth, Orch. Peru, Fieldiana 30(1): 43. 1958; Hamer, Ic. Pl. Trop. pl. 1193. 1984; Soto Arenas, Orqudea (Mx.) 13(1-211. Vanilla phaeantha Rchb.f., Flora 48: 274. 1865. TYPE: Cuba, C. Wright 3351 W; AMES(71001, iso.)! BM(another specimen is V. cf. poiteai)! G(7889/109)! K(iso.)! Vanilla planifolia var. macrantha Grisebach, Vcat. Pl. Cub. 267. 1866. C OMMON NAMES : “Tapia” (Panama). Hemiepiphytic vine, branching, leafy. Stems terete, smooth, green, 4-12 mm thick (in dried condition); internodes 8-15.5 cm long. Aerial, free roots, terete, pale brownish, up to 10 cm long, ca. 2 5 mm wide. Leaves subsessile, the petiole twisted, canaliculate, up to 8 mm long; blade narrowly oblong to narrowly elliptic, base rounded-subcordate, apex acute-apiculate, chartaceous (xeromorphic), green, margins revolute, 10.3-19 x 2.2-5 cm. thick, peduncle 15-29 mm long, rachis, 11-15 mm. Flowers successive, 1 open at once, ephemeral, the segments spreading, at least 75 mm long; tepals greenish-white to cream, lip white stained with brownish at edges or orange-yellow. straight to arcuate, subterete, smooth, 42 mm long, ca. 4 mm thick. Dorsal sepal long oblanceolate acute-subacute, rounded at apex, subcalyptrate; base long unguiculate, attenuate, canaliculate, the claw ca. 24 mm long, ca. 3-4.5 mm wide, blade concave, ca. 10.5 mm wide, 62-74 mm. Lateral sepals strongly oblique, long oblanceolate, arcuate, apex acute, subcalyptrate and minutely warty on the apex of the outer surface; base long unguiculate, attenuate, canaliculate, claw ca. 22 x 5 mm; blade concave, 12-12.5 mm wide, ca. 14 mm. Petals long linear, arcuate, slightly sigmoid, apex subacute-obtuse, base long attenuate, canaliculateconduplicate basally the rest concave, with an elevated triangular, acute, terminal process ca. 1 mm long, ca. 10-veined, thinner than the sepals, 61-73 x 8.5 mm. Lip attached to the column along the margins of the basal half (ca. 42-52 mm), tubular, trumpet shaped, conspicuosly cymbiform, deepest near the middle, axially grooved on the lower surface, when spread out 62-72 mm; unguiculate, the claw canaliculate, with two rows of trichomes on the apical half, 26-34 x ca. slightly undulate, ca. 36-veined, apex emarginate; obliquely long obovate, rounded 30-33 x 12-14 mm; midlobe distinct, transversely oblong, the margins x 11 mm; penicillate callus at 40-45 mm from the base, made up by ca. 12 congested, retrorse, aproximately united each to other along the lateral margins, ca. 9 x 5.5-6 mm, with an apical callus, low, inconspicuous, thickened. Column elongate, conspicuously sigmoid, 51-60 mm long; ventral surface lanuginose towards the

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.328 LANKESTERIANA FIGURE 12. Vanilla phaeantha Rchb.f.. Based on R.S. Williams 588 O.F. Cook s.n. SEL! with leaf) and C.W. Powell 412 Dwyer et al. 4691 (column). Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla329 apex, below the stigma; apex dilated, vertical wings somewhat bilobed, the lower lobe falcate, ca. 2 x 3.5. Stigma trilobed, with a convex, midlobe, ca. 3 x 4.5 mm, lateral lobes transversely oblong. Fig. 12, 16D. DISTRIBUTION : U.S.A. (Florida), Cuba, Lesser Antilles, Venezuela, Trinidad, Mexico (Yucatn and Quintana Roo), ?Costa Rica, and Panama. It is unknown if similar plants found in Ecuador, Guyanas and Brazil belong here or to V. bahiana. ECOLOGY : In Mexico this species grows in tropical deciduous or semideciduous forest near swampy areas, at about 100 m above sea level. Mexican specimens have been recorded. In Panama the species grows in coastal vegetation and in savannas with Curatella, Clethra, and Roupala. A previous report of V. phaeantha from Central V. calyculata. Vanilla phaeantha is very common in the West Indies, but it has been scarcely recorded from the mainland, except from Florida. However, it reaches the coast of Venezuela (G. Carnevali, pers. com.), and also the Yucatan Penninsula and central Panama, in our study area. Mr. Neal Byrd, owner of Finca La Gavilana, Province of San Jos, Costa Rica, cultivates a Vanilla phaeantha that was apparently collected from specimens of this species that have been available to us are from Panama, where it seems to be common on the The oblong-elliptic, xerophytic, usually greenyellowish leaves are usually shorter than the internodes. the lip is white with yellow stripes. It is related to V. bahiana Hoehne from Brazil. From the V. planifolia and V. odorata complexes it can be distinguished by its smooth disc.O THER RECORDS : MEXICO: YUCATAN: Cenote Mucul, Cerca de Sotuta, selva baja caducifolia, ca. 100 m s.n.m., G. Carnevali 4885 (cultivated specimen, not preserved, ITS sequence)! QUINTANA ROO: Mpio. Othn P. Blanco, Ejido Caobas, alrededores de la Sabana del Jaguactal, unos 21 km al sur de la carretera principal Xpujil-Chetumal, A. Cibrin 21, 23, R. Jimnez, G. Carnevali AMO(sterile)! COSTA RICA : LIMON: Guapiles, Los Diamantes, collected 26-12-2000, cultivated at Finca La Gavilana, Ro Savagres basin, N. Byrd II-A-1-5 PANAMA: PANAMA : Vine climbering up trees in low thicket. Farfan Beach; roadside thicket adjacent to beach. 30 March 1969, J.D. *MO(1980689)! Balboa, “Tapia”, alt sea level, grows out in much sunlight, C.W. Powell 412 *AMES(28219)! Ancon Hill, June 4, 1923, W.R. Maxon 6779 *AMES(33689)! *US(1180005)! Bismark above Panama, Fls. nearly white except greenish apex; climbing on trees several yards, March 12, 1908, R.S. Williams 588 NY! *US(678159; det by Schlechter as V. pittieri )! Common, sea level, February 1924, epiphyte, Hort. Powell 137, C.W. Powell 3507 *AMES(28290)! Climbing vinelike on the Farfn Beach area, forward dune-high beach community, 18 June 1971, R.L. Lazor 5358 1966, E.L. Tyson 4106 the old Las Cruces Trail, between Fort Clayton and Corozal, December 3, P.C. Standley 29103 AMES(31440, sterile)! [cf.] Brushy slope; large herbaceous vine, common, Tumba Muerto Road, near Panam, January 6, 1924 P.C. Standley 29731 AMES(31439, sterile)! VERAGUAS : Carretera Panamericana, 4.3 km al W de El Higo, ca. 20 km al W de La Mesa, sabana de Curatella, Clethra y Roupala con selva riparia de Andira, Pithecellobium , Enterolobium y Acacia , bejuco con frutos jvenes, extendido sobre algunos arbolitos M. Soto 9920 PMA(ITS sequence)! DOUBTFUL LOCALITY: . Photo, April 15, 1925 O.F. Cook *SEL(011359; specimen; 111357 photo, the photo in envelope is of a membranaceous species)! 12. Vanilla planifolia G.Jacks., in Andrews, Bot. Repos. 8: t. 538. 1808. LECTOTYPE: West Indies, without proper locality. Introduced into England by the Marquis of Hon. Charles Greville. The plate 538 was prepared from Greville’s specimen and was chosen as the Lobus oblongus aromaticus Clusius, Exoticorum Libri Decem.: 72. 1605. Araco aromaticus Hernndez, Thes. Rev. Med. Myrobroma fragrans Salisb., Parad. Lond. 2: t. 82. 1807, nom. illeg. Based on the same specimen that served as the type of V. planifolia G. Jackson.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.330 LANKESTERIANAVanilla fragrans (Salisb.) Ames, Sched. Orch. 7: 36. 1924; Stehl, Fl. Descr. Antill. Fr. 1: 109. 1939; Portres, Enc. Biol. 46: 234. 1954. Vanilla sativa Schiede, Linnaea 4: 573. 1829. Holotype: “Baynilla mansa HispanoMexicanorum”. Hab. sponte Papantlae, Misantlae, Nautlae et Colipae inqque iisdem pagis colitur”; Schiede, May 29 [single leaf] BM! Vanilla sylvestris Schiede, Schiede, Linnaea 4: 573. 1829. Type: “Baynilla cimarrona, HispanoMexicanorum. Hab. Papantlae, Nautlae, Colipae”, not located. Rolfe, J. Linn. Soc. London 32: 446. 1896. [or probably V. gardneri but the type is not useful] Vanilla duckei Huber, Bol. Mus. Goeldi, Par, 5(2): 327. 1908. Type: A. Ducke GOELD. Vanilla bampsiana Geerinck, Bull. Jard. Bot. Lebrun 1459 BR! C OMMON NAMES : “Vainilla mansa”, “vainilla” (Veracruz), “Xanat”, “shanat”, “caxixnath” (Totonaco, Veracruz), “vainilla colibr” (Oaxaca), “tlilxchitl” antique Nahutl (Badianus manuscript), “Kuoley gm” (Chinanteco, Oaxaca), “zizbic” (Maya, Yucatn); “juju” (Tabasco). Hemiepiphytic or rupicolous vine, much branched, leafy, up to 10 m high. Stems dark green, whitish dotted, sometimes the new stems covered with a whitish wax (Oaxacan plants), (6.5)11-12(13) mm thick; internodes 8-11 cm long. Terrestrial roots pubescent, 1.4-1.6 mm thick; aerial, free roots terete, 2.5-3.0 mm thick; attaching, aerial substrate, 3 mm wide. Leaves subsessile, the blade, elliptic, oblong, narrowly oblong, usually with parallel margins, abruptly acuminate to subacuminate, somewhat oblique (variable), upper surface deep to pale green, paler on lower surface, 9.5-23 x 3.5-7.6 cm, ca. 1.3-2.4 mm thick in fresh. raceme, up to 26.5 cm long. Flowers successive, 1-2 open at once, segments variably spreading, most frequently the tepals ca. 40 with respect to the column; ephemeral, showy, tepals pale green to whitish green, lip pale cream-yellow to yellow with ochre in the throat and papillae; ca. 62-64 mm high, 39 mm wide, fragrance weak, sometimes a cinnamon can be appreciated, herbal, or white (e.g. Hymenocallis -like). arcuate at base, upper part straight, terete, smooth, basally white, mm thick. Dorsal sepal narrow elliptic to oblanceolate, apex subacute, rounded, subcalyptrate, base attenuatesubclawed, slightly concave, canaliculate basally, ca. 9-veined, 55-60 x 10-12 mm. Lateral sepals obliquely narrowly elliptic, to oblanceolate, apex subacute, slightly canaliculate basally, margins slightly involute, ca. 10-veined, 54-60 x 12-13.5 mm. Petals obliquely long oblanceolate, somewhat arcuate, apex slightly thickened, base attenuate; concave, canaliculate at ascending, free process, ca. 2 mm long; ca. 13-veined, 55-58.5 x 10-11 mm. Lip attached to the column along the margins of the basal half (ca. 31-35 mm), tubular, trumpet-shaped, very concave to cymbiform, slightly opening subtriangular, 18 mm wide, axially grooved on when spread out 49-55 x 24 mm; long unguiculate, the claw pubescent, trichomes short, yellow to ochre, 15 x ca. 36-veined, the veins branched in the distal third, and slightly thickened; the lateral lobes obliquely triangularthe midlobe; teeth less than 0.6 mm long, ca. 27 x 10.5 mm; midlobe approximately subquadrate-transversely oblong, emarginate to deeply bilobed, the margin undulate-crenate, 5 x 9 mm; penicillate callus at 3037 mm from the base, 5.8-6 x 4-4.5 mm; made up by scales, pale yellow, the distal ones bigger, sometimes united each to other along the lateral margins, continuous towards the lip apex with 2 conspicuous rows of papillae, and 2-4 inconspicuous rows, apical papillae bright green. Column very elongate and slender, trigonous-semicylindrical, 39-42 mm long, 2.5 erose, acute to rounded, membranaceous wings, 2 mm

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla331long, 4 mm wide. Stigma trilobed, the lobes emergent; midlobe a trapezoidal, convex blade; ca. 3 x 2 mm; covering and parallel to the lateral lobes, longitudinally oblong, slightly divergent each to other, ca. 1 x 1 mm. Anther versatile, attached to the clinandrium margin x 3 mm. Fruit straight to strongly arcuate, cylindricalsubclavate, thickened towards the apex, often 2-3 conspicuously sulcate in wild specimens; smooth in cultivated and many wild specimens; 10-30 cm long, 7-10 cm thick. Fig. 16E—F, 17D. DISTRIBUTION : Imperfectly known. Mexico (Veracruz, Oaxaca, Chiapas, Tabasco, Quintana Roo, and probably escaped in Yucatn and southern Oaxaca), Guatemala, Belize, Honduras, and Costa Rica. It is unknown if the plants in other areas are native or escaped from cultivation. For example, in Panama it is known mostly from Barro Colorado Island and San Blas Province, perhaps an indication that it is escaped there. The species seems to be escaped in Florida (cf. two areas well botanized by experienced collectors. Foldats (1969) indicated that the species is common in Venezuela, but did not cite records; neither did Garay and Dunsterville include it in their works on Venezuelan orchids. South American wild specimens V. planifolia and others from elsewhere outside Central America have proven to be collections from Ecuador that match closely the Central American, cultivated material. It is doubtful that V. planifolia is native in regions outside of Mesoamerica. Suposedly wild specimens of V. planifolia from Rio Palenque Center, Ecuador, have proven to be V. hartii. The original distribution in Mexico is also uncertain. Northernmost, wild-collected specimens come from the region of Crdoba, in Central Veracruz, some hundreds of kilometers southwards from the area where the species supposedly was domesticated in northern Veracruz and Puebla. Most collections come from northern Oaxaca, where great morphological variation has been observed; a single record from rare in Chiapas. There are several old collections from Yucatn, in the xerophytic thorn scrub of the northern part of the peninsula; we suspect that they represent escaped, old relicts from cultivated plants, since this habitat is very different from the moister forests where it is wild at present. ECOLO G Y : Dwelling in tall tropical evergreen or semievergreen forests, from 150 to 900 m, rarely to 1300 m altitude. The species seems to prefer moist forests, seasonally dry in spring, on calcareous terrain. It is absent in volcanic areas and in the wet tropical rainforests of Mexico. In moister areas it can be found in secondary, in March to April, but varies from year to year, and be a response to low winter temperatures, followed by strong sunny conditions in early spring. Vanilla planifolia faces severe conservation problems in the wild. There are very few and scattered locations known at present; the largest populations in northern Oaxaca have been completely removed as a source of cuttings to establish new plantations; it is also almost extinct in Veracruz, where only two clones have been located in recent years. Only four isolated specimens have been located in Chiapas. Vanilla planifolia is the vanilla of commerce, and the most widely planted species for its aromatic fruits. The nomenclature of this species is very complex, since its history began before the Linnean system of nomenclature. The problem has been solved by with a plate based on the original plant (leaving aside the statement of the identity of an earlier Plumier’s polynomial); it seems pertinent to reproduce here their observations: “Jackson regards Plumier’s breviori, corallino” as representing his V. planifolia. A study of this plate, however, convinces us that it represents Vanilla eggersii Rolfe. Salisbury based his Myrobroma fragrans also in Greville’s cultivated specimen, but cited in synonymy Epidendrum rubrum Lam. Since in his publication Epidendrum rubrum is an integral part of the protologue, he should have made a transfer of Lamarck’s epithet. It is of further interest that Salisbury equates or rather confuses Epidendrum rubrum Lam. with Plumier’s polynomial description

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.332 LANKESTERIANAreferable to Vanilla eggersii Rolfe.” Therefore, the widely used name Vanilla fragrans must be rejected, because it was based on an illegitimate name. The plate selected as the lectotype is not accurate enough to permit the recognition of the species; especially the much fringed lip suggests another species of the group, such as V. odorata or V. insignis. Hooker (1891) also expressed concern about the identity of this plate, and recognized the differences among Andrews’ and Salisbury’s plates. However, the same Greville’s individual plant was illustrated and precisely described a year earlier by Salisbury, when he published his Myrobroma fragrans, the plate and description are much more accurate, and undoubtedly belong to the widely cultivated, “Mexican vanilla”, discarding the possibility of it being another related species. The history of Vanilla planifolia has been reviewed by Dillon (1942) and Bruman (1948). The published plate (Flore d’Afrique Centrale, pl. 39) of the recently described V. bampsiana Geerinck, V. planifolia. There are no vanillas with lamellose lips and papillose apices outside of Tropical America. VARIATION AND CULTIVARS : Vanilla planifolia has proven of the lip papillae, intensity of fragrance, development of grooves on the fruit, self-compatibility, etc. This variation initially led us to consider the existence of more than one species involved; especially because of the uniformity of plants in cultivation (the cv. “mansa”) compared with wild-collected specimens. However, it is possible that the narrow morphological variation seen in cultivated plants must be attributed to reduced genetic variation (Cibrian, Soto Arenas, ). Studies of of different forms is composed primarily of the same compounds (1-2-dimethyl-cyclopentane, ethyl acetate, 1-8-cineol and ocimene-trans); 1-8-cineol especially is well known to be a strong attractant for euglossine bees. Since V. planifolia is pollinated by Euglossa viridissima and maybe other species of the same genus, the fragrant, attractive compounds can be considered forms. The growers in Veracruz recognize different types in V. planifolia, and these perhaps represent different genetic individuals (clones), since no genetic manipulation or breeding programs have been conducted in Mexico. These clones are better treated as cultivars. cv. “mansa” or “dura” This is the form widely cultivated in northern Veracruz, and probably in Africa and Asia; the plants from Asia (except the Philippines) and Africa are descendant of the original type specimen grown by Grenville and distributed to many botanic gardens during the 19th Century. It is unknown if more than one clone is involved, but the plants are very uniform in morphology. Different from many wild specimens are the non-sulcate fruits. cvs. “acamaya”, “rayada”, and “variegata”. The stems and leaves have yellowish stripes, alternating with the common deep green of the other cultivars (see Roullet, 1990). There are at least two different genetic individual with variegated plants (Schlter et al. 2007). There seem to be no differences in molecular markers between one of this clones and ‘Mansa’, suggesting that it can be a sport (due to somatic mutation). cv. “albo-marginata” A cultivar with beautiful leaves, margined with white (see Roullet, 1990). It has never been found in Mexico, and it is probably also a sport of Madagascan origin. cv. “oreja de burro” ? Vanilla sylvestris Schiede The cv. “oreja de burro” is occasionally grown as a curiosity in the plantations of the region of Papantla. to distinguish except by experienced growers; the stems are slightly thinner, the leaves rather hanging, because the more elongate petiole, elliptic, abruptly acuminate, concave, with more prominent veins, and fruit up to 30 cm long, usually bisulcate, but almost all the fruits are aborted in July, except sometimes when the pollen came from a different clone or from V. pompona; therefore, it is self-incompatible. Additional watering or feeding does not seem to maintain the fruits

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla333on the plant. The occasional fruits are longitudinally sulcate; it is probable that this cultivar was the base for Schiede’s V. sylvestris, since the sulcate fruits are cited in the protologue. It is said that the cv. “oreja de burro” grows faster and with more vigor than other cultivars of V. planifolia; furthermore the growers say that it is not severely affected by fungal diseases. Some of the indicated differences between “mansa” and “oreja de burro” 1993); however, the vegetative propagation of vanillas in the plantations has made many of them genetically uniform, and it is very probable that the observed differences are among clones and not between different taxa (species, subspecies or varieties) or indicative of a genetic substructure in V. planifolia. The “oreja de and it is also visited by Euglossa viridissima. Cuttings of “oreja de burro” are sold occasionally by unscrupulous growers to discourage the establishment of new plantations. We have observed hectares of plantations that were nonproductive due to large quantity of vines belonging to the cv. “oreja de burro”. Several wild collected plants are notoriously different from the specimens cited above, and it is very probable that they belong to locally differentiated populations. Among them, there are several specimens collected in Belize that look somewhat different, with of lip. These specimens have also rather elongate rachis Schipp S-971, M.A. Soto 8355, P.C. Standley 25064 are from Belize, Izabal, and Chiapas, from riparian habitats and they seem to be wild; however, leaf aspect is much more elliptic, and M.A. Soto 8355 has sequences of MAS 8355 is nested within the variation of V. planifolia, although with several autapomorphies, but we do not know if they represent only sequencing problems. Schlter et al. have shown that V. planifolia has three rather different genetic groups, the cultivated group, the wild specimens from Mexico to Quintana Roo and the wild specimens from Costa Rica. A specimen from Costa Rica [Esquinas forest. Area between the Ro Esquinas and Palmar, sea level. May 2, 1950. Flowers pale green. P.H. Allen 5532 SEL (011362)! ] is somewhat strange, the leaves are clearly petiolate and elliptic, acuminate, but not abruptly and the lip does not seem lobed, and the warts on the apex of the lip are very obscure. Our impression is that its morphology is intermediate between V. hartii and V. planifolia, and therefore it may be a hybrid.O THER RECORDS : MEXICO: : Rancho El Coscoln, El Escoln, Cerro Grande, Mpio. Papantla de Olarte, 350 m s.n.m., 20’N, 97’W, clima clido seco, suelo arcilloso-arenoso, plantacin de vainilla. Hierba del perianto amarillenta. Fruto con la base adelgazada. Aborta 50-100% de los frutos. “vainilla oreja de burro”, 25 Abr 1989, AMO(x2)! MEXU(678227)! Other names applied in northern Veracruz, such as, “negra”, “verde” or “amarilla” are apparently due to different exposure to light intensity. O THER RECORDS : UNITED STATES : FLORIDA : Dade Co. growing on trees in an extensive area in Cutler Hammock, on Biscayne Bay. Fls. greenish. May 15, 1976. Popenoe 47216 NY! Brickell Hammock, Miami, Dade County, J.K. Small , 19 Feb-March 22, 1915, NY! MEXICO : SAN LUIS POTOSI : Tanjasnec, Mpio. San Antonio, bosque tropical perennifolio, dooryard, vine, “vainilla”, planted, 4 May 1979, J.B. Alcorn 3006 *LL! MEXU! PUEBLA : Bejuco, fruto verde, “vainilla”, en cafetal, fecundada por jicotes, Yancuictlalpan, Cuetzalan, el fruto seco se usa como aromatizante y saborizante de atole y otros alimentos, F. Basurto y R. Patrn 263 MEXU! : Rancho El Coscoln, El Escoln, Cerro Grande, Mpio. Papantla de Olarte, 350 m s.n.m., 2029’N, 9733’W, clima clido seco, suelo arcilloso-arenoso, plantacin de vainilla, hierba 1989, AMO(x2)! MEXU(678228)! [cultivated] Predio Escoln, 12 km al NE de Papantla, 370 m s..n.m., trepadora, abundante, fruto verde, selva alta, secundaria, vainillal, suelo rendzina de color negruzco, 22 Junio 1987, A. Garca 3226 MEXU! [cultivated] Papantla de Olarte, 2027’N, 9719’W, “vainilla”; usos, para F. Rosas C.I.P. 709 *XAL! [cf.] San Andrs Tuxtla, El Viga, al E de Cerro Blanco, 28 May 1967, M. Pea 108 *AMES(11744, sterile)! Estacin de Biologa Tropical de Los Tuxtlas, ca. 200 m s.n.m., selva alta perennifolia, G.A. Salazar 2247 AMO(drawing)! AMO(exsiccata, en FAA)! Volcano San Martn, on trees in dense fore st, 5 May 1936, 9511’, 1835’, 600 m, J. Gonzlez

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.334 LANKESTERIANA sub E. Oestlund 5790 *AMES(51851, sterile)! Mpio. Atoyac, a un lado de la va del tren, cerca del puente, selva alta subcaducifolia de y Dendropanax arboreous , terreno crstico, sobre piedras y rboles, 2 mayo 1995, ca. 400 m altitud, M. Soto 7745 *AMO! Vainillal del Sr. M. Soto 7648 AMO! [cf., fruit] A 800 m al norte de la casa de Genaro, ejido Vista Hermosa, Mpio. Tezonapa, 1840’N, 9640’W, alt. 500 m, selva mediana subperennifolia, primaria, suelo oscuro, arcilloso, bejuco perenne, escasa, fruto verde, nom. vul. “vainilla”, 4-8-1986, R. Robles G. 977 *XAL! [cultivated] Papantla, suelo poroso bien drenado con abundante materia orgnica asociada a Gliricidia sepium , Erythrina berteroana , Bursera simaruba amarillenta, nom. vul. “vainilla”, “caxixanath” (Totonaco), 17-02-1985, *XAL! OAXACA : Vine 10 ft high in woods between Los Llanos de Ozumazin and Ro Chiquito, District of Choapam. Lat. 1735’, Long. 9555’, Alt. 700 ft., 1, May, 1939, R.E. Schultes and B.P. Reko 727 *AMES(9034)! Cultivada en los vainillales de Santiago Tlatepuzco, Mpio. Usila, ca. 500 m s.n.m., R. Medinilla s.n. AMO(en FAA)! Cerro Machete, Distr. Pochutla, 800 m, Feb. 1941. B.P. Reko 6279 blanquecina, acahual derivado de selva alta perennifolia, 3 km al W de La Esperanza, Mpio. Ixtln de Jurez, Sierra Norte, R. Lpez G. 2 AMO(16698)! same data E. Morales 1 AMO(16697)! Usila, Mpio. Usila, cultivada en los vainillales de la familia Bautista, cerca de la pista de aterrizaje, 90 m s.n.m. 25 abril 1992, *AMO(x 2; also in spirit)! mismos datos, aparentemente un clon con Hernndez AMO! mismos datos Hernndez AMO! Dto. Tuxtepec, Mpio. San Jos Independencia, Cerro Clarn, en el extremo SW de la Presa Temascal, ca. 120 m s.n.m., selva alta perennifolia, 1 mayo 1994. AMO! Instituto Tecnolgico Agropecuario, San Bartolo Tuxtepec, plantacin a cargo de Jess Prez Meza. Abril 1996. M Soto 8040 AMO! Ejido La Gran Lucha, Mpio. Valle Nacional. Vainillal particular de Nemesio Miguel Martnez. Plantacin proveniente de un slo clon que originalmente creca en las afueras de la comunidad. Plantacin de 1 ha, 2500 plantas, de derivado de selva alta perennifolia en ladera krstica. 17-III1997. “Ligey” (Chinanteco). * AMO! La Cueva, cerca de Cerro Verde, km 25.3 del camino 780 m s.n.m., muy hmeda, ca.1857’N, 9444’W. Plantacin de vainilla de Jos Roldn (aparentemente hay tres clones en la plantacin). 21-IV-1997. Tal vez el mismo clon que Soto 8477 . Flor muy cncava que huele a canela. M. Hernndez AMO! Mun. Santa Mara Jacatepec, predio El Aguila, al O de San Agustn, entrando por la Reforma, 28 km SO de Tuxtepec, carr. a Matas Romero. 1750’N, 9606’W. Elev. 550 m. Selva alta perennifolia. 19 Ene 1988. R. Torres AMO(sterile)! MEXU(x3; sterile)! TABASCO : cultivada en el Jardn Botnico de Puyacatengo, en Teapa, fotografa! (Alderete y Capello, 1988); [cf., sterile] Teapa, 0.34 km al E de la Universidad Agraria de Chapingo, 1731’31’’N, 9255’33’’W, selva alta perenifolia, 28 enero 2002, J. Calnico 21157 et al. MEXU! Poblado de Nacajuca, 20 m s.n.m, selva mediana perennifolia, primaria, suelo negro arcilloso con abundante materia orgnica, bejuco perenne, 3 m, escaso, nom. vul. “juju”, uso comestible, 3-10-1978, R. Ortega, W. Mrquez y B. Guerrero 858 *XAL! CHIAPAS : 17 km NO de Ocozocuautla, carretera nueva a la presa de Mal Paso, selva alta perennifolia, ca. 900-1200 m, J. Castillo 833 (specimen in cultivation)! Palenque, abril de 1892, Alt. 450 m, R.M. Historia Natural No. 7725 *AMES(21309)! 6 km al E de Emilio Rabasa, 1 km al NE de La Reyna, Ocozocuautla, 1655’N, 9338’40’’, 720 m, alrededores de una casa, huerto familiar y cultivos de maz, S. Ochoa 3659 CHIP! [cf., fruit] Reserva Ecolgica “El Ocote”, Mpio. Ocozocuautla, Alt. 750 m, selva alta perennifolia, primaria, suelo calizo, perturbada, hierba perenne, abundante, fruto vaina, trepadora, 14-021986, M. Palacios-Ros 2823 *XAL! Alrededores de la Laguna Ocotalito, cerca de Monte Lbano, selva baja perennifolia, 950 m altitud, M. Soto 2836, 2840 AMO(slides, in spirit)! Mpio. Ocosingo, Estacin de Biologa de Chajul, vereda que conecta la vereda a La Granja con la de la Sabana I, a unos 500 m del Ro Lacantm, selva perturbada con bejucos y Bactris , terrenos ondulados y arenosos pero ocasionalmente inundables, ca. 200 m s.n.m. 1608’ N, 9053’ O. 13-IV-1997. M. Soto 8355 AMO(sterile)! Estacin km de la estacin. 200 m s.n.m. 1608’ N, 9053’. En acahual escasa. 20 junio 1996. M. Soto 7947 AMO(sterile)! Estacin de Biologa de Chajul, junto al Ro Lacantun, acahual con aparentemente es el mismo clon que Soto 7947 , sobre Cecropia obtusifolia , muy escasa, vegetativamente igual a Soto 8355 , que es otro clon. Flor ms verde que las plantas cultivadas, hojas ms anchas no planas y fruto corto y trgono, muy largo. Probablemente silvestre. 15 de abril de 2000, M. * AMO(x3)! Selva del Ocote, km 44.7 de la carretera Ocozocuautla-Malpaso, cerca de la presa, manchn perturbado de selva alta perennifolia con Poulsenia armata, Ficus glabrata, Reinhardtia gracilis y Zamia splendens , muy hmeda, sobre areniscas, 275 m s.n.m., probablemente silvestre. 21 abril de 2000, M. Soto 9922, S. Maldonado, P. AMO! Nah, Mpio. de

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla335 Ocosingo; 27 km al SE de Palenque por la carretera fronteriza hasta el Crucero Chancal, despus 55.6 km por el camino de terracera hacia Monte Lbano, 1656’-17o1’ y 9133’9138’; 900 m de altitud; bosque tropical perennifolio: Planta 1999, A. Durn Fernndez 1166 MEXU! YUCATAN : [sterile] “Vainilla” Yuc., “Sisbic” Maya, Cenote de Mayana, Yucatn, 1866, A. Schott 215 F(276993)! In virgin xerophytic forest, April 1916, forests of Xbac, 23352 AMES(*71360, 71369)! *F(446854)! MO(948146)! NY! In virgin xerophytic forests, Izamal, April 1917, G.F. *F(466398)! *US(1268044; sterile)! [cf., poorly preserved] “Sisbic”, June 4, 1899, G.F. Gaumer F! Virgin forest near Chemax, on tree trunks, 29 Sep 1935, 8756’, 2039’, alt. 30 m, O. Nagel sub E. Oestlund 5050 *AMES(51846, fruit)! QUINTANA ROO : Isle of Cozumel, Cenote Cedral, on tree trunks in mixed forest of Achras zapota , 2 sep 1935, 8659’, 2021’, 5-10 m, alt., O. Nagel sub E. Oestlund 5001 *AMES(51852, fruit)! Ejido Hermenegildo Galeana, predios de Artemio y Constantino Lpez Pascual, 1810’33.2’’N, 8915’08.4’’W selva alta subperennifolia alternada. 10-V-2008. M. Soto 11370, L. AMO! WITHOUT LOCALITY : “1815 de Mexico, Epidendrum vanilla Pavn G! “Mexico, Mr. Cowan G! BELIZE : Machaca, common, 16 May 1934, *AMES(40477)! TOLEDO scented, in broken cohune ridge, Feeders Road, 14 Miles, San Antonio-Punta Gorda Road, April 20, 1949, P.H. Gentle 6721 greenish-white, scented, in acahual, on hill slope beyond San Antonio, January 30, 1952, P. Gentle 7557 MEXU(511464)! *LL(x2)! “Vainilla”, Fls. pale green, in cohune ridge, Bolo Camp, uper reach of Golden Stream, April 12, 1944, P.H. Gentle 4521 cohune ridge, near Jacinto Hills. December 30, 1944, P.H. Gentle 5106 *LL(x3)! Quite common along river bank here material as the squirrels get them. Flowers greenish yellow. Temash River. 100 ft alt., 30 ft, _ in. diam., 13th Mar 1935, W.A. Schipp S-971 *AMES(43554)! Maskal Pine Ridge road, P.H. Gentle 1234 AMES (*41360, *41361)! GUATEMALA : Blt. mattschwefelgelle, sehr himfalling. Frchte. Blht sehr reich. Am Rio Polochic ber Tucur, 900 m, Selten! 30.4.1882. F.C. Lehmann 1436 G! : crests down middle. Vicinity of Finca Yalpemech, near Alta Verapaz-Petn boundary line, alt. 100-120 m. March 24, 1942, J.A. Steyermark 45286 *AMES(62981)! *F(1195546)! PETEN forest about 2 km. 100 m south of village, May 3, 1961; E. Contreras 2239 *LL! : Large herbaceous vine, wet thicket, frequent, vicinity of Puerto Barrios, at sea level, June 2-6, 1922, P.C. Standley 25064 *AMES(22674, fruit)! EL SALVADOR : Finca Las Canoas, 700 m alt. terr., climbing trees, mostly oaks, April 11th, 1969, H.C. Clason sub F. Hamer 268 MO(3092269)! same data, near Ahuachapan, April 9th, 1978 H.C. Clason sub F. Hamer 776 *SEL(049262)! “Vainilla”, San Salvador, in garden, Vanilla seems to be cultivated in Salvador only as a curiosity, 1922, S. Caldern 518 *AMES(22663, sterile) *US (1151498; sterile)! HONDURAS : : [cultivated] “vainilla”, racemes more on Cupressus lusitanica in my yard, campus of EAP El A. Molina 34346 *MO(4248893)! ATLANTIDA : [cf.] “Vainilla”. Wet forest. Large herbaceous vine climbering over trees. Fls. pale green throughout. Lancetilla Valley, near Tela, altitude 20 to 600 m, Dec. 6. 1927, Mar. 20, 1928, P.C. Standley 55583 *AMES(36947, sterile)! F(581081)! Climbing on tree, Lancetilla Valley, 159 ft alt., 8/12/1934, *AMES(105299, sterile)! COMAYAGUA : [cf., fruit] “Vainilla”, frutos muy aromticos cuando secos. Planta 1300 m, Marzo 31, 1963, A. Molina 11777A F(1620024)! NICARAGUA : : Logging camp near Quebrada La Talolinga. Elev. ca. 170 m. ca. 1151-52’N, 8426-27’W. Tropical wet forest. Climbing vine fruit green, J.C. Sandino 1138 MO! COSTA RICA : LIMON : “Vainilla”. Large epiphytic vine; leaves thick; fruit green; Hamburg Finca, on Ro Reventazn below Cairo, altitude about 55 meters. Febr. 19, 1926, *AMES(32697, fruit)! *US(1309657)! in regione Atlantica: Musas G. Cufodonti 735 *AMES(37507)! W(1770)! Forest between Punta Manzanillo and Punta Nona, E of Manzanillo de Talamanca. Elev. ca. 8-12 m. 938’N, 8238’W. Vine appressed-climbing in lower part, the fertile branches pendent, festooning small tree in understory of open forest atop coral bluffs along coast just E of Punta Manzanillo; borne ca. 3-4 m above ground. Flowers whitish-green, without fragrance (11:30 AM). Fruits green. Leaves thick, subsucculent. Flowers and fruits, 2 May 1985, * M.H. Grayum CR(119858)! Pococi Cantn. Refugio de Vida Silvestre Barra del Colorado, Cerro del Tortuguero, summit ridge. 1035N 8332W, ca. 100-119 m. Vining to ca. 3.5 m above ground on small tree in secondary forest. labellum longitudinally pleated internally and more or less lacerate marginally. 22 January 1997, M.H. Grayum, B. INB! Cantn de

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.336 LANKESTERIANA Puerto Vargas, 094420 N, 824930 W, 2 m. Planta trepadora, creciendo a orilla de sendero. Frutos inmaduros verde -claro y maduros verde amarillento. Comn. 9 December 1993, INB! PUNTARENAS Ribera del Ro Esquinas. Boca Ro Esquinas. 084400N amarillentos. 22 December 1993, Quesada INB! CARTAGO : Pejivalle, Very common in forest 1924, G.H. Lankester 847 AMES(28561)! PANAMA : PANAMA : [sterile] Gross Lathrops trail, Barro Colorado Island, June 26, 1931, AMES(37507)! [sterile] Wet forest; large herbaceous vine; occasional, Barro Colorado Island in Gatn Lake, altitude 120 meters or less, Nov. 18-24, 1925, P.C. Standley 40897 *US(1251676)! Barro Colorado Island, Lutz Trail 300, vine, much scandent, seldom branching, sterile, July 10, 1970, T.B. Croat 11177 Island, Oct. 26 1931, O. Shattuck 230 AMES(71355, juvenile, sterile)! [cf., sterile] Wooded swamp; large herbaceous vine. Coln, January 9, 1924, P.C. Standley 30160 US(1225400)! SAN BLAS : Flowers pale green, April 28, 1933, G. Proctor Cooper 287 AMES(*40380, *71354)! *F(771391)! NY! *US(1557502)! REFERENCES : Hooker, Bot. Mag. 117: t. 7167. 1891; Rolfe, Journ. Linn. Soc. Bot. 32: 463. 1896; Duss, Ann. Inst. Col. Marsaelle 3: 601. 1897; Cogniaux in 16. 1952; Cabrera, T. 1999. Orqudeas de Chiapas: Salvador 2: 372-373. 1974; Hall, Fl. Nov.-Caledonie 8: 407. 1977. 13. Vanilla pompona Schiede, Linnaea 4: 573. 1829. TYPE: “Baynilla pompona HispanoMexicanorum”; Mexico, Papantla et Colipa, Schiede 1043, “Vanilla pompona nob. interim Vaynilla pompona Papantlensis in sylvis Papantla, Ja. 29” BM (lectotype; Garay Fl. Ecuador Orch.)!; isolectotype W (sterile)! Vanilla pompona is a variable, widely distributed species. We consider it rather to be a species complex. collected in central Veracruz. Mexican populations are apparently isolated from the rest of the populations of the species because there are no known populations in Chiapas, Guatemala, Belize, or El Salvador. The taxon reappears only in Central Honduras and Nicaragua. This geographical isolation is correlated and probably the reason that Mexican samples in the ITS tree form a clade with high bootstrap support. The Mexican plants also have slightly different morphology, since the lip deeply emarginate, but the general aspect of the vine and South America. reveal more differences, but there is a notable lack of V. pompona group. The rest of the samples of V. pompona have a paraphyletic relationship with the Mexican samples, but are more or less clust ered into two groups. One of the groups encompasses the samples from Honduras, Nicaragua, Costa Rica and Panama. Differently from the Mexican and the South American vines, rather small leaves, that is, they keep the juvenile stem morphology. Even though they do not form a monophyletic group within the tree it seems better to consider them another group within the species. All the South American samples have commom synapomorphic DNA characters, like peculiar nucleotide bases in certain positions and common deletions or insertions, although they also form a paraphyletic group. We have considered recognizing these three groups in the V. pompona complex as different species, but the topology of the ITS tree, with the Mexican samples forming a monophyletic, derived group, and the other two geographic groups being rather paraphyletic, and the lack of constant, morphological traits have subspecies pompona , pittieri , and are here recognized. 13a. Vanilla pompona Schiede subspecies pompona COMMON NAMES : “vainilla platanillo”, “platanillo”, “vainilla pompona”, “pltano”, “vainilla cimarrona”, “bania” (oaxaca), “nuguyu” or “nejuyu” (Oaxaca, ?Popoluca); “vainilla gruesa” (Guerrero and S Oaxaca).

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla337 Hemiepiphytic, massive vine, little branched, leafy, up to 16 m high. Stems terete, smooth, pale green dotted with whitish, 10-24 mm thick; internodes 11-14.5 cm long. Aerial, free roots thick; terrestrial roots pilose, 4 the lower surface, whitish-greenish. Leaves subsessile; blade variously shaped, ovate, widely elliptic, oblongfalcate, base widely rounded to subcordate, apex acute green, 22-29 x 8-13.7 cm, 3 mm thick. Bracts widely ovate, obtuse, rounded, concave to cymbiform, apex somewhat conduplicate, ca. 8 x 10 mm. Flowers successive, 1-2 open at once, ephemeral, tubular or with spreading segments, 58-83 (69.679.96) mm long; 7.5-10 cm in diameter; pale yellow, lip orange yellow, very showy; fragrance very strong, spicy, mint-like. arcuate, oval-subtrigonous in cross section, very slightly grooved on one face, smooth, green, 37-45 (42.292.71) mm long, 5-7 (5.850.78) mm wide, 3-4 (3.290.36) mm thick. Dorsal sepal long oblanceolate to spatulate, apex obtuse, rounded, subcalyptrate and minutely warty on the apex of the outer surface; base long unguiculate, long attenuate, canaliculate, the claw ca. 17-30 mm long, 8 mm wide in the broadest part, blade concave to somewhat conduplicate, 1116.5 (14.061.73) mm wide, ca. 14 veined, smooth, Lateral sepals long oblanceolate, oblique, apex obtuse, rounded, subcalyptrate and minutely warty on the apex of the outer surface; base long unguiculate, long attenuate, strongly canaliculate, the claw ca. 28 mm long, 9 mm wide in the broadest part, blade concave, 1316 (14.871.13) mm wide, ca. 13-veined, smooth, thick Petals oblanceolate-oblong, arcuate, slightly sigmoid and oblique, the upper margin more straight, apex truncaterounded, base attenuate, basally slightly canaliculate, keel on the lower surface, ending in a triangular, acute, at the apex of the outer surface, ca. 12-veined, 59-75 (67.835.58) x 11-17 (14.252.15) mm. Lip attached to the column along the margins of the basal half 3040 mm (35,40,30,37,39) tubular, trumped-shaped, cymbiform, deepest near the middle, axially grooved on the lower surface; when spread out 60-80 (718.52) x 29-40 (35.84.26) mm; unguiculate, the claw canaliculate, with rows of trichomes, 22-30 (24.22.92) x very obscurely trilobed, margin undulate, ca. 40 veined, apex deeply emarginate-bilobed; lateral lobes long triangular-ovate, oblique, rounded, 30-39 (35.23.48) x 10-15 (12.62.06) mm; midlobe inconspicuous, the 6-10 (82) mm x 13-22 (173.4) mm; penicillate callus at 32-46 mm from the lip base, made up by ca. 10-14 congested, retrorse, approximately trapezoidal, laciniate scales, the scales regularly united each to other along the lateral margins, ca. 5.5-7 x 5-5.5 mm, continuous with 3 thick keels that soon become fused in a cushion like, claviform, rounded, rugose callus ending at the very apex; 18-23 (20.61.85) x 3.5-4.5 mm. Column elongate, conspicuously sigmoid, 43-59 (535.54) mm long, 3.2 mm wide; ventral surface lanuginose on the apical half, below the stigma; apex dilated, 4.5-6.5 mm margin acute, uncinate, ca. 2 x 4.5 mm. Stigma trilobed, the lobes emergent, with a convex rostelum, ca. 3 x 4.5; lateral lobes divergent each to other. Fruit variable, usually arcuate, trigonous, thick, green, blackish when rippen and strongly aromatic, 7.3-15 cm long, ca. 14 mm thick (n = 6). Fig. 13. DISTRIBUTION : Mexi co (Nayarit, Jalisco, Colima, Michoacn, Guerrero, Oaxaca, Veracruz). Specimens cited as V. pompona from Guatemala and Belize, and most from Honduras have turned out to be other taxa. Vanilla pompona has a disjunct distribution. The type came from Veracruz, Mexico and this group of populations extends into N Oaxaca until the Tehuantepec Istmus, but not reaching Chiapas. Subspecies pompona occurs in W Mexico (Nayarit to S Oaxaca), where spread widely are very common. In the ITS tree all the samples of V. pompona from form a nested subclade with high bootstrap support (97%). Another clade within V. pompona includes material cultivated in Guadaloupe, and from Suriname and Guyanas, whose bootstrap value is moderate (70%). The rest of the samples of the complex have unresolved relationships at the base of the V. pompona clade, which has a 100% bootstrap value and is sister to V.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.338 LANKESTERIANA FIGURE 13. Vanilla pompona Schiede subsp. pompona. Based on M. Soto 7746. Drawing by M. Soto.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla339 calyculata, a clearly different species. The distributiona, molecular, and morphological data suggest that subsp . pompona may be the result of a relatively recent colonization of Mexico through a rather long-dispersal event from Central American plants. We believe that we can recognize at least two taxa in the complex: V. pompona subsp pompona and V. pompona subsp. (Lindl). Soto Arenas. ECOLOGY : from sea level up to 1200 m, more common at 300-900 m; in several types of tropical forest (deciduous, subdeciduous, evergreen, lower mountain, gallery forests, in savannas or warm pine-oak forest, dry areas; this species is usually absent from the wet forests, but occasionally can be located there in limestone outcrops. Flowering period from April to early June. Vanilla po mpona was described from sterile specimens collected by Schiede near Papantla and Colipa, in northern Veracruz, about 1829. The big plants, of Mexican plants. The large, deep yellow, and strongly segments (rarely in western Mexican populations); the short, thick fruit is strongly aromatic and very much appreciated by peasants, and also by vanilla conoisseurs. The odor of the cured fruits is typically heliotropin-like in the V. planifolia plantations, as a curiosity or charm, where it is known as “vanilla gruesa”, is very common in the coffee plantations, protected, but used only for local consumption. Vanilla pompona subsp. pompona is somewhat and the perianth segments apparently never spreading; vegetative features probablly represent a plastic some specimens of southern Oaxaca approach those of are found in Nayarit and Jalisco.O T HE R R E CORDS : MEXICO : NAYARIT : Camino de Ruiz a Jess Mara, ca. 200 m altitud, cafetal derivado de selva mediana subdecidua, abril 1986, 2009, 2010, 2011, 2013, 2017 AMO! El Caimn, 100 m sobre la brecha hacia El Naranjo, que sale del km 10 del Quercus cf. conspersa , Byrsonima crassifolia , Bursera acuminata , 490 m s.n.m., 21 26’ 22.2’’ y 105 06’09.6’’. Nos gui el Sr. Benito Decena Jimnez, de El Cora. 26/ junio/1998. M. Soto 8613 y E. Huerta AMO! M. Soto 8617 y E. Huerta AMO(x4)! JALISCO : La Playa (oeste de Los Llanitos), Mpio de Puerto Vallarta, bosque subdeciduo, R. Gonzlez Tamayo 884 *AMO! *LL! Mpio. Puerto Vallarta, entre Los Llanitos y Los Almacenes, Gonzlez Tamayo s.n. dibujo! Mpio. Cabo Corrientes. above El Tuito, road to Mina de Cuale, R. McVaugh 26397 MICHOACAN : Mpio. Aquila, San Pedro, Coalcomn, 550 m, 6-19-39, “Vanilla”, forest, vine, G.B. Hinton 13815 *AMES(57796)! NY! *US(1805771)! Aquila, Tizupan, Coalcomn, 4-8-41, “Vanilla”, forest, G.B. Hinton 15910 *AMES(60655)! NY! GUERRERO : Vallecitos, Montes de Oca, 9-13-1937, “vanilla”, on a tree, G.B. Hinton et al. 11387 K! *US(2020421)! Sierra Madre, GuerreroMichoacn, 400Endroits ombrags,”Vainilla”, E. Langlass 941 *AMES(71358)! G(x2)! K! *US(386288)!. OAXACA Madre, near Copalita and below Pluma Hidalgo, climbing on tree trunks, 26 Jul 1936, 500-1000 m, Gonzlez sub E. Oestlund 6102 *AMES(51840)! km 190 Oaxaca-Pto. Angel. 14 mayo 1990. Flor prensada de material en lquido O. Surez 1153 AMO(15906; also in spirit)! Sanguijuela, Mpio. Juquila, 1200 m s.n.m., 21 sept. 1992, AMO(only fruit)! ca. San con encinar seco en las lomas. ca. 600 m s.n.m. 17 de abril de 1992. M. Soto 6607, N. Prez y E. Martnez AMO! Cerca de San Gabriel Mixtepec, cafetal en la zona de selva mediana subcaducifolia y encinar caliente, 800 m s.n.m. 11 mayo M. Soto 7632, A. AMO! OAXACA (specimens from the Gulf slope): Consoquilla, /42, Liebmann 288 W(11761, drawing)! 1859, Cuming s.n. fruit very fragrant, 9501’ 170’, 100 m, 18 Feb 1935, O. Nagel sub E. Oestlund 4574 *AMES(51841)! F(1257235)! climbing on trees, 9501’ 170’, 100 m, 20 Apr 1935, H. Knape sub Oestlund 4781 *AMES(51842)! F(1257236)! *MO(1145435)! *SEL(011366)! *US(1809788)! San Juan Guichicovi, “vainilla”, hierba, pachine seco, en la orilla del camino, uso, con la fruta machacada, VIII-1986, Nereyda MEXU(431242)! ! Mpio. Sta. Mara Chimalapa: Arroyo Huahuagtz, ca. 5.5 km al E

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.340 LANKESTERIANA de Sta. Mara; lat. 1655’ N, Long. 9439’ O, alt. 250 m; acahual en rea plana en orilla de arroyo (anteriormente selva); suelo negro. Bejuco; fruto verde, carne amarillaverdosa con semillas negras; comn; n.v. “bainia”, “nuguyu” (en zoque); se usa en forma de aceite para el cabello para que de ms olor, 5 septiembre 1984, H. Hernndez G. 396 *AMO(8094, fruit)! CHAPA! Mpio. Sta. Ma. Chimalapa: Arroyo Rancho Viejo, ca. 2-4 km al E de Sta. Mara; lat (rodeada por encinar) con Poulsenia , Ficus , Brosimum etc., (en castellano), “nejuyu” (zoque); frecuente; se usa para darle fragancia al cabello (se seca primero), 6 abril 1987, H. Hernndez 2418 , AMO(16964)! 4 km al W de la frontera Oaxaca-Chiapas, carretera a Tapanatepec, Dto. Juchitn, rocoso, 27 Marzo 1984 [botones], Martnez MEXU! NY! Mpio. Santa Mara Guienagati, Loc. Dto. Tehuantepec. 3 km al N de Santa Mara Guienagati, carretera a Guevea de Humboldt. 1643’N, 9522’W; elev. frecuente, 27 junio 1991, A. Campos 3721 MEXU(578082)! ca. Valle Nacional, 1996, J. Prez Mesa s.n. AMO(in spirit)! Terrenos comunales de San Agustn, Mpio. Sta. Ma. Jacatepec; a unos 2 km al W de San Agustn, selva hmeda con Manilkara , Brosimum , Pseudolmedia , con cafetal, a 350 m s.n.m. Planta silvestre de ca.18 m de alto, protegida, con M. Soto 8495, M. Hernndez, F. Snchez y M. Tensohua AMO! trees, 12 feb 1932, 9652 and 1912’ ca. 900 m, O. Nagel sub E. Oestlund 2684 *AMES(41477, sterile)! From Mexico. Secured by Prof. P.H. Rolfs, in charge of the Subtropical Laboratory, Miami, Fla., while traveling in April, May, and June, 1905. This species is a very stronggrowing vanilla. Produces the largest fruits and in considerably quantity. Secured near Papantla, Person interested: David Fairchild. P.I. No. 14440 SEL(011358; photo)! Valle de Crdoba, 20 april 1865-6, Bourgeau 2332 m, Jun 1935, C.A. Purpus sub E. Oestlund 4852 *AMES(51839)! H. de la Higuera, 510 m s.n.m., vegetacin riparia, bejuco perenne, abundancia regular, fruto aplanado [sic], septiembre 8, 1966, M. Sousa 2721 *AMO(3443, fruit)! MEXU(277445, 296430, 277427; x3)! MO(4273614)! perennifolia primaria, perenne, “litsmoya”, colectada en fruto. Usos: seca se pone en vaselina y se unta en el pelo, septiembre 30, 1967, M. Sousa 3276 *AMO(8592, fruit)! MEXU(fruit, 284554)! a 1 km al NE de Soteapan, 400 m [estril], M. Sousa 3272 MEXU(284553)! “vainilla”, La Ceiba, Municipio de Puente Nacional, bosque de rboles F. Ventura 12355 *AMO(785, fruit)! MEXU(fruit, 235724)! Barranca de Monterrey, entre el manantial y la cueva del abono, Mpio. Axocuapan, alt. 380 m s.n.m., selva mediana subperennifolia, bejuco, fruto verde, nom. vul. “vainilla cimarrona”, trepador, 02-01-1987, *AMO(9026, fruit)! XAL(fruit)! El Remudadero, por la carretera Conejos-Totutla, Mpio. Paso de Ovejas, (1917’N, 9633’ W), selva mediana subcaducifolia primaria, suelo arcilloso, pedregoso, en la barranca, bejuco perenne, 10 m, escasa, fruto verde, nom. vulgar “platanillo”, fruto comestible, bejuco trepador, 01-11-1982, G. Castillo C. 2747 Mpio. de Atoyac, 1857’N, 9649’W, alt. 900 m s.n.m., selva mediana subperennifolia, primaria, suelo crstico, 18-05-1985, *XAL! [cf., fruit] Lmites de Veracruz con Tabasco, Mpio. Coatzacoalcos, 1803’N, 9606’W, selva baja inundable, primaria, suelo negro pantanoso, hierba trepadora, perenne, 6 m, fruto verde en vainas, nom. vulgar “vainilla”, 18-05-1980, J.I. Calzada 6096 F(1975502)! *XAL(fruit)! Alrededores de San Fernando, Mpio. Soteapan, 1818’N, 9453’W, alt. 720 m s.n.m., pinar-encinar, primaria, suelo rojo arcilloso, bejuco perenne, 1 m, escaso, fruto verde, 18-12-1978, R. Ortega et al. 1021 Palmillas, Mpio. Puente Nacional, 1913’N, 9646’W, 600 m s.n.m., selva baja caducifolia, primaria, suelo delgado, arcilloso, pedregoso, caf, con poca materia orgnica, bejuco perenne, 4 m, fruto verde, 13-03-1985, M.E. Medina 4284 N. Demneghi G. 558 *XAL(fruit)! Mecayapan, regin de Los Tuxtlas; acahual derivado de encinar-pinar, 13/01/1998, S. Cruz Ramrez sub M. Soto 8777 ca.750 m s.n.m. Acahual de selva mediana perennifolia submontana muy hmeda. Protegida o cultivada. Planta muy vigorosa, tallos 13-26 mm de dimetro, hojas hasta de 30 x ptalos verdes exteriormente, parte interna amarillo-verde limn, labelo amarillo yema cocida con algunas lneas ocre, garganta profunda con la parte apical amaillo-naranja, aroma especioso, hierbabuena y algo ms. 29-IV-1995. M. *AMO! El Angostillo, 34 km de Huatusco, ca. 14 km de Manuel Gonzlez, Congregacin El baja caducifolia en barrancas baslticas; aroma fuerte a mentol, visitada por Eulaema en la casa de Julio Arguijo. 30-IV-1995. M. Soto 7747, M.A. Flores, J. Arguijo, D. Demneghi, M. Demneghi * AMO(x2)!REFERENCES: McVaugh, Fl. Novo-Galiciana. 16: Orchidaceae: 351-353, , 1985. 13b. Vanilla pompona Schiede subsp.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla341(Lindl.) Soto Arenas, comb. et stat. nov. BASIONYM: Lindl.,Gen. Sp. Orchid. Pl. 435. (1840). TYPE: Guyana Francesa, Martin s.n. (holotype, K-L!). Illustr. Dunsterville y Garay Venez. Orchid. Ill. 1: 435 (1050, as V. pompona). N.v.: Palanda vainilla, Ecuador: vainilln. Vanilla claviculata Duss non Sw.; V. lutescens Moq. ex Dupuis COMMON NAMES: “vainilln” (French and British Antilles), DISTRIBUTION: Apparently naturalized in the West Indies. There is no adequate material from NW South America, but the specimens from the Amazon basin, Guyanas, Coastal Venezuela, and S Brazil are considered here to belong to. It is unknown if the plants from W Ecuador are V. pompona or V. grandifolia. ECOLOGY (2006) report the pollination of V. (pompona subsp.) by male Eulaema meriana, one of the largest of the euglossine bees. They also photographed a male Eulaema cingulata gathering scents from the mature fruit of this species. There has been at least one other report of euglossine bees brushing on Vanilla fruits (Madison, 1981). It is quite possible that the Euglossini may play a role in the dispersal of Vanilla Vanilla pompona is very similar to Lindl. (“French Guiana: without precise locality, Martin s.n., K-L!), and the latter is has been recognized as a synonym by most orchid taxonomists. However, V. is different because it has a rather oblong V. pompona. In the lip > 80 mm long, the callus 9-10 mm long and originated at ca. 60 cm from the base and the column is 60-75 mm long; also the apical thickening is less conspicuous in usually spreading segments, while V. pompona is usually closed, although occasionally, specimens with spreading segments on sunny days are found, as the 13. Vanilla lutescens Moq.-Tand. described from material of Paraguay [M. Coudert, cultivated at the Facult de Mdecine de Paris, the type, not seen; probable duplicate at W(35455)!] is also similar to V. REFERENCES Les orchides de Guyane Franaise: 138, photo p. 93. 13c. Vanilla pompona Schiede subspecies pittieri (Schltr.) Dressler, comb. et stat nov. BASIONYM: Vanilla pittieri Schltr., Fedde Rep. 3: 106. 1906. TYPE: Costa Rica: In der Wldern an Ufern des Ro Ceibo bei Buenos Aires, c. 200 m. M-H. Pittier no. 6600, blhend im Januar 1890, B(destroyed); AMES(24329; drawing of the holotype)! BR! DISTRIBUTION : Honduras, Nicaragua, W Costa Rica The available Costa Rican material is not very well-preserved, but seems to be more similar to name V. pittieri Schltr. is available to these Costa Rican populations. The type of V. pittieri was lost in Berlin bombing; a sketch of the type made under R. Schlechter’s supervision, and a rather crude shown as entire, with a very elliptic blade, as it had been prepared from an immature bud. There are additional specimens from the area of Buenos Aires, Puntarenas, that match our concept of V. pompona. OT HE R R E CORDS : NICARAGUA : CHONTALES : ca. 2.8 km above (N of) Cuapa; ca. 1217’N, 8523’W, elev. 400-500 m; roadside, pastures, disturbed evergreen forest on hillside, and bank of small stream. One seen, climbing up trunk of large tree in disturbed forst near road, section of lower stem collected, W.D. Stevens, B.A. Krukoff 3690 *SEL(049330; sterile)! SEGOVIA : Oersted W(19359)! Plant Large thick (up to 2 cm) climbing leafy stems. FlowersLarge light yellow sepals and petals, yellow-orange lip. Light yellow column, orange on anterior face [mounted with description, notes an analytical drawing] A.H. Heller s.n. *SEL(003850)! NUEVA SEGOVIA : Quilal, ca. 1334?N, 8601’W, elev. 430 m, cultivated, said to have been collected wild in the aromatic, 16 March 1980, W.D. Stevens, B.A. Krukoff 16829

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.342 LANKESTERIANA *SEL(049331)! : Cerro Livico, 7 km northeast of Siuna, forest slope; elev. 500 m. Climbing trunk of tree 28 April 1978, D. Neill 3670 *SEL(047695; sterile)! COSTA RICA : GUANACASTE : Flores amarillas. Enredadera subiendo tronco de rbol, en chaparral o bosque poco denso dentro de sabana cerrada. A 5 millas al sur de La Cruz, Liberia. Alt. 200 m, febrero 11, 1963, A. Jimnez M. 313 *F(1606969)! 550 m, Calvo 1136 F(1456238)! Parque Nacional Guanacaste, Estacin Maritza, sendero a la cima del Volcn Oros. Bosque primario y secundario, 1057.6’N, 8529.6’W, 600 m. Planta de una quebrada. Frutos verdes. 2 julio 1989, INBio 132 INB! *CR(fruit)! PUNTARENAS : Cantn de Buenos Aires. Along the Ro Ceibo, Ujarrs. 091400 N, 831800W, 500 m. Thick stemmed vine; climbing high on trunk of tree M. Grayum 10237 INB! Esparza Macacona, margen derecha, quebrada G. Herrera MO(3594340)! WITHOUT PRECISE LOCALITY : “ Vanilla reichenbachii ”, Endres 270 W(16231, 16175, 16176)! Sepals and petals greenish yellow. Lip orange yellow. Pie del Turrubares, 150 m, March 6, 1926 A. Alfaro 269 *AMES(31487, *31487)! In forest between Colonia *US(474659)! “Costa Rica”, dried leaf from M. Godefroy-L. , Paris, 1895 K! PANAMA : PANAMA : Low places contiguous to City, Alt Sea level. Grows in damp places. Climbs trees up to 40-60. Flowers yellow, with orange throat. Flowers FebMarch. 1919, C.W. Powell 137 AMES(*23763; AMES 28290 is V. phaeantha )! Moist woods; large herbaceous vine, Ro Paraso, above East Paraso, January 7, 1924, P.C. Standley 29919 *AMES(31437, sterile)! *US(1225393)! Vine on west shore, Barro Colorado Island, March 11, 1931, C.L. Wilson 119 *F(636191)! Barro Colorado Island, shores of Gatum Lake, South of Lab., Aug 28th 1929, W.N. Bangham 458 * C.V. Piper s.n. *AMES(36948)! Low thick scrub along R. Tecumen, north of Chepo road, up to 30 m. March 10, 1935, A.A. Hunter and P.H. Allen 852 *AMES(42162)! *F(1329667)! *US(1976167)! Coastal thicket; large herbaceous vine; common, Punta Paitilla, January 12, 1924, P.C. Standley 30788 *US(1229496)! Parque Nacional Altos de Campana. este del campamento de los guardabosques de INRENARE. Bosque hmedo tropical premontano. Elevacin entre 800 y 900 m. 840’N; 7955’W. 23 de abril de 1993, M.D. Correa, PMA! Cerca de Cerro Azul, 14.8 km al N de la carretera Cd. de Panam-To cumen, camino Cerro Azul-Cerro Jefe, vegetacin secundaria con Terminalia y Byrsonima , ca. 600 m s.n.m., trepadora, estril, toda la poblacin con hojas elpticas, grandes y tallos gruesos, 4 junio 2001, M. Soto 9921, G. Salazar, J. Linares y R. Gregg PMA! in cult., at Smithsonian Tropical Research Institute in Balboa. Voucher: fragrance collection. 18 Mar 1979, J.D. Ackerman 1354 SEL(039338)! In Powell’s garden; large vine, Balboa, Non., 1923-Jan., 1924, P.C. Standley 28570 *US(1225365; sterile)! COCLE : ca. 5 km al N de El Cop, entrada del Parque Nacional Omar Torrijos, vegetacin secundaria abierta, derivada de bosque hmedo tropical premontano con Vochysia , ca. 700 m s.n.m. 2 junio 2001 , G. Salazar J. Linares, M. Soto y R. Gregg PMA! REFERENCES : Hamer, Orch. Nicaragua, Ic. Pl. Trop. pl. 1195. 1984.14. Vanilla sarapiquensis Soto Arenas, sp. nov. TYPE: COSTA RICA: HEREDIA: Sarapiqu. Llanura de San Carlos. Sardinal. Bosques residuales y tacotales camino a Finca Dos Loas. 10’’N, 84’’, 70 m. Bejuco herbceo. Spalos verde crema, labelo blanco, columna con la punta negra, 10 May 1995, holo. J.F. Morales 4082, E. Lpiz, INB! iso. apparently at MO. Vanillae costaricense aemulens differt longioribus, labello suborbiculari, 3-lobato, callo duobus carinis incrassatis prominentibus in apicem Hemiepiphytic vine, leafy. Stems 4 mm thick; internodes 6.5-7.8 cm long. Aerial roots ca. 2 mm wide. Leaves obscurely petiolate, the petiole canaliculate, ca. 7 mm long, 3.5 mm wide; blade elliptic, long acuminate, somewhat cuneate at base, membranaceous-chartaceous in dried condition, 15.9-19.5 x 4.8-7 cm. very different cm long, internodes up to 1.5 cm long; peduncle ca. 26 mm, rachis ca. 2 mm thick. Bracts scale-like, ovate to lanceolate, acuminate, concave, 9 x 4 mm. Flowers successive, with rather spreading segments, showy, ca. 7 cm in diameter; tepals green-cream colored, lip white; of heavy substance. rather sigmoid, terete, somewhat thickened at base, smooth, inconspicuously sulcate, 42-53 mm long, 3-4 mm thick. Dorsal sepal rather spreading, convex, the undulate; elliptic, apex acute-subacute, extreme apex

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla343 FIGURE 14. Vanilla sarapiquensis Soto Arenas, based on the type specimen, J.F. Morales 4082, Drawing by M. Lpez.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.344 LANKESTERIANAconduplicate-subcalyptrate, rounded; base obtuserounded, ca. 10 veined, sparsely warty on the dorsal surface, 44 x 17 mm. Lateral sepals basally concave, the apex recurved, widely elliptic, obtuse, base obtuserounded, margins scarcely undulate, sparsely warty on the dorsal surface, 11-veined, 41 x 20 mm. Petals spreading, recurved at apex, slightly undulate on the upper margin, oblong-elliptic, oblique, subacute, base obtuse-rounded, 8-veined, smooth but axially sulcate on dorsal surface, 45 x 16 mm. Lip attached to the column 3 mm, ovate-suborbicular, obscurely trilobed in outline, constricted at 2/3; 32 x 28 mm, the base subcordate, slightly concave, with the lateral margins erect forming a throat around the column; midlobe 8 x 16 mm; the lateral lobes semielliptic, the margins minutely pleated, 21 x 10 mm; callus arising above the middle, builted up by a pair of broad, elevated apically botton to the crest, more elevated at distal part, widely triangular in cross-section on their distal part, 18 mm long, 5 mm wide. Column arcuate, semiterete, with a dilated apex, smooth, 21 x 4 mm. Anther protruding, semierect the distal part perpendicular to the column body; the anther separated from the surface of the lip. Fig. 14. DISTRIBUTION : Known only from northern Costa Rica, in the Llanura de San Carlos; it is to be expected from adjacent Nicaragua. ECOLOGY : Apparently in rainforest, at 70 m elevation. This species of the membranaceous group is known from a single collection from N Costa Rica. Vanilla costaricensis, is known from the same geographic area (Llanura de San Carlos), but V. sarapiquensis is like bracts (V. costaricensis has big bracts like those of V. inodora tepals, suborbicular, trilobed lip, and especially by the very different callus made up of two elevated, V. costaricensis with the apex truncate or somewhat trilobed, but the lateral lobes at the apex are subequal or slightly longer than the mid lobe. In V. costaricensis the callus line, from the base to the begining of the apical third, then separated in 3 low, erect, congested keels ending at the apex. Vanilla mexicana has a distinctly trilobed lip, with the midlobe more protruding, 3-keeled, with the keels parallel. Also related is V. martinezii with much larger is similar.15. Vanilla trigonocarpa Hoehne, Arq. Bot. Estado Sao Paulo, nov. ser. 1(6): 126, t. 139. 1944. TYPE: BRAZIL:PARA 7-10-1926, A. Ducke s.n.. holo., HEPF(19445)! Dressler, Orquideologa 13(3): 229-232. pl. 1979. TYPE: PANAMA: Prov. Panam, carretera El Llano-Cart, 10-15 km al norte de El Llano; 3 marzo 1976; spalos verde plido, ptalos crema verdoso, labelo blanco, lmina cafamarillo, fragante, R.L. Dressler 5290, holo. US(not seen). Hemiepiphytic vine, leafy, up to at least 8 m high. Stems 4.8-7.5 cm long. Adventitious attaching roots dorsiventrally compressed, at least up to 6 cm long, ca. 4 mm wide; adventitious free roots subterete, ca. 3 cm long, 2 mm wide. Leaves petiolate, the petiole 1.5-2.5 cm long; blade elliptic or elliptic-oblong to obovate, long acuminate, leathery, relatively thin, 1530 x 5.5-7 cm. usually on the lower long. Bracts lanceolate, 15-20 mm long. Flowers with spreading segments, sepals pale green, petals greenish cream, lip cream, the blade yellow-brown with a white margin, the throat with yellow-brown veins. ca. 4 cm long. Dorsal sepal lanceolate, obtuse, subclawed, the claw ca. 40 mm long, 8 mm wide, 11-11.5 x 1.6-1.8 cm. Lateral sepals elliptic, subclawed, the claw ca. 40 mm long, 8 mm wide; 10.5-11 x 1.8-2 cm. Petals narrowly oblanceolate, obtuse, with a prominent rib on the outer surface, 10.8-11 x 1.6-1.8 cm. Lip attached to the column along the margins of the basal half (ca. 6.5x 5.5 cm; unguiculate,

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla345 FIGURE 15. Vanilla trigonocarpa Hoehne. Based on McPherson 11286 Dressler 4750 (fruit), Dressler s.n. published picture. Drawing by M. Lpez and M. Soto.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.346 LANKESTERIANAclaw ca. 4 cm, then expanding abruptely and somewhat penicillate callus made up by several laciniate scales, 5 x 5 mm; with a verruculose ridge running toward the apex; lip apex strongly veined. Column elongate, ca. 80 mm long, slightly arched, ventral surface pilose toward the apex (for ca. 3 cm); with small wings. Stigma trilobed, the lobes emergent, the mid-lobe the largest. Anther ventral, 6 x 4 mm. Fruit cylindrical, 20-23 cm long, ca. 1.8 cm wide; somewhat triangular in cross-section, black and shiny, ca. 0.6 mm long. Fig. 15. DISTRIBUTION : Costa Rica, Panama, and Brazil, and probably in the intermediate areas. ECOLOGY : From very wet rainforest, from 120 to 350 m altitude; it can be common in certain areas. Flowers in Central America in January-February, and July; perhaps all year. Hoehne (1945) indicates that the fruits is pleasant. Dressler (pers. com.) has observed males of Euglossa asarophora pollinating this species in Panama. This species has been known in recent years in Central America as Dressler, but the Central American specimens are apparently by Hoehne as V. trigonocarpa. Vanilla trigonocarpa is somewhat variable in the form of the apical lobe of the lip, deeply retuse in the type and truncate to slightly emarginate in the Central American specimens. It is also variable in the development of the lacerate margins. The type specimen is minutely lacerate, as it is the specimen T. Antonio 3645, however, other specimens are deeply lacerate, as G. McPherson 11286. Vanilla trigonocarpa is distinguished by the very characteristic. It is probably related to V. espondae (and perhaps to V. sprucei the same color, but with shorter and broader segments, and the lip with an oblong-rounded midlobe covered with digitiform papillae, up to 3.5 mm long. Molecular analyses show that V. trigonocarpa is the most basal species among the American penicillate Vanillas. OTHER RECORDS : COSTA RICA : PUNTARENAS: at base of Osa Pennsula, ca. 25 km E of Rincn along newly improved road; elev 350 m, climbing on tree in mature rainforest, 4 Jan 1986, AMES! PANAMA: PANAMA: El Llano-Cart road, 10.1 miles from highway, on trail to west; 9’N, 79’W; c. 350 m. Epiphytic on trunk; perianth green-white, the frilly portion of lip brown, 16 July 1987, G. McPherson 11286 *MO(3499115)! about 10 km north of El Llano, 29 January petals greenish cream, lip cream, streaked with brownyellow within, center of limb brown-yellow, R.L. Dressler 4571 (PMA, SEL); El Llano-Cart highway, 10-12 km north of El Llano, 12 September 1974; climbing vine, about 8 m, R.L. Dressler 4750 PMA, SEL *US(2952908)! COCLE: On the Atlantic side, ca. 5 hr walk from sawmill at El Cop. Along slopes above Norte Ro Blanco near small village of 2 Feb 1980, T. Antonio 3645 *MO(2928700)! WITHOUT LOCALITY: leg.: Collector unknown. Selby Accession: 89-0281. Growing in Selby Display House. 26 April 1991. Coll. *SEL(065176)!R EFERENCES : Hoehne, Fl. Bras. 13(2): 28, t. 20. 1945.Excluded speciesVanilla barbellata Rchb.f., Flora 48: 274. 1865. A horticultural specimen grown in Austria, supposedly from Cahuita, Costa Rica, seems to be the West Indian V. barbellata. There is a plate prepared by the artists of the Royal Botanical Expedition to New Spain, circa 1800 (Hunt Library; tracing at Library of MEXU (as 12 Vanilla mexicana”), G, and tracings probably elsewhere) that represents this species. Mexican origin of the plant is doubtful, since there are Mesoamerica. Many other paintings labeled as from Mexico (“de Mxico”) are actually Cuban plants, prepared during the stay of the Expedition at the island. As far as we know, Vanilla barbellata has been found only in Bahamas, Florida, Cuba, Puerto Rico, and theVirgin Islands.Vanilla mexicana Miller, Gard. Dict. ed. 8: n. 1. 1768. TYPE: Haiti, Plumier s.n., the holotype is Plumiers original drawing at P; reproduction Plumier, Nov.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla347 FIGURE 16. A — Vanilla cribbiana. B — Vanilla dressleri. C — Vanilla hartii. D — Vanilla phaeantha. E, F — Vanilla planifolia .

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.348 LANKESTERIANA FIGURE 17. A — Vanilla calyculata. B — Vanilla guianensis . C — Vanilla insignis. D — Vanilla phaeantha .

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla349Pl. Amer. Genera 25, pl. 28, 1703! Epidendrum vanilla L., Sp. Pl. ed. 1, 2: 952. 1753. Lectotype (icon designated by Cribb, Taxon: 48(1): 47. 1999): “Volubilis siliquosa plantaginis folio” in Catesby, Nat. Hist. Carolina 2 (3): app. t. 7. 1747. V. vanilla Porto Rico 5: 185. 1924. Syn: V. aromatica L. (based on the same specimen). V. aromatica Sw., Nov. Act. Soc. Sc. Upsal. 6: 66. 1799. TYPE: Haiti, Plumier s.n., the holotype is the original drawing by Plumier. V. anaromatica Griseb. Fl. Brit. W. Ind. 638. (?) V. epidendrum Mirbel, Hist. Pl. ed. II. 9: 249. 1802-6. V. vanilla Karst. Deutsch Fl. ed. 2, 1: 474. 1895. V. vanilla Huth, Helios 11: 136. 1893. Vanilla mexicana has been repeatedly cited from Central America because the prevailing confusion between this species and Vanilla inodora. In spite of its name, it is a plant from the West Indies and northeastern with apple-green tepals and white, distinctly trilobed, 3-keeled lip. Other specimens labeled as V. mexicana from Costa Rica belong to the new species here proposed as V. costaricensis. Vanilla planifolia Jackson x V. pompona Schiede x V. planifolia Jackson It was imported to Costa Rica, where is rather common, and it is said that it is very resistant to root rot disease (David Gardella, com. pers.). ITS sequences are polymorphic and similar to both V. pompona subsp. and to V. planifolia. MatK and rbcL sequences are grouped with V. planifolia sequences, which suggest that V. planifolia was used as the maternal progenitor, since the plastid genome is inherited maternally. Vanilla x tahitensis J.W.Moore, Bishop Mus. Bull. Bot. 102: 25. 1933. TYPE: Flora of the Society Islands, Island of Raiatea, altitude 20 m; on trees, escaped from cultivation; 3rd valley south of Faaroa Bay, 2-3 dm high. Native name Vanira tahiti, Nov. 11, 1923, J.W. Moore 294, holo. Bernice P. Bishop Museum Herbarium (559503)! V. hirsuta Lasianthera 1(1): 47. 1996 TYPE: Papua New Guinea, M.A. Clements 6742 CANB(holo.) iso(NCBG) not seen, photo ! A DDITIONAL SPECIMENS : FRENCH POLYNESIA: Raiatea: H. St. Johns 17310 K! Mongareva Is. H. St. Johns 14459 K(drawing)! Vanilla tiarei 1915 161: 202, nomem TYPE: Tahiti. DISTRIBUTION : Known from French Polynesia (Society Islands), and New Guinea. The Tahitian vanilla certainly is not known in Central America. Vanilla tah itiensis J.W. Moore was described from material from Raiatea, Society Islands. The type corresponds to the plant widely cultivated in French Polynesia and known as “Vanille de Tahiti”, and the second most important source of commercial vanilla. The same taxon was also recently redescribed by Clements V. hirsuta , based on a specimen collected in Papua New Guinea ( M.A. Clements 6742 CANB(holotype), NCBG(isotype) not seen, photos!). There are also other different plants in Tahiti, including V. planifolia , and some variants of V. tahitiensis that may represent different cultivars or different species (e.g. a plant labeled as cv. Tahita is apparently V. bahiana ). Porteres (1954) indicated that the origin of V. tahitiensis was very enigmatic. It has been suggested also that the distribution of V. tahitiensis is far from other vanillas, that its occurrence in Tahiti is perhaps due to a human introduction, and that it is very similar to V. planifolia and not to the species from New Guinea, which is the area with native vanillas closest to Tahiti, such as V. giulianettii F.M. Bayley, V. kaniensis Schltr., V. kempteriana Schltr., (J.J. Sm.) J.J. Sm., V. wariensis Schltr. Except for V. tahitiensis , all the Vanilla species with aromatic fruits are restricted to the Neotropics (Soto, 2003). Occasional reports of fragrant fruits in Asia (e.g. Heyne, 1922) are probably based in wrong observations, or in odors unlike the typical vanilla fragrance. Vanilla

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.350 LANKESTERIANA disjunctions are better explained by ancient vicariant events than by long distance dispersal (Soto et al., in prep). Vanilla (in the generic sense) was introduced to Tahiti from Manila about 1848. Ptard (1986: 123) claimed that Admiral Hamelin was responsible for bringing the plants to the Botanic Garden at Papeete, whereas Herman et al. (1989: 20) asserted that William Ellis, a missionary from London Misionary Society, brought cuttings from the Philippines (Smith et al ., 1992). Ryan (1986) agrees that the origin of V. tahitiensis is uncertain, she said that it may be that some Vanilla planifolia plant stock was hybridized or that it mutated, allegedly in the Philippines and that V. tahitiensis is not more data about this statement, although she indicates Tahitian growers as the source of the information. Morphological variation in Vanilla planifolia is rather great, but it is clear that traits exhibited by V. tahitensis depart from the known variation of V. planifolia than could cause such conspicuous differences. Sexual reproduction of vanilla is almost impossible under plantation conditions, and the life span is long enough of such “novelties”. Therefore, the hybrid hypothesis is more plausible. The hybrid origin of V. tahitiensis was suspected by Portres (1954) and Ptard (1986), who suggested that it arose from a cross between V. fragrans (= V. planifolia ) and V. pompona . Vanilla tahitiensis is morphologically very similar to V. planifolia and its allies, especially V. odorata and V. insignis, but not to V. pompona . The hybrid between [ V. pompona x V. planifolia ] x V. planifolia has thicker stems, leaves, and different V. tahitensis . Compared to V. planifolia, V. tahitensis has retrorse warts on the midlobe of the lip. Since this trait is a synapomorphy of the group of V. odorata (including and V. insignis , and also V. cristagalli and V. ribeiroi from South America), if of hybrid origin one of these taxa is the candidate for parental species in addition to V. planifolia . V. helleri has a very short lip claw, and widely elliptic leaves, broader than either in V. planifolia or V. tahitensis . V. insignis has conspicuously sulcate, papillose stems, and leaves similar in shape to V. planifolia . Since V. tahitensis has slender stems, narrower leaves, longer perianth segments and column, the best candidates for a parental species are V. odorata and (if this is V. odorata ), both with slender stems, narrower leaves, longer perianth segments and column, than V. planifolia and V. tahitensis . Vanilla odorata is an important source of vanilla fruits from the wild. Most vanilla beans collected by peasants in Chiapas, Mexico are from V. odorata , since wild V. planifolia is extremely rare or absent. Ames also reported that it was the best quality species in Ecuador; and herbarium records from Bolivia indicate that “it was the best one there”. Furthermore, V. odorata has some interesting features that make it a good candidate in a Vanilla hybridization program, such as the higher tolerance to more open habitats, its origin from places with higher annual rainfall and more equatorial its resistance to the attack of “chinche roja” ( Tentecoris confusus ) (M. Soto, pers. observations); nothing is known about its resistance to fungal and bacterial diseases, but it seems not as susceptible as V. planifolia (M. Soto, pers. obs.). Vanilla planifolia and V. odorata may be sympatric in several regions of Central America. In Northern Oaxaca, Mexico, both species have been Both species are probably pollinated by Euglossine bees (apparently Euglossa viridissima in the case of V. planifolia ) involving a pollination system involving deception. Fragrance analysis Vanilla tahitiensis is the only Vanilla species from which there are reports of diploid and tetraploid chromosome counts. Tonier (1951) reported 64 somatic chromosomes for “ Vanilla haapape ”(sic), considered to be a variety of V. tahitiensis (R. Porters, 1954), while Heftimiu-Heim (1941) reported 2n=32 for V. tahiti ( sic ) and V. papeno ( sic ), the later a common name of V. tahitiensis var. haapape (Porters op. cit .). ITS sequences of a specimen of V. tahitensis cultivated in France, and another gathered from the Gene Bank as V. hirsuta are nested into one of the clades of V. odorata with 100% of bootstrap support (other specimens of V. odorata have an ITS sequence more similar to that of V. helleri ) . V. tahitensis to make a quantitative analysis of the

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla351intermediate morphology character of it between V. planifolia and V. odorata , qualitative data support the hybrid origin between these species. ITS sequences indicate an extremely close relationship with V. odorata . However, Schlter found that was no evidence of a hybrid origin analysing. In conclusion, it is suggested that V. tahitensis is a taxon of hybrid origin between V. odorata and V. planifolia introduced to Tahiti from Phillipines. Lubinsky et al. (2008) provide strong support for the hybrid origin of V. tahitensis from cross(es) of V. planifolia and V. odorata. It is not improbable that this cross was taken from Mexico to the Philippines when both countries were ruled by the Spanish crown and subject to a large commercial exchange thanks to the Manila Galleon, that connected Acapulco with the far east. The large variation observed (Maurice Wong, pers, com.) and detected by RAPDs in the tahitian vanilla is an additional point that supports its hybrid origin. Lubinsky et al. (2008) have suggested that V. xtahitensis can be an artificial hybrid and even postulated a postclassic origin. Orchid hybridization and subsequent germination of the sclerotic seeds is almost impossible under primitive conditions. Even now, Vanilla seeds are planted from immature fruits in which the testa have not become hard and it is thought that mature seeds in ripe fruits must pass through the digestive tract of a bat to germinate under natural conditions. Given the sympatricity of V. planifolia and V. odorata , the same flowering season, and the prevalence of the similar herbal aroma with cinnamon-like notes in both species, is not unlikely that they share the same pollinator, apparently Euglossa viridissima in V. planifolia . Therefore, a spontaneous hybrid between V. planifolia and V. odorata may be the source of V. x tahitenss , with a later selfing or backcrossing with V. planifolia , already in Tahiti, that gave origin to the variation in cultivars, genotypes and chromosomic numbers. The following specimens are sterile, but their leaf morphology is so different to the rest of the Central American Vanilla, that we are convinced that they represent a different, undescribed species. The leaves are rather thin, but not as thin as those of subsect. Membranacea; the leaves are conspicuously obovate and abruptly mucronate.COSTA RICA [Interamericano de Ciencias Agrcolas], Turrialba, en el bosque, 600 m alt., 23-VIII-49, J. Lon 1692 US(2021492)! GUANACASTE: Large epiphytic vine in moist forest, El Silencio, near Tilarn, altitude about 750 m, Jan. 13, 1926, US(1309513)! There are very few records on vanilla dispersal, but bat dispersal had been suggested in V. insignis (M. Soto, unpublished data) and recently observed in V. pompona (N. Byrd, pers. com.). Bird dispersal is expected in some Asian species, as J.J. Smith and Rchb. f., and also in the closely related genera Cyrtosia. ACKNOWLEDGMENTS . Thanks are due to Bush Boake Allen work, especially to Drs. John Janes, Angela Ryan and Neil C. Da Costa. Dr. Phillip J. Cribb, Royal Botanic Gardens, Kew, encouraged us to conduct this work. The Comisin Nacional para el Conocimiento y Uso de la Biodiversidad, analysis of nucleotidic sequences of Mexican taxa. We are in debt to the keepers of the following herbaria, for permission to study the collections in their charge: AMO, AMES, BM, BR, CHAPA, CR, EAP, ENCB, F, G, GOEL, HEPF, INB, US, UV, W, WU, and XAL. Dr. Kenneth M. Cameron, Wisconsin, for sharing his ITS sequences of Epistephium and Lecanorchis. To the Unin de Vainilleros de San Felipe Usila, Oaxaca, to Ing. Heriberto Larios, for the facilities to visit and work in plantations in Oaxaca and Veracruz, Mexico. To Mariana Hernndez and Anglica Cibrin, placed at our disposal the facilities of AMO Herbarium. To the Direccin General de Conservacin Ecolgica, Secretara de Medio Ambiente, Recursos Naturales y Pesca (now SEMARNAT), Mexico, for the permits for collecting vanillas in Mexico, particularly in regions under the National System of Natural Protected Areas. To the staff of the Biosphere Reserve of Calakmul, Campeche, particularly to Esteban Martnez, who made it possible to study the populations of Vanilla insignis. Julio Arguijo guided MAS in Central Veracruz to locate wild plants of V. planifolia and V. pompona. To the staff of the Station of Tropical Biology, Chajul, Chiapas, for the facilities to work in the reserve. To Ulises Snchez, who collected spirit preserved material of the region Mazateca, in Oaxaca. To Dr. Thomas Wendt and the National Geographic Society for the logistics for the expedition to the region of UxpanapaSierra Tres Picos. To Dr. Germn Carnevali (CICY, Mrida, Mex.) and Joao Batista da Silva, from Belm, Brazil, who

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gave us material from Venezuela and Brazil, extremely taxa. To Jos Linares, Escuela Agrcola Panamericana, V. calyculata, and other species. To Febronio Tun Lpez, for information on the vanillas of northern Guatemala. To Prof. William Stern, Florida State Museum for material of V. trigonocarpa. To Dr. Thomas Croat, Missouri Botanical Garden, for access to material grown at the garden. To Mr. Neal Byrd, Ca. Agrcola La Gavilana Ltd., Costa Rica, and Professor Franco Pupulin for invaluable material and information on Costa Rican species. To M.Sc. Elvira Salas, Universidad de Costa Rica, for her help with Costa Rican materials. To Rolando Jimnez and Marco Lpez, who prepared several illustrations. To Dr. Gerardo A. Salazar for constant help, support and comments during the preparation of this work. To Msc. Laura Mrquez Valdelamar, Lab. de Bologa Molecular, Instituto de Biologa, UNAM, by sequencing labour. Finally, to our team in the Laboratorio de Gentica Molecular y Evolucin Instituto de Ecologa, UNAM, especially to Ms. Angeles Cortz, M.Sc. Rosalinda Tapia, Drs. Alicia Gamboa, Caroline Burgeff, and Francisca Acevedo for all their help in the laboratory. LITERATURE CITED cubanas: 22. Ames, O. 1934. An addition to the genus Vanilla . Bot. Mus. 2(8): 101-103. Ames, O. and D.S. Correll. 1952. Orchids of Guatemala. Fieldiana: Bot. 26: 54-60. Fontqueria 29: 160-161. Bouriquet, G. 1954. Le vanillier et la vanille dans le monde. en R. Portres [ed.] Encyclopdie Biologique 46. Paul Lechavalier. Paris. : 83-84. Bruman H. 1948. The culture history of mexican vanilla. Hisp. Am. Hist. Rev. 28(3), 360-376. Cameron, K.M. 2000. Gondwanan biogeography of Vanilloideae (Orchidaceae). Southern Connections Congress, Program and Abstracts: 25-26. Lincoln, New Cameron, K. 2003. Vanilloideae. Pp. 281-285 In: Pridgeon, Genera Orchidacearum vol. 3. Orchidoideae (Part two), Vanilloideae. Oxford University Press. Oxford. Carnevali F.-C., G., J.L. Tapia M., R. Jimnez M., L. Peninsula and a tentative checklist of the orchidaceae of the Yucatan Peninsula Biotic Province. Harv. Pap. Bot. 5(2): 383-466. Caracterizacin de dos tipos de Vanilla planifolia . Acta Bot. Mex. 25: 49-59. Chase, M.W. 2001. The origin and biogeography of Orchidaceae. pp. 1-5 In: A.M. Pridgeon, P.J. Cribb, Orchidacearum vol. 2. Orchidoideae (Part one). Oxford University Press. Oxford. studies. Taxon 40(2); 215-220. Vanilla The orchid of commerce. Pp. 477-508 in: C.L. Press Co. New York. Cibrin, J.A. 1999. Variacin gentica de Vanilla planifolia en Mxico. Thesis. Faculty of Sciences, National University of Mexico, Mexico City. 60 pp. Correll, D.S. 1944. Vanilla : its history, cultivation and importance. Lloydia 7, 236-264. Correll, D.S. 1965. Supplement to orchids of Guatemala and British Honduras. Fieldiana: Bot. : 31(7): 725. Croat, T.B. 1984. Flora of Barro Colorado Island. Stanford University Press. A revised annotated checklist. Monogr. Syst. Bot. Missouri Bot. Gard. 54 p. Dressler, R.L. 1979. Una Vanilla notable de Panam. Orquideologa 13(3): 229-232. Dressler, R.L. 1993. Field guide to the orchids of Costa Rica and Panama. Comstock Publishing Associates. Ithaca and London. Pp. 325-326. (Vanilla pompona Schiede). J. Essent. Oil Res. 9: 427431. In: Flora of Jamaica. Foldats, E. 1969. Orchidaceae. In: Flora de Venezuela. vol. In: Flora of the Lesser Antilles. Hgsater, E., M.A. Soto Arenas, G.A. Salazar Chvez, R. 2005. Las orqudeas de Mxico. Instituto Chinon A.C. Mxico City. Hamer, F. 1974. Las orqudeas de El Salvador. Ministerio de Educacin. San Salvador. Hamer, F. 1984. Orchids of Nicaragua, part 5. Ic. Pl. Trop. fasc. 12. The Marie Selby Botanical Gardens, Sarasota, Florida.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.352 LANKESTERIANA

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studies. Phytologia 14(1): 1-37. Rentier. 1989. Fleurs et plantes de Tahiti . Les Editiones Hoehne, F.C. 1945. Flora Brasilica XII, II. Orchidaceae. (Texto e Ilustrao). Secretaria da Agricultura, Industria e Comrcio de So Paulo. Brasil. Tab. 20. Kenny, J. 1988. Native orchids of the Eastern Caribbean. London, Macmillan Caribbean . of Vanilla and evolution in Orchidaceae. Lindleyana [Orchids] 76: 926-929. Lubinsky, P., K. M. Cameron;, M. C. Molina, M. Wolng, S. Lepers-Andrzejewski, A. Gmez-Pompa, y SK. SeungChul. 2008. Neotropical roots of a Polynesian spice: the hybrid origin of Tahitian vanilla, Vanilla tahitensis (Orchidaceae). Amer. J. Botany 95(8): 1040-1047. Luer, C.L. 1972. The native orchids of Florida. The New York Botanical Garden. orchids of Belize. A.A. Balkema. Rotterdam. high weight plant DNA. Nucl. Acids Res. 8: 4321-4325. Ptard, P. 1986. Quelques plantes utiles de Polynsie franaise et Raau Tahiti. Editions Haere Po No Tahiti, Pepeete. Portres, R. 1954. Le genre Vanilla et ses espces. In: Lechevalier, P. (ed.). Le vanillier et la vanille dans le monde. Paris. Purseglove, J. W. 1975. Tropical crops: Monocotyledons. J. Wiley, New York. R .J. 1981. Spices, vol. 2. Longman, London. history of hemiepiphytes. Selbyana 9, 61-69. Rain, P. 1986. Vanilla cookbook. Celestial Arts. Berkeley. Ray, T. S. 1990. Metamorphosis in the Araceae. Amer. J. Bot. 77: 1599-1609. Rodrguez C., R. L., D.E. Morta, M.E. Barahona, N.H. Williams. 1986. Gneros de orqudeas de Costa Rica: 70-71. Rolfe, R.A. 1896. A revision of the genus Vanilla . J. Linn. Soc. 32: 439-478. L’Orchidee 12:28. of euglossine orchid-bees (Apidae: Euglossini) in Panama. Oecologia (Berlin) 73: 321-333. Savolainen, V., P. Cunoud, R. Spichiger, M.d.P. Martnez, herbarium specimens in DNA phylogenetics: evaluation and improvement. Pl. Syst. Evol. 197: 87-98. Genetic variation in Vanilla planifolia (Orchidaceae). Econ. Bot. 61(4): 328-336 Schultes, R.E. 1960. Native orchids of Trinidad and Tobago: 39-40. J. P. 1992. Tropical forests and their crops, pp.357-364 . Cornell University Press. Ithaca. Soto Arenas, M.A. 1989. Listado actualizado de las orqudeas de Mxico. Orqudea (Mx.) 11: 233-277. Soto Arenas, M.A. 1994. Vanilla odorata, una especie de amplia distribucin. Orqudea (Mx.) 13(1-2): 295-300. Soto Arenas, M.A. 1999. Filogeografa y recursos genticos de las vainillas de Mxico. Proyecto J-101. Mexico City. Soto Arenas, M.A. 2003. Vanilla (generic treatment). Pp. 321-334 in: A.M. Pridgeon, P.J. Cribb, M.W. Chase vol. 3. Orchidoideae (Part two), Vanilloideae. Oxford University Press. Oxford. Soto Arenas, M.A. 2003. Vanilla insignis Ames. Pl. 700 in Mexico, parts 2 and 3. Icones Orchidacearum fasc. 6 and 6. Herbario AMO. Mxico D.F. Vanilla Plum. ex Mill. (Orchidaceae, Vanillinae). Lankesteriana 9(3): 355-398. Vanilla . Pp. 583587 in Manual de plantas de Costa Rica, vol. Missouri Botanical Garden,-INBio-Museo Nacional de Costa Rica. tcnica de Vanilla planifolia . In: Soto-Arenas, M. A. (ed). Informacin actualizada sobre las especies de orqudeas del PROY-NOM-059-ECOL-2000. Instituto Chinoin A.C., Herbario de la Asociacin Mexicana de Orquideologa A.C. Bases de datos SNIB-CONABIO. Proyecto No. W029. Mxico. D.F.. Toledo, V. M. 1982. Pleistocene changes of vegetation in tropical Mexico. Pp. 93-111 in: G.T. Prance (ed.) University Press, New York. Wendt, T. 1989. Las selvas de Uxpanapa, Veracruz-Oaxaca. Anales Inst. Biol. Univ. Nal. Autn. Mx. ser. Bot. 58: 29-54. rain forest. In: T.P. Ramamoorthy, R. Bye, A. Lot and J. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & DRESSLER — A revision of Central American Vanilla353

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Fa (eds.), Biological diversity of Mexico. Origins and distribution. Oxford Univ. Press Williams, L.O. 1951. The orchidaceae of Mexico. Ceiba 2: 34-35. Williams, L.O. 1956. An enumeration of the Orchidaceae of Central America, British Honduras and Panama. Ceiba 5(1): 18. Williams, N. H. 1982. The biology of orchids and euglossine bees. Pp. 119-171 in: J. Arditti (ed.), Orchid Biology, Reviews and Perspectives, 2. Cornell University Press, Ithaca, New York. Wood, J.J. 2003. Orchids of Borneo, vol. 4. Revised and Kew Publishing.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.354 LANKESTERIANA

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A NEW INFRAGENERIC CLASSIFICATION AND SYNOPSIS OF THE GENUS V ANILLA PLUM. EX MILL. (ORCHIDACEAE: V ANILLINAE) MIGUEL A. SOTO ARENAS 1 & PHILLIP CRIBB 21 Laboratorio de Gentica Molecular y Evolucin, Instituto de Ecologa, Universidad Nacional Autnoma de Mxico and 2 The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, UK355. 2010. LANKESTERIANA 9(3): ABSTRACT . An updated, annotated checklist of the species of the genus Vanilla is presented, with information on nomenclature, distribution, literature and examined specimens. A key for the determination of the species Vanilla is provided. RESUMEN . Se presenta un listado actualizado, comentado de las especies del gnero Vanilla , donde se proporciona informacion nomenclatural, distribucional, literatura y especmenes examinados. Se incluye una infragenrica de Vanilla . KEY WORDS / PALABRAS CLAVE : Orchidaceae, Vanilla infragenricaThe pantropical genus Vanilla Plum. ex Mill. belongs to the subtribe Vanillinae and the subfamily Vanilleae, it comprises some 106 species of hemiepiphytic vinous orchids (Soto Arenas, 2003). Recent phylogenetic analyses have shown that it is closely related to the South American genera Dictyophyllaria Garay and Epistephium Kunth and to the achlorophyllous genus Lecanorchis Blume (Cameron, 1996; Cameron et al., Information on vanillas has been collected for a phylogenetic analysis, but since no modern revision exists, it has been necessary to evaluate the status of all the proposed taxa. The results of this survey are presented here, pending the production a formal revision. Vanilla named before the establishment of the binomial system of nomenclature (Linnaeus, 1753) and their taxa were described from sterile material, and like other hemiepiphytic plants, vegetative traits vary determination. A particular problem is that several taxa have been described from immature buds, in which measurements, and shape of the segments can be substantially different from the morphology of uncertain. Furthermore, numerous species are known from a surprisingly small number of specimens, forests, and the low density of some populations. Vanillas from densely populated areas in the tropics apparently face severe conservation problems and their populations are often depleted. Additional complications are the poor preservation of the rotting before they dry, and the presence of some is subtle. A very large group of American species is pollinated by male Euglossine bees attracted by chemical compounds (Roubik and Ackerman, 1987; pers. obs.). In other orchids with similar pollination systems, there may there be little morphological differentiation, yet the species can be effectively separated by barriers imposed by the fragrance-

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collecting behavior of their pollinators (van der Pijl Our knowledge of Vanilla is therefore far from adequate to produce a good, modern revision of the group, although there have been important contributions, such as two revisions of Vanilla (Rolfe, 1896; Portres, 1954), the treatment by Klotzsch (1845) and regional accounts by Hoehne (1945), Cribb (1995), Luer (1972), Summerhayes (1968), Seidenfaden (1992), and Szlatchetko and Olzewski (1999). The revision by Portres, the last full account of the genus, although useful, is somewhat out-ofdate because, although we accept a similar number of species in this checklist compared with his revision (110 species), there are substantial differences. The description of several new species, improved collections of some poorly-known taxa clarifying their status, and our understanding of the distribution of the species and their phylogenetic relationships (Soto Arenas, 2003; Soto Arenas et al., unpublished results) have improved our understanding of the genus. The fragmentary or poor material of some taxa available to Portres (1954) led him to propose of the group. Therefore, the aims of this paper are to cladistic analysis of morphological and molecular data presented elsewhere (Soto Arenas, 2003; Soto updated checklist of the species and to stress where more information is needed. This revision will also a base for future research. Material and methods A bibliographic revision was conducted. Specimens were examined in the following herbaria: AMES, AMO, BM, F, G, K, L, NY, P, W. Special attention was paid to type and critical specimens, for the species were described from sterile or immature material. When possible, we examined cultivated of many taxa have been assessed with the aid of molecular data, published elsewhere (Soto Arenas, 2003; Soto Arenas et al., in press). Vanilla The genus Vanilla is a rather diverse assamblage of species of pantropical distribution. Despite the lack of morphological synapomorphies in the genus, there is strong molecular evidence that in prep., Cameron, unpubl. data). The following combination of characters are diagnostic for Vanilla : the hemiepiphytic vinous monopodial growth habit, with roots produced at each internode (these characters are found in some Galeolinae, a subtribe that may be considered part of Vanillinae, and Clemastepistephium wingless seeds with a hard seed coat (similar seeds are apparently found in Dictyophyllaria ). Vanilla is an ancient group, and most morphological geographic distributions. In many ways it is a very diverse genus compared with others in the Orchidaceae. Some authors might favor the idea that the genus should be split into more uniform assemblages with a more restricted geographic distribution. The type species, Vanilla mexicana , and most of Portres’ section Membranacea could be easily considered as the genus Vanilla with the rest of the species separated in a different genus. Strong molecular and morphological information supports this view, although it is equally correct (in terms of recognizing monophyletic taxa) to consider all of them in a single genus, although it has few morphological synapomorphies. The inner lineages of the genus may be supported by molecular characters, but most of them lack morphological synapomorphies that permit their recognition and we found that their acceptance as separate genera is not favoured by other authors. One of the main reasons to keep Vanilla in its traditional sense is to keep the traditional use of the name. The type species, V. mexicana , is in one of the two main clades, and the species cultivated for their aromatic fruits fall in the other one. To recognize both lineages as different genera would produce the following problems: the economically most important species in the LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.356 LANKESTERIANA

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Orchidaceae, to which the name “vanilla” or “vainilla” has been attached since pre-Linnean times. nomenclature has been historically complex, and conservation of the name for the cultivated species. We believe that to keep Vanilla in its traditional sense is congruent with the phylogeny of the group and to maintain the nomenclatural stability that has now been reached. Rolfe (1896) proposed a formal infrageneric Vanilla, recognising two sections: Foliosae and Aphyllae. Portres (1954) accepted Rolfe’s sections and, in Foliosae, proposed 3 subsections: Papillosae, Lamellosae and Membranacea. However, his subsections are invalid as he provided them with neither a Latin diagnosis nor a type. Furthermore, his sections and subsections are polyphyletic, except for subsection Membranacea which is paraphyletic (Soto Arenas, 2003). Section Aphyllae is polyphyletic, because the in the genus, or maybe is the plesiomorphic condition for the clade containing the Caribbean-Paleotropical taxa, in which leafy species are also included. The (V. penicillata) in the clade of V. planifolia. If sect. Aphyllae is polyphyletic, then sect. Foliosae is also polyphyletic. These sections are also hierarchically incongruent, because they do not reveal the existence of the two main clades in the genus: the membranaceous species and all the others. Porters included both V. wrightii (= V. bicolor) and V. palmarum, the most basal species in the clade containing the American nonmembranaceous leafy species, and V. weberbaueriana (= V. ruiziana) and V. pleei, to which he had no access to appropriate material (see comments under these species in the checklist) in subsect. Membranacea. Subsections Papillosae and Lamellosae are very heterogeneous and incongruent with our cladistic analyses. In order to be congruent with the phylogenetic analyses, we propose Vanilla to be formed by two subgenera: Vanilla and Xanata subgenus Xanata comprises two sections: Xanata and Tethya. Vanilla Species groups are indicated in an informal way. They are easily recognized morphologically, but there is little evidence of their monophyly, and in some cases they may well be paraphyletic. A. Vanilla subgenus Vanilla Vanilla subsect. Membranacea Portres in Bouriquet, Le Vanillier et La Vanille: 159 (1959). nom. illeg. scarcely or not at all differentiated from vegetative axis. Lip without a penicillate callus. Column united to the lip only at the base. Column with a subperpendicular, protruding anther (incumbent only 90), concave stigma, and smooth ventral surface. Type species: V. mexicana Mill. DISTRIBUTION: 15 species in the Neotropics, most diverse in southern Brazil. Subgenus Vanilla has never been used as a name for this taxon, but it is automatically applicable to the group of V. mexicana Mill., the type species of the genus. The genus Dictyophyllaria is practically identical V. parvifolia group. Its recollection and comparison with this subgenus and other Vanillinae is much needed. Two groups can be recognized, but their limits are not clear-cut: 1. V. mexicana group: Petals undulate; sepals usually inrolled; lip very wide, 3to 5-lobed, with a wide, obtuse apical lobe and numerous high keels or an basal triangular keel on the abaxial surface. Species placed here include Vanilla costaricensis, V. guianensis, V. inodora, V. martinezii, V. methonica, V. mexicana, V. oroana and V. ovata. 2. group: Petals not undulate; sepals not inrolled; lip 3-lobed, with a rounded-quadrate apical lobe, a blade with axial rows of warts; and column with a basal triangular keel on the abaxial surface. Species included here are V. angustipetala, V. parvifolia and V. verrucosa. Similar to these, but LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla357

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midlobe of the lip are V. bertoniensis, V. bradei, V. edwallii and V. organensis. B. Vanilla subgenus Xanata subgen. nov . a sectione typica plantis foliosis vegetato valde differt; callo penicellato labelli adaxialis glaber vel pubescens; anthera ventralis ad columna parallela; lobis stigmatibus emergentibus distinguenda. Typus: Vanilla planifolia G. Jackson. vegetative ones. Lip with or lacking a penicillato callus. Column united to the lip usually more than a half of its length. Column with a parallel, somewhat ventral anther, emergent stigmatic lobes, and with a smooth or often hairy ventral surface. DISTRIBUTION: pantropical. DERIVATION OF THE NAME: from the Mexican Indian Totonaco name for Vanilla, “xanat” or “shanat”. Totonaco Indians are thought to have brought Vanilla planifolia to cultivation. a. Vanilla subgen. Xanata sect. Xanata 3. Vanilla palmarum group: Ovaries calyculate; and united to the column only about a half of column length, with longitudinal pubescent to hirsute lines on the distal half of the lip. DISTRIBUTION: West Indies and South America. Species included here are V. bicolor, V. palmarum and V. savannarum. 4. V. trigonocarpa group: much crenulate margins; vegetative habit mesophytic; grade rather than a clade. DISTRIBUTION: Tropical America, from Costa Rica to Amazonia. Species included here are V. espondae, V. hartii, V. sprucei and V. trigonocarpa. 5. V. planifolia group: Flowers greenish; lip concave and with small papillae in the apical region. The South American species have an apical callus connected with the penicillate callus in the middle of the lip by rows of papillae and with the lip apex usually recurved DISTRIBUTION: Widespread in Tropical America, including the West Indies. Species included here are V. appendiculata, V. bahiana, V. cristagalli, V. denticulata, V. dubia, V. dungsii,, V. helleri, V. insignis, V. odorata, V. phaeantha, V. planifolia, V. ribeiroi, V. schwackeana, V. tahitiensis and V. uncinata. 6. V. penicillata group: to the American leafy, penicillate species. Petals subunguiculate, lip long-unguiculate, with penicillate callus, and villose apex. Fruit calyculate. Similar to the leafy V. ribeiroi belonging to the V. planifolia group. DISTRIBUTION: Guyana-Amazonia. The only species included here is V. penicillata. 7. V. hostmanii group: Flowers large, usually white with a yellow-orange lip; lip with thickened, radiating veins on the distal half that can be somewhat warty rather distichous bracts; Ovary, sepals, and petals granulosely keeled, very conspicuously so in some species; Lip surface puberulent-cellular-papillose in some species; perianth segments sometimes with crystal druses included. DISTRIBUTION: Widespread in Tropical America, except for the West Indies. Species included here are V. cribbiana, V. dressleri, V. gardneri, V. hostmannii and V. ruiziana. 8. V. pompona group: thick, xerophytic; lip rather smooth, except for the penicillate callus, and with a slightly thickened axial cushion running from the penicillate callus to the lip; fruits thick, trigonous, banana-like. DISTRIBUTION: Widespread in Tropical America, except for the West Indies. Species included here are V. calyculata, V. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.358 LANKESTERIANA

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chamissonis,, V. pompona, V. pseudopompona and V. vellozii. b. Vanilla subgen Xanata sect. Tethya Cribb, sect. nov. a sectione Xanata foliis absentiis vel praesentis, carnosis vel coriaceis; ovario non calycoso; manifeste vel obscure trilobato, ad columna liberato, in partim apicali lineis pubescentibus destituto distinguenda. Typus: Vanilla phalaenopsis Rchb. f. ex van Houtte. DERIVATION OF THE NAME: From the ancient Tethys Sea, thought to be the migration route of Vanilla into the Palaeotropics. 9. V. kinabaluensis group: Leaves usually very broad, and V. kinabaluensis); ovary stout, especially in New Guinean species; fruit thick and relatively short; tepals concolorous, greenish-yellowish, usually obtuse; lip species, trumpet-shaped, not unguiculate, more or less trilobed, with the midlobe often recurved and with rows of reddish subulate papillae that may be very V. kinabaluensis, or whitish and longer in V. siamensis. Most species of this group are seldom collected and many are imperfectly know, but several of them seem to be closely related or maybe Malay Archipelago and New Guinea. DISTRIBUTION: From S China to New Guinea, most abundant in Indonesia. Species included here are , V. giullianetii, V. kaniensis, V. kempteriana, V. kinabaluensis, V. klabatiensis, V. ovalis, V. platyphylla, , V. siamensis, V. seranica, V. sumatrana, V. utteridgei and V. wariensis. 10. V. albida group: and plants similar in appearance to the American V. planifolia unguiculate; tepals long; and lip with short apical papillae. Some species, such as V. moonii and V. yersiniana, and perhaps other have sulcate stems in the way of V. insignis. V. albida, V. andamanica, V. montana and V. yersiniana are closely related, differing in morphological details and distribution. V havilandii seems to belong here, but has a very V. moonii has buff-colored tepals and yellow lip with apical papillae arranged in two rows, representing perhaps a shared character with V. annamica and V. somai. DISTRIBUTION: Sri Lanka to SE Asia and the Malay Archipelago across to Borneo. Species included here are V. albida, V. andamanica, V. havilandii, V. montana, V. moonii, V. sanjappae and V. yersiniana. 11. V. annamica group: shaped; and the lip has two apical, parallel rows of papillae. DISTRIBUTION: S China, Taiwan and Indochina. Species included here are V. annamica and V. somai. 12. group: V. palembanica is very imperfectly known, but it seems to be allied to V. obtuse tepals, a lip with a deeply retuse midlobe, woolly rather than scaly penicillate callus, and truncate buds. DISTRIBUTION: Sumatra, Borneo and peninsular Malaysia. Species included here are and V. palembanica. 13. V. borneensis group: Callus penicillate, continuous with the pilose lip apex; plants with small, of corpuscles at base. They seem to be leafy versions of the V. aphylla-V. calopogon-V. wightii group. DISTRIBUTION: From India and SE Asia to Indonesia. Species included here are V. borneensis and V. diabolica. 14. V. imperialis group: Flowers large; lip trumpetshaped, similar to that of the V. planifolia group but shorter; callus absent; lip apex papillate.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla359

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DISTRIBUTION: Continental Africa. Species included here are V. grandifolia, V. imperialis, V. ochyrae and V. polylepis. 15. V. franoisii group: Flowers very small, whitish green with the lip rose-tinted at the base and with purple margins; lip entire or six-lobed, densely front. DISTRIBUTION: Madagascar. Species included here are V. franoisii and V. coursii. 16. V. chalotii group: Lip sacciform; midlobe reduced, recurved; side lobes well developed. DISTRIBUTION: West and equatorial Africa. Species included here are V. chalotii, V. nigerica and V. seretii. 17. V. africana group: Lip not sacciform; side lobes of lip united to the basal 1/2-2/3 of the column and forming deep sinuses with the midlobe; callus formed by scales or warts; column apex exposed. DISTRIBUTION: West and equatorial Africa, with a single species in Eastern Africa. Species included here are V. acuminata, V. africana, V. crenulata, V. cucullata, V. hallei, V. heterolopha, V. ramosa and V. zanzibarica. 18. V. barbellata group: Flowers green to greenish with a reddish-purplish lip; callus incipient, penicillate. DISTRIBUTION: Caribbean. Species included here are V. bakeri, V. barbellata, V. claviculata, V. dilloniana and V. poitaei. 19. V. aphylla group: Lip with very long trichomes and sometimes an incipient penicillate callus. Scarcely different from V. borneensis group, except for the lack of leaves. DISTRIBUTION: India to the Philippines. Species included here are V. aphylla, V. calopogon and V. wightii. 20. V. phalaenopsis group: rows of hairs besides the axial line; petals obtuse, very wide. DISTRIBUTION: Indian Ocean basin (East Africa, Comores, Mascarene Islands, Madagascar, with a single species in Sri Lanka and S India). Species included here are Vanilla decaryana, V. humblotii, V. madagascariensis, V. phalaenopsis, V. perrieri, V. roscheri and V. walkeriae. KEY TO VANILLA 1. Plants leafy, the blades well-developed and persisting several years, bracts persistent until the fruit ripens 2 2. Leaves soft, membranous, very thin in herbarium specimens; lip margins united to the column only at base for less than 4 mm; penicillate callus absent column length, usually for more than 8 mm; penicillate callus absent, wanting or often present 3 3. Lip deeply saccate at base 3. Lip not saccate at the base 4 4. Stem 4 mm in diameter; leaves very small, 2 cm or less long, much shorter than the internodes 5 4. Stem stouter, 5 mm or more in diameter; leaves much larger, usually more or less as long as the internodes or longer 6 5. Lip side lobes lacerate or deeply erose on front margin, the central callus on lip circular, densely papillate, the midlobe of lip densely papillate V. penicillata 5. Lip crenulate towards apex, the disc densely hairy V. franoisii 6. Lip hexagonal in outline V. coursiiLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.360 LANKESTERIANA

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6. Lip entire to 3-lobed 7 7. Lip 3-lobed in basal half 7. Lip entire or 3-lobed in apical half 8 9. Lip hexagonal V. coursii 9. Lip not hexagonal 10 10. Lip entire, broadest in middle, callus densely papillose, running from base of lip almost to tip V. annamica 10. Lip 3-lobed, broadest across the apex; callus on disc, elongate-elliptic, papillose, with a few scattered papillae on the mid lobe V. somai 11. Lip disc lacking a central almost quadrate tuft of erose or lacerate callus; callus papillose, hairy or of several erose or papillose ridges running the length of the lip 11. Lip with a central tufted or convex callus 12 long as broad or less; species from Old World 12. Lip with a prominent narrowed claw when spread; lip usually more than 1.5 times as long as broad KEY I maroon blotches deep in the throat of the lip; petals much broader than sepals; penicillate callus absent, but with rows of longitudinal papillae; species from the Indian Ocean basin: E Africa, Madagascar, Sri-Lanka, India, Seychelles, Comoros 2 callus and other trichomes towards the apex; species from America 8 2. Column of 14-17 mm long; sepals 25-30 mm long V. decaryana 2. Column 18-30 mm long; sepals 50-70 mm long 3 3. Flowers white with a yellow throat; veins of the petals and lip thin, almost straight V. walkerae 3. Flowers white with an apricot or pink throat or yellow with orange or maroon markings in the throat of the lip; veins of the petals and lip partially or totally sinuous-undulate 4 5. Lip papillose; column 20 mm long V. humblotii 5. Lip not papillose; column 27-30 mm long V. perrieri 6. Sepals 75-80 mm long; stem 12-20 mm in diamet. V. roscheri 6. Sepals 50-65 mm long; stem 8-10 mm in diam. 7 7. Lip oblong, acute, the margins undulate V. phalaenopsis 7. Lip obovate, obtuse, the margins not undulate V. madagascariensis column fused to the lip margins until stigmatic region; margins of the obscure lateral lobes of the lip lacerateV. penicillata 8. Stems thick, about 1 cm or more in diam. when mature in living condition; lip less than 50 mm long; lip apical 9. Leaf present at each internode, persistent, to half or more the length of the internode, leaves plane, apices hooked; lip green with maroon margin and veins, hairs yellow, margins irregular V. poitaeiLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla361

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9. Leaves not present at each internode, most or early deciduous, usually much less than half the length of the internode, leaf margin at least somewhat revolute, apices straight, curved, or hooked; lip not colored as above, margins more or less entire 10 of the lip to the apex, stem squarish V. claviculata 11. Lip less than 4 cm long, deeply 3-lobed at the truncate apex, with sinuses prominent V. barbellata 11. Lip more than 4 cm long, broadly rounded above with a projecting apical lobe, without distinct V. dilloniana KEY II neatly 3-lobed 13 2. Lip entire, pentagonal or hexagonal 3 2. Lip 3-lobed 4 3. Lip < 5 cm long, with basal, warty keels, species from Guatemala V. martinezii 3. Lip > 5 cm long, basal veins of the lip smooth, species from Amazonia and Guyanas V. guianensis 4. Midlobe of the lip strongly emarginate 5 4. Midlobe of the lip broadly elliptic, ovate, subquadrate 7 V. inodora 5. Lip with 3-keeled callus, converging and thickened and rugose near the apex 6 6. Bracts small, 5.0-7.5 mm long; lip not papillose V. methonica 6. Bracts foliaceous, 110-150 65– 71.5 mm; lip papillose V. oroana 7. Midlobe of the lip triangular, elongate, acuminate 8 7. Midlobe of the lip not conspicuously emarginate, acute, truncate or obtuse 11 8. Fertile bracts foliaceous 9 8. Fertile bracts not foliacous 10 9. Lip distinctly 3-lobed, 29 mm long V. bradei 9. Lip obscurely 3-lobed, 30-37 mm long V. organensis 10. Leaves 7-8 cm broad; column 25 mm long V. acuta 10. Leaves 5-7 cm broad; column 20-25 mm long V. bertoniensis 11. Midlobe of the lip very short, quadrate, truncate, scarcely extending beyond the lateral ones V. costaricensis 11. Midlobe of the lip ovate-elliptic, conspicuously longer than the lateral ones 12 12. Petals > 40 mm long V. mexicana 12. Petals < 38 mm long V. ovata 13. Lip with longitudinal, smooth keels, lacking papillae V. parvifolia 13. Lip with rows of papillae and keels 14 14. Lip 20 mm long V. verrucosa 14. Lip 23-29.5 mm long V. angustipetala KEY III 1. Flowers small; sepals < 22 mm long; lip conspicuously 3-lobed, the midlobe very short and recurved V. chalotii 1. Flowers large; sepals > 34 mm long; lip obscurely 3-lobed, the midlobe very broad; stems papillose-verrucose 2 2. Lip midlobe strongly recurved-inrolled, the base saccate but without 2 gibbae, the throat slightly hairy V. sereti 2. Lip midlobe recurved, but not rolled, the base with 2 gibbae, the throat densely and coarsely hairy V. nigericaLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.362 LANKESTERIANA

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KEY IV 1. Lip midcolumn straight; lip margins united to column in basal third only; species from Asia 1. Lip apical lobe acute or truncate; lip callus not papillose; column strongly arcuate; lip margins united to the column for only 2/3 of column length or less, leaving the column apex very exposed; species of continental Africa 2 2. Lip lacking a penicillate callus, but with a warty axial zone 3 2. Lip with a penicillate callus with scales, not warty, rather thin in substance 4 3. Midlobe of the lip truncate, broader towards the apex V. heterolopha length of midlobe V. hallei 4. Leaves small, long cuspidate; raceme dense, simple; midlobe of the lip oblong or obovate-elliptic, not axially thickened on the lower half ; callus a tuft of raised toothed crests 5 4. Leaves large, elliptic, acuminate; raceme lax to subdense, simple or branched 6 5. Mid-lobe of the lip oblong or obovate-elliptic, not axially thickened on the lower half V. africana 5. Mid-lobe of the lip oblong-elliptic, with an axial, thickened keel in the basal half V. cucullata lip blade strongly gibbose-conduplicate, with a penicillate callus at the top of the gibba V. crenulata 7. Lip blade slightly gibbose, strongly apiculate at tip; leaves elliptic to very broadly elliptic; species from Kenya, Tanzania, Uganda V. zanzibarica 7. Lip blade not gibbose, lip obtuse, apiculate; leaves oblong to elliptic; species from western Africa V. acuminata KEY V 1. Lip callus a more or less circular papillose mound; lip midlobe deeply emarginated, the margins of the sinus swollen into a U-shaped callus 1. Lip not as above 2 V. sanjappae 2. Lip central callus elongate-oblong, of papillae, dendritic hairs of irregularly lacerate low ridges; mid-lobe of papillae, ridges or glabrous 3 3. Leaves oblong-elliptic, up to 21 cm long and 8 cm broad, rather succulent; floral bracts less than 1.5 cm V. polylepis specimens V. grandifolia Costus-like); leaves less than 25 cm long 5 5. Lip apex triangular V. imperialis 5. Lip apex elliptic to semi-circular V. ochyrae KEY VI with reddish blotch at the lip apex 2 V. sumatrana LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla363

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2. Lip almost entire, cuneate; mid lobe of lip lacking a longitudinal papillate ridge 3 3. Column with dendritic hairs below stigma; apex of lip with a horseshoe shaped callus with a lacerate margin 3. Column glabrous; apex of lip with a V-shaped callus of papillae V. kinabaluensis V. siamensis 5. Flowers conspicuously purple-red blotched on the lip, at least in the papillae area 6 6. Lip cuneate, neatly 3-lobed, species from Philippines to Moluccas and New Guinea 11 7. Lip with a pair of rounded calli at the base V. diabolica 7. Lip lacking any basal corpuscular calli 8 8. Column hairy on ventral surface facing the lip V. utteridgei 8. Column glabrous 9 9. Lip apical margins lacerate-irregularly erose V. kempteriana 9. Lip apical margin not lacerate or erose 10 10. Lip as broad as long, apex shallowly emarginate V. kaniensis 10. Lip longer than broad, apex more or less rounded V. wariensis 12. Stems angular in cross-section V. giullianetii 12. Stems terete 13 13. Midlobe of the lip long haired V. ovalis 13. Midlobe of the lip papillose to short-haired 14 14. Column pubescent on inner surface; lip disc papillate at base V. platyphylla 14. Column glabrous; lip disc glabrous at base 15 15. Lip mid lobe slightly em n arginate; callus on mid-lobe oblong, densely papillose V. sumatrana 15. Lip mid lobe subacute; mid-lobe sparsely papillose all over V. seranica than 3 cm long V. havilandii lip more than 3.5 cm long 17 17. Tepals tan-colored; lip buff-colored; species from Sri Lanka V. moonii 17. Tepals greenish, cream or white; species from Andaman Islands to Indonesia 18 18. Tepals greenish, the lip white V. albida 18. Tepals whitish-cream, the apices sometimes greenish, the lip white 19 19. Apical trichomes conspicuous, c. 4 mm long V. montana 19. Apical trichomes short, less than 3 mm long 20 V. andamanica 20. Callus small, with lacerate scales; species from Indochina V. yersiniana KEY VII 1. Ovary calyculate, penicillate callus absent, leaves shorter than the internodes of the stem 2 V. bicolorLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.364 LANKESTERIANA

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2. Flowers concolorous, yellow, self-pollinated, with thick, strongly recurved about the middle, banana-like pods; plants from savannas, usually on palms 3 3. Lip less than 4.5 cm long; species from Cuba V. savannarum 3. Lip more than 5 cm long; species from South America V. palmarum V. espondae 6. Flowers large; lip more than 8 cm long V. trigonocarpa 6. Flowers small, lip less than 6 cm long V. sprucei 7. Flowers with white to greenish tepals, the lip usually white to greenish, the throat usually lined, the lip may be suffused with yellow, orange or brown on a white-green basal color, distal part of the lip often with papillae or 7. Flowers greenish-yellow, cream-yellow to yellow with the lip bright yellow to orange, without lines on the lip throat, distal half of the lip lacking papillae or trichomes apart from the penicillate callus 22 8. Leaves narrowly lanceolate to ensiform; lip distal part somewhat warty, with the margins erose, denticulate or 8. Leaves ovate-lanceolate, oblong to elliptic, not narrowly lanceolate to ensiform, distal part of the lip smooth, 9. Flowers small; lip 25-30 mm long; species from the Guyanas 9. Flowers large; lip 35-45 mm long; species widespread in the V. odorata 10. Distal part of the lip smooth, except for the penicillate callus, occasionally the axial keels of the lip slightly prominent 11 10. Lip with warts, small papillae or conspicuous, retrorse trichomes on distal part 13 11. Flowers small, white to greenish-white; lip less than 35 mm long; leaves short, up to 4 2 cm, much shorter than the internodes; stems less than 5 mm thick V. hartii 11. Flowers large, greenish cream; lip more than 60 mm long; leaves medium-size, more than 8 3 cm, slightly shorter than the internodes; stems stout, more than 7 mm thick 12 V. bahiana V. phaeantha 13. Lip bearing warts, papillae and conspicuous retrorse trichomes on the distal half 14 13. Lip without conspicuous retrorse trichomes on the distal part 20 14. Lip subentire, distal half of the lip completely covered by trichomes V. ribeiroi 14. Lip trilobed 15 15. Lip mid lobe linear, recurved; leaves spatulate-oblanceolate, mucronate V. appendiculata 15. Lip mid lobe not linear, leaves not spathulate-oblanceolate 16 16. Trichomes covering all the surface of the distal half of the lip, although they may be longer axially 19 17. Trichomes from the penicillate callus to near the base of the midlobe V. dungsii 17. Trichomes from the penicillate callus to almost the lip apex 18 18. Leaves 6 times as long as broad, tapering at base into an obscure petiole; lip midlobe much longer than the side lobes, acute V. dubia 18. Leaves about 3.5 times as long as broad, abruptly constricted at the base into a distinct petiole; lip oblongobovate, mid lobe emarginated V. cristagalliiLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla365

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19. Flowers very large; lip more than 8 cm long; stem conspicuously grooved and minutely papillose; leaves oblong V. insignis 19. Flowers not very large; lip less than 6 cm long; stem terete, smooth; leaves elliptic V. helleri 20. Leaves elliptic; species from Brazil V. schwackeana 20. Leaves oblong-elliptic; species from Central America (but escaped in all tropical areas) 21 21. Leaves oblong, about four times as long as broad V. planifolia 21. Leaves lanceolate, about six times as long as broad V. tahitiensis lip with a thickened axial cushion near the lip apex, the veins of the distal part of the lip not thickened 23 conspicuously distichous, lip without a thickened axial cushion near the lip apex, the veins of the distal part of the lip thickened 28 23. Lip conspicuously longer than the petals, the midlobe of the lip strongly emarginated 24 23. Lip about as long as the tepals, the midlobe rounded to slightly notched 25 V. calyculata V. chamissonis 25. Leaves very large, usually more than 22 11 cm, widely elliptic 26 25. Leaves medium-sized, less than 16 7 cm, oblong to oblong elliptic 27 V. pompona 26. Midlobe of the lip obscure 27. Lip strongly concave V. pseudopompona 27. Lip gently concave V. columbiana 28. Lip entire; sepals not conspicuously papillose, the papillae, if present, scattered 29 28. Lip obscurely 3-lobed; ovary, sepals and dorsal keel of the petals conspicuously granulose in the outer surface 30 29. Lip acute to subacute; leaves oblong, not acuminate V. gardneri 29. Lip apex widely rounded; leaves oblanceolate, acuminate-cuspidate V. dressleri lobe broadly ovate, with c. 10 thickened keels in the lip apex extending to the lateral lobes; species from Amazonia and Guyanas V. hostmanii 31. Mid -l obe of the lip subquadrate-ovate, with 5-6 thickened keels on the apical, central part of the lip, not extending to the lateral lobes, axial keel low; sepals rounded, slightly subcalyptrate; species from Central America V. cribbiana 31. Midlobe of the lip ovate, with c. 8 thickened keels, rather warty on the apical part of the lip, extending to the lateral lobes, 1-3 elevated, axial keels; sepals acute and conspicuously calyptrate; species from Peru and Bolivia V. weberbauerianaSYNOPSIS OF VANILLALANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.366 LANKESTERIANA Listed species, recognized as valid appear in bold face. Those described in Vanilla and considered to be synonyms are italicised. Vanilla Plum. ex Mill., Gard. Dict. abridged de. 4, 3: without page number (1754). Lectotype: Epidendrum vanilla L. = Vanilla mexicana Mill. (Mansf. in Myrobroma Salisb., Parad. Lond. t. 82 (1807). Vanillophorum Neck., Elem. Bot. 3: 184 (1790), nom. inval. 1. J.J. Sm. in Bull. Jard. Bot. Buitenzorg ser. 3, 2: 21 (1920). Types: Borneo, Koetei, Soengei Djongkang, Tengaroeng, March 1918, W.C. van Gelder 1; Soengei Landak, J.E. Teysmann s.n.; Kapoeas, J.E. Teysmann 109898 (all syn. BO). DISTRIBUTION : Borneo (Kalimantan Timur, Sarawak). Similar to V. kinabaluensis because of its long and large leaves, but and the apical hairs of the lip are less conspicuous.

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It has branched trichomes on the column below the stigma. Carr 3417 V. sumatrana, a plant apparently related to COLLECTIONS . SARAWAK: (K!); SABAH: Koung, Carr 3417 (K!). SOURCES : Smith, J.J. 1920. Bull. Jard. Bot. Buiten. 2: t. 6; Portres, R. 1954. Le Genre Vanilla et ses Espces: 2. V. acuminata Rolfe in J. Linn. Soc. Bot. 32: 456 (1896). Type: Gabn, Munda Prov., Soyaux 134 (holo. K!). DISTRIBUTION : Gabon, Congo. This species is very close to V. africana and stout plants of it are indistinguishable. The lip is said to have an oblong, obtuse, apiculate midlobe, and the disc bears a smaller callus with only about three scales. The lip is missing in the type specimen. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. acuta Rolfe = V. guianensis V. acutifolia Lodd. ex W.Baxt. in Suppl. Hort. Brit. 3: 655 (1850). nomen (Venezuela) 3. V. africana Lindl. in J. Linn. Soc. Bot. 6: 137 (1862). Types: Upper Guinea, Brass R., Barter 47 (syn. K-LINDL! isosyn. K!); Hort. Loddiges (K-LINDL! watercolor only). DISTRIBUTION : Guinea, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Gabon. Among the African complex, V. africana is characterized by its oblong-lanceolate, long acuminate leaves, the triangular, broadly rounded side lobes of lip, and the ovate midlobe. It is not always easy to of V. ramosa. COLLECTIONS : EQUATORIAL GUINEA: Mann s.n. (K!); W.W. Sanford 6032 (K!); Whyte (K!). GABON: Griffon de Bellay 501125 (W!); van der Burg 6691 (K!). CAMEROUN: Braun s.n. (K!). NIGERIA: Richards 3251 (BM!); Hort. Rotterdam Bot. Gard. (K!). GHANA: Johnson 798 (K!); Johnson 859 (K!); Bowling 36605 (K!). IVORY COAST: Hepper (K!); Prez-Vera 561 (K!). LIBERIA: Adam 20267 (K!). WITHOUT LOCALITY: Hort Schiller (W!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. ramosa); Summerhayes, V.S. 1968. Flora of West Tropical Africa 3(1): Orchidaceae: 205; Szlachetko, 100, A-F. 4. V. albida Blume, Catalogus 100 (1823). Type: Java, Gunong Salak, Parang, Burungrang, Blume s.n. (holo. L). DISTRIBUTION : Indonesia (Java and Sumatra). We interpret this species as the plant illustrated by Comber (1990) in Orchids of Java with conspicuous trichomes at the lip apex. Vanilla albida is similar to V. montana and V. yersiniana, and they have been submerged in the synonymy of the former by some authors. However, V. montana has larger trichomes at the lip apex, broader leaves, and V. yersiniana has shorter trichomes and whiter tepals, longer sepals and shorter lip. The plant illustrated by Comber (1990) was collected on Gunong Hambali, Lenkong, SW Java, not far from Blumes original locality. Comber (pers. comm.) indicated that the sepals become yellower with age. Blumes plate 67 Combers plant rather than to the Thailand plants V. albida in recent years that must treated as V. yersiniana. COLLECTIONS : INDONESIA, JAVA: Blumes Rumphia t.67!; Zollinger 961 (W!); C. Doran (K!); cult. Bogor s.n. (K! W!); (BM!); Curtis 2271 (BM!). SOURCES : Blume, C. 1825. Bijdr. 422; Portres, R. 1954. Le Genre Vanilla 97. V. anaromatica Griseb. = V. mexicana 5. V. andamanica Rolfe in Bull. Misc. Inform. Kew 1918: 237 (1918). Type: Andaman Islands, Betapur Valley, 25 Feb. 1916, C.E. Parkinson 1139 (holo. K!). DISTRIBUTION : India: Andaman Islands only. Often relegated to the synonymy of V. albida to which is closely related. However, it has a larger penicillate callus in which the scales are so divided, that they look more like hairs than scales. This, together with its isolated distribution in Andaman Islands, suggest that is better considered as a distinct species.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla367

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SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 220. V. angustifolia Willd., Sp. Pl. 4: 121. (1805). 6. V. angustipetala Schltr. in Anex. Mem. Inst. Butantan Secc. Bot. 1(4): 19 (1922). Type: Brazil, Sao Paulo, near Morro das Pedras, Iguap, Dec. 1916, A.C. Brade 7775 (holo. RJ; iso. AMES 114509! K!, R!). V. rojasiana Hoehne in Arq. Bot. Estado Sao Paulo n.s. 1: 61. t. 82 (1941). Type. Paraguay, Estancia Princeza, Caguazu, , cult. Asuncin, April-May 1927, T. Rojas 5082 AMES 58883! sterile). DISTRIBUTION : S Brazil, Paraguay and Argentina. We have examinated the type of V. angustipetala and it matches well the published drawings of V. rojasiana. This species seems to be close to V. parvifolia, based on Rodrigues’s original plate, which lip, and it lacks the papillae on the disc of the lip, and to V. verrucosa from them. COLLECTION : PARAGUAY: E. Hassler 1294 (K!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses 7. V. annamica Gagnep. in Bull. Mus. Natl. Hist. Nat. sr. 2, 3: 686 (1931). Types: Vietnam, Annam, col de Mangiang, Qui-nhon Prov., Poilane 17973 (syn. P; isosyn K!); Lang-bian, between Bdle and Dankia, Poilane 18635 (syn. P; isosyn. AMES 65778!); Langbian, Jacquet 622 (syn. P); Bihn-thuan Prov., Djiring, Mangnein 82 (syn. P). DISTRIBUTION : S China and Vietnam. This species was described from immature buds. are characteristic. Flowers structure of modern Vietnamese specimens are similar to those of V. somai apex. COLLECTIONS : CHINA: Guizhou, McGough ASBK 37 (K! PE!); Yunnan, Cavalerie 3427 (K!). SOURCES , A . 1932. Flore Genrale de LIndo-Chine 2. Orchidaces: 584 Vanilla Vietnamese Orchids: 76. V. anomala Pseudovanilla anomala 8. V. aphylla Blume, Bijdr. Fl. Ned. Ind. 8: 422 (1825). Type: Java, near Sadjvam Province, Bantam; Nusa Kambanga islands, Nov.-March, Blume 1719 (holo. L) Limodorum aphyllum Roxb., Pl. Coromandel 1: 34, pl. 41 (1795). Vanilla parishii Rchb. f., Otia Bot. Hamburg 1: 39 (1866). Type: Myanmar, Tanasserim, Parish 286 (holo. K!). D ISTRIBUTION : Myanmar, Laos, Thailand, Vietnam, Malaysia, Indonesia (Java). Vanilla of SE Asia. The lip of the specimen of V. parishii at Kew bears scattered hairs on the side lobes of the lip, and it is not clear if this represents a distinct character or if, as proposed by Seidenfaden, they are pieces of veins that have broken due to much boiling. Vanilla aphylla is closely related to V. calopogon of Philippines and V. wightii of India. COLLECTIONS : THAILAND: Kerr s.n. (K!); Kerr 0791 (K!); (K!); Put 1907 (K!). VIETNAM: Pierre 6565 (K!). INDONESIA: Java, Zollinger 599 (G7888/100! 101! K-LINDL!) SOURCES : Ridley, H.N. 1924. The Flora of the Malay Flore Genrale de LIndo-Chine 2. Orchidaces: 582; Averyanov, L. 1988. Predvaritelnyt spisok orkhidnykh: Orchids of Thailand: a preliminary list, part 1: 57, Thailand VI. Dansk Bot. Arkiv. 32(2): 142, 145-146, Vietnamese Orchids: 77; Holttum, R.E. 1964. Flora of Malaya. Orchids: 98; Comber, J.B. 1990. Orchids of Java: 75. V. aphylla Wight = V. wightii V. aphylla Eggers = V. poiteai 9. V. appendiculata Rolfe in Bull. Misc. Inform. Kew 1895: 178 (1895). Type: Guyana (British Guiana), LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.368 LANKESTERIANA

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Corentyne River, im Thurn s.n. (holo. K!). V. purusara Barb. Rodr. ex Hoehne, Arq. Bot. Estado. Sao Paulo, n.s. 1: 127. t. 140 (1944). Syntypes: Brazil, Amazonas, Bom-Lugar, Ro Purs, May 1904, Huber s.n. (MG 4721!); Par, Igap edge of the Pirapitinga, tributary of the Juta de Almeirim, 19 April 1923, Ducke s.n. (RJ 18704!). V. marowynensis Pulle, Enum. Vasc. Pl. Surinam: 118, pl. 4 (1906). Type: Surinam, Upper Marowyne River, near Poeloegoedoe , July, Versteeg 623 (not seen). DISTRIBUTION : Guyana, Surinam, Brazil and Peru. V. appendiculata is distinguished by its large, spatulate, abruptly mucronate leaves, and large Vanilla marowynensis illustration, based on an immature bud is inadequate. This species seems to be common in Amazonia. COLLECTIONS Cavalcante 659 (MG!); PERU, Loreto, (K!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses V. marowynensis). V. argentina Hicken = V. chamissonis V. aromatica Willd. = V. planifolia V. aromatica Sw. = V. mexicana V. articulata Northr. = V. barbellata 10. V. bahiana Hoehne in Arq. Bot. Estado. Sao. Paulo, litornea da Bahia, Nov. 1947, Ferreira Da Cunha s.n. (holo. SP 54661). DISTRIBUTION : Known only from Brazil. A xerophytic inhabitant of the coastal caatinga resembling those of V. pompona, with the midlobe of the lip very short. Its closest relative seems to be V. phaeantha. Like V. phaeantha, the leaves are usually much shorter than the internodes. Some specimens listed as syntypes of V. gardneri actually belong to V. bahiana. COLLECTIONS Harley 17366 et al. (K!); (K!). V. bakeri Schltr. = See V. barbellata . V. bampsiana Geerinck = V. planifolia 11. V. barbellata Rchb. f. in Flora 48: 274 (1865). Type: Cuba, C. Wright 3352 (holo. W; iso. AMES 70985! K! US!). V. articulata Northrop in Mem. Torrey Bot. Club. 12(1): 31, pl. 3 (1902). Types: Bahamas, New Deep Creek, June, Northrop 545 (syn. NY according to Correll, 1946). V. bakeri Schltr. in Repert. Sp. Nov. Regni Veg. 8: 561 (1910). Type: Cuba, Cojimar, Havana Prov., June 1905, F.C. Baker 5127 (holo. B); same locality and date, Baker 5127 (lectotype here designated HAC!). DISTRIBUTION : Florida, Bahamas, Cuba, Dominican Republic, Puerto Rico, Virgin Islands. Vanilla barbellata is one of the commonest species in the Caribbean Islands and Florida. It is conspicuously 3-lobed, truncate lip. The holotype of V. bakeri was destroyed at B; the only duplicate of it is housed at HAC, but it has only that it may have been based in an immature bud of V. barbellata, but details of the lip ornamentation do not coincide with the morphology of that species. The lectotype, although sterile, corresponds with the morphology of V. barbellata, the fruits being smaller those of V. dilloniana corresponds to that of V. barbellata and there is also an additional collection of V. barbellata from the same locality (Len 6278 HAC!). COLLECTIONS : Wright 3337 (BM!). CUBA: L.B. Smith et al. 3041 (AMES!); cf. P. Wilson 4592 (AMES!). BAHAMAS: N.L. Britton (K!); D. Morris s.n. (K!); N.H. (AMES!); (AMES!); L. Zemaitis (AMES!). DOMINICAN REPUBLIC: A. Liogier 11610 (AMES!) 15640 (AMES!). WITHOUT LOCALITY: cf. Pavn (actually Sess and Mocio) 20-1 (G7889/179!); Mocio 112 (watercolor!). SOURCES , 1962. The LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla369

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Bahama Flora: 84 (as V. articulata Catlogo Descritivo de las Orqudeas Cubanas: 21-22 (as V. articulata, 1946. Flora de Cuba 1: 351 (as V. articulata); Hawkes, A.D. 1948. Notes on Vanilla barbellata. Amer. Orchid Soc. Bull. 17(11): 652-653; Correll, D.S. 1946. The Soc. 15(7): 328-332. Luer, C. 1972. Native Orchids of Florida: 76, pl. 20; Ackerman. J.D. 1995. An Orchid Flora of Puerto Rico and the Virgin Islands: 171-174, V. barrereanaV. hostmanii V. beauchenei A. Chev., Etudes Fl. Afr. Centr. Franc. 1.297 (1913), nomen. 12. V. bertoniensis Bertoni in An. Ci. Parag. ser. I, 8: 10 (1910). Type: Paraguay, Puerto Bertoni, Bertoni 6039 (holo. not located). DISTRIBUTION : Paraguay. A scarcely known plant, described as larger than V. perexilis (= V. parvifolia), with contorted sepals and a lip with a triangular acuminate midlobe. However, the ovary was described as calyculate, a trait unknown in other members of this group. We suspect that this may be the earliest name for the plant currently known as V. edwallii. SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 262-264. 13. V. bicolor Lindl. in Edward’s Bot. Reg. 24: Misc. 58 (1838). Type: Guyana (British Guiana), Schomburgk s.n. (K-LINDL, not traced); lectotype (designated by Christenson, 1995) Guyana, Demerara, watercolor illn. by Schomburk s.n. ex Hort. Loddiges (K-LINDL!). V. claviculata sensu Lindl. in Ann. and Mag. Hist., ser. 3, 1: 334 (1858), non Sw. V. wrightii Rchb. f., Flora 48: 273 (1865). Type: E Cuba, 1856-7, Wright 672 (holo. W! iso. AMES V. cf. barbellata) BM! G7889/135! K! MO! P!). V. gratiosa Griseb., Cat. Pl. Cub. 267 (1866). Type: Based on same type as V. wrightii Rchb.f. V. palmarum sensu Grisebach, Cat. Pl. Cub. 267 (1866), non (Salzmann ex Lindl.) Lindl. DISTRIBUTION : Cuba, Jamaica, Hispaniola, Venezuela, Guyana, Surinam, French Guyana, Trinidad, and Brazil. A very distinct Caribbean and Guyanan species. The slender plants with small, elliptic leaves, which are much shorter than the internodes, elongate pods, tan-colored tepals, a cream-yellowish lip with scattered hairs on the disc, and a poorly penicillate callus are characteristic among American vanillas. It differs from other Vanilla species, in that several populations are found in mountain habitats up to 1700 m elevation. Molecular data suggest that it is the sister species of all the American penicillate vanillas. This species is widely known as V. wrightii, a by Sarah Thomas, from Kew and Eric Christenson published a well illustrated paper on the correct identity of this species. COLLECTIONS : J.A. Steyermark, G.C.K. (AMES!); B. Maguire 32967 (AMES!); (AMES!). J.B.F. da Silva 263 (MG!); Amazonas: B.G.S. Ribeiro 1028 (MG!); ?col. 7885 (BM!). GUYANA: G.S. Jenman 803 4561 (K!); E.F. im Thurn 43 (K!); 23176 (K!); (K!); Tillett, R. Boyan 44880 (K!). SURINAM: Hostmann s.n. (K!); B. Maguire 24258 (K!). JAMAICA: H. Wood (BM! drawing); [?Langley] 7885 (BM!). DOMINICAN REPUBLIC: B. A.H. Liogier 11474 11742 (AMES!); W.L. Talbot 2363 (AMES!). HAITI: E.L. Ekman 4355 (AMES! K!). CUBA: Eggers 5248 (K!); J.A. Schafer 3218 4047 (AMES!); B. Clemente 1956 6626 (AMES!); B. Alain 373 (AMES!); (AMES!); F. Len, F. Clement, F. M. Roca 9992 (AMES!(; B. Len 15646 (AMES!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 222 and 259-260 (as V. wrightii); Christenson, E.A. 1995. The long-lost Vanilla bicolor. Amer. Orch. Soc. Bull.: 64(8): 844-847; Schultes, R.E. 1960. Native Orchids of Trinidad and Tobago: 44 (as V. wrightii); Foldats, E. 1969. Flora de Venezuela. Orchidaceae 1: 127-129 (as V. wrightii); Gloudon, A. V. wrightiiLANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.370 LANKESTERIANA

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las Orqudeas Cubanas: 22-23 (as V. wrightii); Len, V. wrightii); Martius, F.P., Fl. Brasiliensis 3: t. 4: 149. 1896. 14. V. borneensis Rolfe in J. Linn. Soc. Bot. 32. 460 (1896). Type: Kalimantan, Banjarmassing, Motley 1248 (holo. K!). V. pilifera Holl ttum in Gard. Bull. Singapore 13: 251-253 (1951). Type: Malaysia: Kota Tingii, Johore, le Doux s.n. (holo. SING). DISTRIBUTION : Reported from NE India, Thailand, Malaysia and Indonesia (Kalimantan). This species was described from a fruiting specimens are known from southern Kalimantan, where the species was originally collected. The elliptic-oblong, long acuminate leaves are different from other Bornean vanillas, such as . The V. havilandii (or the Javanese V. albida V. and V. kinabaluensis. However, it looks very similar to specimens of V. pilifera collected in Sabah, northern Borneo, especially in leaf shape V. pilifera have much narrower, lanceolate, acute leaves, and shorter V. montana, from peninsular Malaysia, has oblanceolate, abruptly acuminate leaves that are also similar. It is easily distinguished by its lanceolate to elliptic, often narrow, short leaves and conspicuously hirsute disc of lip. It is closely related to V. diabolica, from Sulawesi. A Burbidge specimen (Burbidge s.n., BM) from Sabah, NW Borneo (Tawaran and Tampasuk rivers), may belong to V. kinabaluensis, rather than to V. borneensis, as suggested by Rolfe. COLLECTIONS : THAILAND: Put 2285 (BK C K!). MALAYSIA: SABAH: A. Lamb 87/83 (K! specimen and drawing); Daim Andau 440 (K! sterile); SAN 122464 (K! sterile); SAN 68739 (K! sterile); A. Talip 54914 (K! sterile); J. Beaman 7804 (K! sterile); L. Madani SAN 89511 (K! sterile). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 215-216; Portres, R. 1954. Le Genre Vanilla et ses Espces: 169-170 (as V. pilifera); Teo, C.K.H. 1985. Native Orchids of Peninsular Malaysia: 108109 (photo. of V. aphylla?; as V. pilifera); Seidenfaden, illustration based on TS 4075, may be V. yersiniana??; as V. pilifera); Seidenfaden, G. 1978. Orchid Genera 88 (as V. pilifera Orchids of Penninsular Malaysia and Singapore 125 (as V. pilifera). Holttum, R.E. 1964. Flora of Malaya. V. pilifera). 15. V. bradei Schltr. ex Mansf in Repert. Sp. Nov. Regni Veg. 24: 243 (1928). Type: Brazil, Sao Paulo, A.C. Brade 7573 (holo. B, AMES drawing!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 258-259. 16. V. calopogon Rchb. f., Otia Bot. Hamburg. 1: 40 (1878). Type: Philippines, Luzn, Cuming 2070, 1841 (holo. W!; iso. BM! G7889/99 168! K-LINDL! K!). DISTRIBUTION : The Philippines (Luzn). This species is very closely related to V. aphylla and V. wightiana. However, it has slightly larger COLLECTIONS Loher 630 (K!). SOURCE : Valmayor, H.L. 1984. Orchidiana Philippiniana 1: 322. 17. V. calyculata Schltr. in Repert. Sp. Nov. Regni Veg. 7: 42 (1920). Type: Colombia, M. Madero s.n. (holo. B). DISTRIBUTION : Honduras, El Salvador and Colombia. The status of V. calyculata remains uncertain and no original material has been located. However, the description suggests that the name must be applied to the specimens listed below. Its calyculate ovary, long emarginate midlobe of the lip, small leaves and the relatively high elevation of its habitat are characteristic. This species is closely related to V. chamissonis from Brazil. Specimens from El Salvador V. phaeantha (Hamer, 1974). COLLECTIONS : EL SALVADOR: Pank in Hamer 203 (AMES!). HONDURAS: Comayagua, al. 6045 (AMES! G78889/152! K!). COLOMBIA: LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla371

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Valle, Dryander 2379 (BM!); Tolima, Lehmann 2263 (BM! G7889/175!); Lehmann 8378 (K! watercolor); La Plata, cf. Lehmann 6278 (AMES!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 247. Hamer, F. 1974. Las Orqudeas de El Salvador 2: 375, t. 41 (as V. phaeantha) V. carinata Rolfe = see in excluded species 18. V. chalotii Finet in Bull. Soc. Bot. France, 56: 102 (1909). Type: Gabon, near Libreville, Chalot 4027 (holo. P). DISTRIBUTION : Endemic to Gabon. This species has a conspicuous, shallowly saccate lip, with a very reduced, recurved, apical lobe. It is similar to V. nigerica and V. seretii, in which the stems much larger. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Olszewski. 1999. Fl. de Cameroun: 280-282, pl. 100, A-F. 19. V. chamissonis Klotzsch in Bot. (1846). Type: Brazil, Chamisso s.n. (lecto. LE! photo). DISTRIBUTION : Brazil. Two different taxa have been called V. chamissonis in recent times. The true V. chamissonis has a very long midlobe of the lip (e.g. drawings of Portres, illustration of V. chamissonis var. longifolia); the other one is V. vellozii, reduced by most recent authors to a variety of the former. Vanilla chamissonis is similar to the V. calyculata from Colombia and Central America. Vanilla vellozii is different in its shorter segments and subentire lip, and it is not clear if it is a member of the V. chamissonis or V. pompona complexes. COLLECTIONS Glaziou 15661 (K!); The specimen Glaziou 14302, “Environs of Rio Janeiro”, was assigned by Rolfe (1896) to V. vellozii, but, have the very long lip characteristic of V. chamissonis; a similar case occurs with the Glaziou specimen from Copacabana (apparently that considered by Rolfe to be his number 11621). Goias, cf. A. Glaziou 22151 (G 7889/115!). BAHIA: M. Blanchet s.n. (G 7889/114)!; Dusn s.n. (AMES 86539!). Rio de Janeiro, Miers (BM!); L.A. Gattoni s.n. (AMES 66126! AMES 66339! AMES 66345!). SOURCES : Martius, F.P., Fl. Brasiliensis 3: pt. 4: 148, t. 32. 1896; Portres, R. 1954. Le Genre Vanilla et ses V. vellozii), p. 191-192 (as V. argentina); Pabst, G.F.J. 1966. As. Orqudeas do Estado da Guanabara. Orqudea 28(2-6): 169-170. 20. V. claviculata (W.Wright) Sw. in Nov. Act. Soc. Sc. Upsal. 6: 66 (1799). Type: Jamaica, Swartz s.n. (lecto. BM!, here designated). Epidendrum claviculatum W. Wright in London Med. J. 8(3): 254 (1787). DISTRIBUTION : Cuba, Jamaica, Hispaniola, Grand Cayman, Puerto Rico, St. Thomas. It bear fugaceous, small leaves, up to 8 1 cm, which are conspicuosly apiculate. The lip is not conspicuously trilobed at apex, as its allies, but rather rounded, and it has a penicillate callus of dense forked hairs and rows of similar hairs towards the apex, but not dense. COLLECTIONS : CUBA: (AMES!); R.A. Howard 6522 (AMES!); A.H. Liogier 11493 (AMES!); R.M. Gray s.n. (AMES! K!). DOMINICAN REPUBLIC: (AMES!). PUERTO RICO: A.G. Kevorkian s.n. (AMES!). JAMAICA: H. Wood (BM! drawing); C. Whitefoord 1423 (BM!). GRAND CAYMAN: M. Brunt 2186 (BM!); W. Kings G.C. 412 (BM!). SOURCES : Sloane, H. 1725. Jamaica Nat. Hort. t. 224; Catlogo Descriptivo de las Orqudeas Cubanas: 23 (as V. parasitica); Correll, D.S. 1946. The American 1995. Orchids of Jamaica.: 206-207; Castillo Mayda, Rico and the Virgin Islands: 52, pl. 10; Ackerman. J.D. 1995. An Orchid Flora of Puerto Rico and the Virgin 5. V. claviculata sensu Lindl. in Mem. Am. Acad. n.s. 8: 219 (1861) = V. bicolor 21. V. columbiana Rolfe in J. Linn. Soc. Bot. 32: 468 (1896). Type: Colombia [New Granada], Valley of the Magdalena, February, Goudot s.n. (holo. K!).LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.372 LANKESTERIANA

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Th e type material is very unsatisfactory, making an accurate description of this species problematic. The has a very neat midlobe to the lip. We have assigned to this concept specimens from the V. pompona complex with small leaves, resembling those of V. phaeantha , lip with a yellow edge, but we would like to examine more material from the Magdalena Valley. It is possible that they might better be attributed to the plant we are treating here as V. calyculata . COLLECTIONS : COLOMBIA: Valderama 35275 (AMES!); H. Garca-Barriga 15373 (AMES!); (AMES! MO). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces V. correlliiV. poitaei 22. V. costaricensis Soto Arenas ined. DISTRIBUTION : Endemic to Costa Rica. This new species of Vanilla is closely related to V. mexicana, from the West Indies and NE South America. However, V. costaricensis has a very short midlobe to the lip and large bracts similar to those of V. inodora Schiede. The species was illustrated by Rafael Lucas Rodrguez (1986), but apart from this record, it is known only from the type and another collection from the same locality. As far as is known, V. mexicana is not found in Central America. SOURCES : Lucas Rodrguez, R., D.E. Morta, M.E. Orqudeas de Costa Rica: 70-71. 23. V. coursii H. Perrier in Rev. Int. Bot. Appl. Agric. Trop. 30: 435 (1950). Type: Madagascar, N of Antalaha, Cours G6 (holo. P!) DISTRIBUTION : Endemic to Madagascar. Very poorly known. We have been unable to examine material of this leafy Madagascan species, of the island. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 1 C. Hermans. 1999. The Orchids of Madagascar: 207. 24. V. crenulata Rolfe in J. Linn. Soc. Bot. 32: 477 (1896). Types: Sierra Leone, Bumban to Lokko, Scott Elliot 5733 (syn. K!; isosyn. BM!); Ashanti, Prahsu, Cummins 4 (syn. K!). DISTRIBUTION : W and WC Tropical Africa. Another member of the V. africana complex, V. ramosa. It is characterized by its broad leaves, unbranched acute crenulate front lobe of the lip. Characteristic is the presence of very broad, oval bracts at the base of the raceme. Summerhayes (1968) also indicated a larger lip, midlobe and column in V. crenulata, compared with V. ramosa. In living condition, V. crenulata distinguished by their conduplicate, gibbose midlobe of the lip, with a penicillate callus at the top of the gibba. In their relatives (e.g. V. ramosa) the lip considered V. crenulata to be a synonym of V. africana. COLLECTIONS : GUINEA: Schnell 3086, 799 (K!). LIBERIA Bunting s.n. (BM!); Barker 1265 (K!); Jansen 1013 (K!); Baldwin 10661A, 10966, 11116 (K!); Whyte s.n. (K!); Blickenstaff 92 (K!); Linder 46, 76 (K!). GHANA: Fishlock 57 (BM! K!); Wilson 1036 (K!); Bowling 38137 (K!); Westwood 112 (K!); Irvine 1186, 1808 (K!); Vigne 3062 (K!); Darko 1018 (K!); Morton GC6553 (K!). IVORY COAST: PrezVera 642 (K!); de Wilde 576 (K!); Chevalier B22383 (K!); Leeuwenberg 3054 (K!). NIGERIA: Miles s.n. (BM!). GABON: Le Testu 1413 (BM!); van der Laan 834 (K!). SIERRA LEONE: Deighton 2496, 2963, 6151 (K!); Dawe 439 (K!); Bobob s.n. (K 37879!); Segerbck 1537 (K! also photos); Thomas 3432 (K!). PRINCIPE: Exell 623 Lebrun 3241 (K!). WITHOUT LOCALITY: Johnson 24 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses 242; Summerhayes, V.S. 1968. Flora of West Tropical Africa 3(1): Orchidaceae: 205-206. Johansson, D. 1974. Acta Phytogeographica Suecica 59: 44-45. 25. V. cribbiana Soto Arenas, ined. DISTRIBUTION : Known from Mexico (Veracruz, Oaxaca, Chiapas), Guatemala, Belize, and Honduras.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla373

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.374 LANKESTERIANA Vanilla cribbiana is a member of the V. hostmanii group. Its fruits are fragrant, and probably of some interest, but the vine is not a strong grower and does not produce fruits regularly. COLLECTIONS C. Whitefoord 1816 (BM!). HONDURAS: O. Ames II-211 (AMES!). 26. V. cristagallii Hoehne in Arq. Bot. Estado. Sao. Paulo, n.s. 1: 125 (1944). Type: Brazil, Amazonas, Sao Paulo de Olivena, 23 Oct. 1931, Ducke s.n. (holo. RJ 14512!). V. cristato-callosa Hoehne in Arq. Bot. Estado. Sao. Paulo, ns .s. 1: 126. t. 137 (1944). Type: Brazil, Amazonas, Manus, 20 Oct. 1935, Ducke s.n. (holo. RJ 37468!). DISTRIBUTION : Amazonian Brazil. This species, like several other Amazonian species, has an apical lobe adorned with retrorse trichomes or transverse crests. The difference between the types of V. cristagalli and V. cristato-callosa is mostly in the the same species. SOURCE : Hoehne, F.C. 1944. Arq. Bot. Estado Sao Paulo, n.s. 1: 125. 27. V. cucullata Mitth. Deutsch. Schutzgeb. 2: 161 (1889). Type: Cameroon, Braun 4 (holo. B). V. africana Lindl. subsp. cucullata (Kraenzl. ex J. Cameroun 34: 286 (1988). Perhaps not different from V. crenulata. In this species of the V. africana complex, the oblong-elliptic lip is acute, and the petals and sepals rather broader. COLLECTIONS : CAMEROUN: Zenker 525 (AMES! BM! G 7889/116 /117 /118! P). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 195-196. 28. V. decaryana H. Perrier in Bull. Mus. Hist. Nat. (Paris) 6: 194-195 (1934). Types: Madagascar, Domaine du Sud-Ouest: Morondava, Greve 39 (syn. P!); Ambovombe, Jan. 1925, Decary 3531 (syn. P!; isosyn. K!); Behara, E of Ambovombe, 17 Nov. 1931, Decary 9313 (syn. P!); Mahatomotsy, N of Ambovombe, 9 Dec. 1931, Decary 9503 (syn. P!). DISTRIBUTION : SW Madagascar. A member of the V. roscheri-V. madagascariensis-V. phalaenopsis group, but with smaller more numerous long and with narrower rhombic lip with the keels of the disc just to near the border of the lip, more COLLECTIONS : MADAGASCAR: O’Connor 98 (K drawing!). SOURCE Hermans. 1999. The Orchids of Madagascar: 207. V. denticulata Pabst = V. odorata 29. V. diabolica P.O’Byrne in Malayan Orchid Rev. Tenggara-Buton Island, near Maligano, O’Byrne SB47V (holo. SING; drawing at K!). This species is closely related to V. borneensis, but V. diabolica shorter and broader tepals and a lip in which the veins of the throat are orange rather than purple. 30. V. dilloniana Correll in Amer. Orch. Soc. Bull. 15(7): 331, f. 1-7 (1946). Type: USA, Florida, Brickell Hammock, Dade County, Humes s.n. (holo. AMES). DISTRIBUTION : Cuba, Florida, Hispaniola, Puerto Rico, and Saint Thomas. trumpet-shaped 3-lobed lip, edged with rich purple, in which the midlobe is triangular and longer, and with yellowish papillae towards the lip apex. The only known population in Florida has been wiped out, although plants survive in cultivation. material is at hand. Vanilla clavicula has an entire lip; from V. barbellata it is distinguished by its larger specimens bear small leaves, similar to those found in V. claviculata and V. poitaei, but rarely seen in V. barbellata. Vanilla poitaei has a larger lip and the margins are crisped. COLLECTIONS : CUBA: Wright s.n. (K-LINDL!); Jack 5200 (AMES!); Wright s.n. (AMES 70987, in part)! PUERTO RICO: Pennington s.n. (AMES 45836!); Pennington s.n. (AMES 43655!); Sintenis 3932 (BM!); Sintenis 4934 (K! fruits only). DOMINICAN

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla375REPUBLIC: Fuertes s.n. (AMES!). ST. THOMAS: Ostenfeld 235 (G7889/119!). SOURCE S : Ackerman, J.D. 1995. An Orchid Flora 93. Britton, N.L. and C.F. Millspaugh. 1962. The Bahama Flora: 83 (as V. eggersii ). McCartney, C. V. dilloniana ). V. domestica Hort., nomen = ?V. planifolia V. dominiana Hort. ex Gentil, Liste Pl. Cult. Serres Jard. Brux. 192 (1907), nomen. 31. V. dressleri Soto Arenas ined. DISTRIBUTION : Costa Rica, Panama and W Colombia. V. hostmanii group. Among the group, it is distinguished (c. 1 cm apart), patent bracts, subentire lip, with a subrhombic blade, and very scarcely papillose sepals; V. cribbiana, V. hostmanii and V. ruiziana have conspicuously granulose sepals. From V. cribbiana it is additionally distinguished by the narrower, thinner leaves which are more long acuminate-cuspidate, of the disc more conspicuously thickened, and August. Vanilla hostmanii from Amazonia has very (e.g. 40-50) and the leaves are larger, thicker, not as acuminate-cuspidate. Vanilla ruiziana, from Peru and Bolivia has larger, broader, less acuminate-cuspidate, and thicker leaves, but is similar to V. dressleri in their rather elevated axial keels of the lip. Vanilla gardneri from Brazil has a subacute lip apex and oblong leaves, not cuspidate. COLLECTIONS : PANAMA: (MO!); Pridgeon s.n. belong to V. pseudopompona); Taylor 768 (SEL!). COSTA RICA: Cruz (CR MO!); Herrera 1840 (CR MO 3864177!); Liesner 14435 (MO!); Todzia 1291 (LL!); Lankester s.n. (SEL!). COLOMBIA: Cuatrecasas 16550 (AMES!); Cuatrecasas 15220 (AMES!); Agualimpia 325 (MO!); Choco, Ingram 1421 (SEL!); Ingram 1422 (SEL!). 32. V. dubia Hoehne in Arq. Bot. Estado Sao Paulo n.s. 1: 126 (1944). Type: Brazil, Minas Gerais, BeloHorizonte, Schwacke 11107 (holo. RJ 37014!). DISTRIBUTION : Brazil. Closely related to V. appendiculata, V. cristagallii and V. schwackeana, and with the characteristic papillose apical apex of the lip of many Brazilian species. V. duckei sensu Hoehne (1945) = V. planifolia 33. V. dungsii Pabst in Bradea 2, 10: 49 (1975). Type: Brazil: Estado do Rio de Janeiro, near Novo Friburgo, 1300 m, Febr. 1973, Dungs s.n. (holo. HB 59052!). DISTRIBUTION : Brazil. Little is known of this species. It was stated in the original description that it is allied to V. cristatocallosa (= V. cristagallii). The clump retrorse papillae towards the apex of the lip is certainly characteristic of that group of Vanilla. V. eggersi Rolfe = V. poitaei V. ensifolia Rolfe = V. odorata V. epidendrum Mirb. = ?V. mexicana 34. V. edwalli Hoehne in Arq. Bot. Estado Sao Paulo n.s. 1: 61. t. 81 (1941). Type: Brazil, Sao Paulo, Serra da Cantareira, Jan. 1902, Gustavo Edwall 6003 (holo. SP). DISTRIBUTION : Brazil and Argentina. A species with long acuminate segments and a rather unornamented lip, except by the longitudinal lamellae. It is very probable that V. edwalii is a synonym of V. bertoniensis Bertoni, which original material has not been available to us. From V. parvifolia differs by its acuminate midlobe of the lip. COLLECTIONS E.P. Heringer 5001 et al. (MG!). 35. V. espondae Soto Arenas ined. DISTRIBUTION : Endemic to Colombia. This is one of the most beautiful Vanilla species. It and on a tributary of Magdalena River. Although is not closely related to any other Vanilla species, it could be related to V. trigonocarpa. It is distinguished by the very large papillae covering the lip apex. V. fasciola Spreng. = Taeniophyllum fasciola

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.376 LANKESTERIANA36. Rolfe in Bull. Misc. Inform. Kew 1899: 133 (1899). Type: Guyana (British Guiana), Barima River, Jenman 6771 (holo. K!; iso. K!). DISTRIBUTION : Brazil and Guyana. Very close to V. odorata The type material is not good enough to check the differences with V. odorata smaller (the lip 25-50 mm long whereas it is 3545 mm in V. odorata). This could be a result of the preservation process. However, the leaves are rather other V. odorata specimen that we have examined. COLLECTIONS : PARA: Plowman et al. 9868 (MG!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses V. fragrans Ames = V. planifolia 37. V. franoisii H. Perrier in Notul. Syst. (Paris) 8: 37 (1939). Type: Madagascar, S Toamasina, Ambilo, cult. Franois (holo. P!). DISTRIBUTION : Endemic to Madagascar. A very rare species of which we have seen the type and one other collection. It was described as having slender stems, very small leaves and small, fewV. perrieri. COLLECTION : MADAGASCAR: between Ampanihy and Andraka, Capuron 479 (P!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 180-181; Du Puy, D., P. Cribb, 207. 38. V. gardneri Rolfe in Bull. Misc. Inform. Kew 1895: 177 (1895). Types: Brazil, Morro do Flamengo, near Rio de Janeiro, Gardner 245 (lecto., K! here designated ); Parnagua District, Piauhy Prov., Gardner 2733 (syn. K!); Natividade, Goyaz, Gardner 3449 (syn. K!); Pernambuco, at Iguarassa, Ridley, Lea and Ramage s.n. (syn. K!); Burchell 894 (syn. K! sterile); near Rio, San Jos da Laranjeira, Gardner 9829 (syn. K! sterile). DISTRIBUTION : Brazil. The entire, rhombic, subacute to obtuse lip of this species is very characteristic, and similar only to the Central American V. dressleri. It has also the thickened veins in the lip apex and the distichous arrangement of the V. hostmanii group. The species seem to be rare and we are basing our concept of this entity upon Hoehnes published drawing and the lectotype, since other materials are fragmentary or inadequate, and characters of the surface of sepals and ovary, typical of the V. hostmanii group, have not been examined. The information on the lectotype, in Rolfe’s handwriting, was probably obtained from a Gardner’s note in Hooker, London Journal of Botany 1: p. 542; cited in the protologue. The original data are . Vanilla sp. Scentless. Climbing on the stump of an old Tree. Rio Comprido [?] common in the woods about The syntypes Ridley, Lea and Ramage s.n., Gardner 2733 and 3449 probably belong to V. bahiana, Gardner 245, selected here as the lectotype. Burchell and the leaves are very different from the lectotype COLLECTIONS Glaziou 20709 (BM!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 192; Pabst, G.F.J. 1966. Orqudea 28(2-6): 169-170. 39. V. giulianettii F.M. Bailey in Queensland Agric. J. 7: 350 (1900). Type: Papua New Guinea (British New Guinea), 15 Dec. 1909, Giulianetti 1906 (holo. BRI; iso. K!). DISTRIBUTION : New Guinea. This scarcely known species has very large leaves, different from those of the New Guinean . to compare it with the New Guinean vanillas described by Schlechter, which also have large leaves (V. kaniensis, V. kempteriana and V. wariensis). COLLECTIONS : PAPUA NEW GUINEA: Morobe, Gillison NGF 25021 (K!); Western Dist., Galore NGF 33728 (K!); Northern Dist., Carr 17133 (BM!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 221. 40. Lindl., Gen. Sp. Orch. Pl. 435 (1840). Type: French Guiana, Martin s.n. (holo. K-LINDL!).

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla377V. clavicula Duss, nomen. V. lutescens Moq. ex Dupuis, Rev. Hortic. ser. 4, 5: 121, f. 24 (1856). DISTRIBUTION : Vene Peru, and Brazil. that V. pompona (lateral sepals up to 8.4 cm in the type specimen). The sepals are narrower, subacute, the petals less rounded, and the lip has a subquadrate apex, not deeply notched as V. pompona. COLLECTIONS : GUYANA: Smith 2474 (G 7889/122! K!); im Thurm s.n. (K! Pittier 9460 Hart 3 (K!); Broadway 4565 (BM!); Broadway 41912 (BM!); Hance 5285 (BM!). PERU: Klugg 3685 Rosa et al. 5060 (MG! fruit only); Rosa et al. 4915 (MG!); Roraima, Pires et al. 16778 (MG! fruits only). SOURCES : Schweinfurth, C. 1958. Orchids of Peru I. Fieldiana: Botany 30: 43-44 (as V. pompona); Schultes, R.E. 1960. Native Orchids of Trinidad and Tobago: 42 (as V. pompona 1959. Venezuelan Orchids Illustrated 1: 434-435 (as V. pompona); Foldats, E. 1969. Flora de Venezuela. OrchidaceaeV. pompona). 41. V. grandifolia Lindl. in J. Linn. Soc. Bot. 6: 138 (1862). Type: Principe (Prince’s Island), Barter 1981 (holo. K! drawing K-LINDL! tracing W!). DISTRIBUTION : Democratic Republic of the Congo, Gabon, Principe Island. This species is characterized by its large, broadly elliptic to suborbicular leaves, abruptely apiculate of rather thin substance. Floral morphology suggests a close relationship with V. imperialis, with a very large rostellum and a trumpet-shaped, concave lip, which is apiculate at apex (after drawings sent by de Wildeman undulate, entire lip lacking the characteristic apicule at apex. COLLECTIONS : PRINCIPE: Henriquez (K!); Monteiro (K!). Democratic Replublic of the Congo: Louis 2599 10800 (K!) de Wildemann s.n. (K fragment ex Berlin)!. GABON: Le Testu 6077 (BM! K!), 8445 (BM! K!) 9308 (BM! K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Olszewski. 1999. Fl. de Cameroun: 292, t. 103, G-K. V. grandifolia var. lujae (De Wild.) Geerinck = V. imperialis V. gratiosa Griseb. = V. bicolor 42. Rchb. f. in Bonplandia (Hanover) 2: 88 (1854). Types: Malacca, (lectotype W, here designated); Tabong, Malacca, 1844, s.n. (syn. K-LINDL! probable duplicate at AMES 71377!). V. tolypephora Ridl. in Trans. Linn. Soc. London, Bot. 3: 276 (1893). Type: Malaysia, Penang and the Island Pulau Ubin, lying in the strait between Singapore and Johore, and in Selangor, Singapore collector 3924 (holo. BM!). DISTRIBUTION : Peninsular Malaysia, Singapore, Borneo (Brunei), Sumatra. Expected from S. Thailand. Va is a distinct species because the “ball” of hairs instead of the penicillate callus of allied buds are very truncate-rounded. Sterile collections can COLL E CTIONS : MALAYSIA: s.n. (K-LINDL!); (K-LINDL!); (AMES!); Ridley 1969 (BM!); Curtis 2271 (K!); Wray in King 2537 (K!); King 2996 (K!); King 7528 (K!); Maingay 1660 3255 (K!); Scortechini s.n. (K!); Curtis 1179 (K!); Henderson 20378 (K!); Ngadiman in Henderson s.n. (K!); Ahmad S.324 (K! sterile); Sidek bin Kiah S.248 (K!); Whitmore FRI 3706 (K!); Negri Sembilan, FRI 27529 (K!); Evans s.n. (K!); Tahit 1233 (K!); Mondi 289 (K! fruits only); Amin SAN 86237 (K! fruits only); Collenette 643 (K!); Scortechini 1080 (K!); Cheng FRI 27529 (K!); Perlis, Kaboch 15197 (K!). SINGAPORE: Lobb 226 (K!); Curtis (K!); Ridley 3924 (BM!). BRUNEI: Johns 7456 (K!); Wong 974 (K!); Boyce 452 (K!). SOURC E S : Ridley, H.N. 1924. The Flora of the Malay Peninsula: 201; Portres, R. 1954. Le Genre Vanilla et ses Espces: 168-169; Teo, C.K.H. 1985. Native Orchids of Peninsular Malaysia: 107-108. Seidenfaden, a preliminary list, part 1: 58; Seidenfaden, G,. 1978. Orchid Genera in Thailand VI. Dansk Bot. Arkiv. 32(2): 142, 144 .

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.378 LANKESTERIANA Rchb. f. var. ronoensis (Hayata) S.S. Ying = V. somai 43. V. guianensis Splitg. in Ann. Sci. Nat. Bot. s. 2, 15: 279 (1841). Type: Surinam, Splitgerber 409 (holo. P?). ?V. surinamensis Rchb. f. in Ned. Kruidk. Arch. 4: 321 (1859). Type: Based upon same type as V. guinanensis Splitg. V. acuta Rolfe in J. Linn. Soc. Bot. 32: 453 (1896). Type: Surinam, Marowyne, Aug. 1846, Kappler (holo. K-LINDL!). V. latisegmenta Orchid . 8: 2 (1925). Type: Guyana (British Guiana), Upper Rupununi River, near Dadanawa, 29 May 1922, de la Cruz 1404 (holo. AMES 22973!; iso. GH 7850!). DISTRIBUTION : Guyana, Surinam, and Brazil. Vanilla guianensis has been regarded as an obscure species. Splitgerber, in the original description, mixed a species of the membranaceous group with the fragrant fruits of a penicillate species, perhaps . The illustration that appeared in De Vriese, Tuinb.-Fl. 3: 33. 1856, clearly illustrates the mistake, and it is not small bracts for a species of the Membranaceous 3-lobed. Rolfe (1896) recognized Splitzgerber’s mistake and reduced V. guianensis to the synonymy of V. inodora (his concept including specimens that here we recognize as V. inodora, V. mexicana, and V. guianensis) and considered Reichenbach’s proposal of V. surinamensis as an error. However, Rolfe described Vanilla acuta in the same revision, based on a Kappler specimen and cited a Schomburgk illustration at the British Museum from the Berbice River, as probably belonging to the same species. We agree that Kappler’s although it is evident that the original description was based upon an open bud, very damaged by the bracts, and rather chartaceous texture of leaves are very characteristic for a membranaceous species. Later, Ames and Schweinfurth (1925) described V. latisegmenta. Their description is very good and the type well-preserved and with analytical drawings and matches very well with the several specimens listed below from the Guyanas and Amazonia. very close to V. mexicana, but it is distinguished by its much broader lip (45-50 vs. 30 mm broad), which is not as distinctly 3-lobed, but approaching hexagonal with the apical lobe very broadly ovate, or obscurely long and the small, ovate, acuminate bracts. Vanilla martinezii from Guatemala has also a subentire, very broad lip, but in this species the veins are warty in the basal half, and the raceme is longer. In dried specimens the very broad lip in which veins are strongly marked is very distinct. COLLECTIONS Huber s.n. (BM! MG 1864!); Huber 1864 (G 78889/105!); Photo of plant in habitat, Arboretum amazonicum (as V. aromatica Sw.) (K). GUYANA: Schomburgk s.n. (BM NY! photo, K! tracing from drawing at BM).; Kaituma River, M.W.D. 211 (K!); Im Thurm 64 (K!); Jenman 689 (K!); Jenman 1614 (K!); Barkley s.n. (K! painting). SURINAM: Hostmann 71 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. acuta); Werkhoven, M. 1986. Orchids of Suriname: 228, 236 (as V. mexicana Mill.). 44. V. hallei 34: 282 (1998). Type: Gabon, Belinga, N. Hall 3065 (holo. P!). DISTRIBUTION : Endemic to Gabon. This species is closely related to V. africana. basal 1/4, and with papillae in the disc rather than long bristles. 45. V. hamata 55: 563 (1846). Type: Peru, Pavn s.n. (holo. B , photo. AMES 38629! sterile, iso. BM! G 7889/123 124, W!). DISTRIBUTION : Endemic to Peru. V. pompona group, with large tepals (e.g. petals at least 63 8 mm), with a rather smooth lip, a callose, penicillate callus, and a thickened axial cushion in the apical

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla379part of the lip. The lip is trilobed, with the midlobe projecting and perhaps emarginate. However, it seems to be distinct from all the known species in that group, particularly by the widely elliptic, rounded leaves. Another, well-preserved bud from the G specimen is perhaps an Epistephium species, because of its numerous retrorse hairs and lip scarcely fused to the column. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 233; Schweinfurth, C. 1958. Orchids of Peru 1: 42-43. Fieldiana: Botany 30. 46. V. hartii Rolfe in Bull. Misc. Inform. Kew 1899 (151/152): 133 (1899). Type: Trinidad, Cabasterre Arima, Hart 6355 (holo. K!; iso. AMES 67785!). V. leprieurii Portres in Bull. Soc. Bot. France, 98: 94. 1951. Type: French Guyana, Cayenne, 1846, Leprieur s.n., (holo. P not seen, drawing!). DISTRIBUTION : Mexico to Brazil and French Guyana. Vanilla hartii is an interesting species with a weak vegetative habit similar to that of V. bicolor. However, with the structure of the V. planifolia group, with which it has been confused. Vanilla lepriurii is considered here to be a synonym of V. hartii. Portres (1954) stressed that his V. leprieurii was distinct because the nervation in the center of the lip is denser, with the zone before the nervation thicker, more imbricated scales in the penicillate callus, and the apex of the lip more emarginate, apiculate and pleated. Vanilla hartii has not been previously reported from Brazil, but we have a herbarium specimen sequences indicate that they belong to V. hartii. The collections from Costa Rica are very stout. COLLECTIONS : COSTA RICA: Grayum 7998 (MO!). HONDURAS: Standley 52824 Davidse 36251 Britton 2841 (AMES!); Broadway s.n. (AMES! in part K!); Fendler 1007 (BM! K!). DOMINICA: Hodge 387 (MG! sterile). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. leprieurii) and 250-251, Trinidad and Tobago: 39. 47. V. havilandii Rolfe in Bull. Misc. Inform. Kew 1918: 236 (1918). Types: Sarawak, Kuching, Haviland s.n. (syn. K!); Sarawak, Matang, Ridley s.n. (syn. K!). DISTRIBUTION : Borneo. A species somewhat similar to the Javanese V. albida, leaves are rather thin in substance. COLLECTIONS: MALAYSIA: SARA W AK, BROOKE 8181 (G 7889/181!); LAMB SAN 89632 (K!); SANDS ET AL. 3702 (K!); CHAN 927 (K!, DRA WING). INDONESIA: KALIMANTAN, AFRIASTINI 310 (K!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 209-210. 48. V. helleri A.D. Hawkes in Phytologia 14 (1): 19, t. 13. (1966). Type: Nicaragua, Dept. Chontales, 2 miles S of La Libertad on the road to Santo Toms, 1900 ft, April 1962, Heller 7946 (holo. AMES not located; iso. SEL!). DISTRIBUTION : Nicaragua and Costa Rica. This is one the rarest of the Central American Vanilla distinct 3-lobed lip, in which the midlobe is covered with retrorse, complanate, elongate trichomes, resembling in some way the South American V. ribeiroi. It is a member of the V. odorata-V. insignis complex, also with retrorse, orange trichomes on the midlobe. COLLECTIONS : COSTA RICA, (F!). SOURCES : Hamer, F. 1984. Ic. Pl. Trop. Pl. 1192; Hamer, F. 1990. Orch. Centr. Amer. Selbyana 11: 847. 49. V. heterolopha Summerh. in Bull. Misc. Inform. Kew 1938: 149 (1938). Type: Gabon, Upper Ogou River, Matoro, Oct. 1929, Le Testu 7494 (holo. P!; iso. BM! K!). DISTRIBUTION : Gabon and Congo. This species is a member of the V. africana group, resembling vegetatively V. africana, but it is easily distinguished by its truncate, rectangular, sharply recurved midlobe of the lip, and warts and papillae in the lip appears slightly broader towards the apex. In the other members of the V. africana group, the apical

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.380 LANKESTERIANAlobe of the lip is acute to acuminate, with retrorse, COLLECTIONS : GABON: Le Testu 5769 (BM! K drawing!); Le Testu s.n. (K!); Le Testu 7494bis (BM!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Olszewski. 1999. Fl. de Cameroun: 282, pl. 100, G-M. V. hirsutaV. tahitiensis 50. V. hostmannii Rolfe in J. Linn. Soc. Bot. 32: 462 (1896). Type: Surinam, Hostmann 306 (holo. K!, iso.K-LINDL!). var. gigantea Hoehne, Com. Lin. Tel. Est. M.G. Amazonas, Bot. 1: 217 (1910). V. barrereana Hist. Nat., B, Adansonia 16 (2-4): 220 (1995). Type: French Guiana, Cremers 4523 (holo. P, iso. CAY). V. porteresiana Natl. Hist. Nat.,B, Adansonia 16 (2-4): 219 (1995). Type: French Guiana, Oldeman 2087 (holo. P). DISTRIBUTION : Colombia, Guyana, French Guiana, Surinam, Venezuela and Brazil. Vanilla hostmanii is a member of a small group of species that includes also V. cribbiana, V. dressleri, and V. weberbaueriana, characterized by the conspicuously distichous arragement of the basal bracts, its entire to obscurely trilobed lip adorned with thickened, papillose veins on the lip apex, and with the ovary, sepals and the petals keels oddly granulose. racemes, larger leaves, a more thickly veined lateral Vanilla barrereana and V. porteresiana, described thickened veins and granulose sepals surface but have smaller measurements and different lip tips. We have compared them with immature buds of V. hostmanii and we believe than they are better considered synonyms of V. hostmanii. Several specimens of V. hostmanii have been annotated as V. planifolia var. gigantea, but we have been unable to locate the type of V. hostmanii or of (= V. pompona sensu Hoehne), as stated by Hoehne in 1945. Vanilla hostmanii seems to be a common species in Guyana and the Amazon. COLLECTIONS : GUYANA: Anderson 71 (K!); Jenman (K!); im Thurn 65 (K!). COLOMBIA: Lehmann H.K. 1187 (K!); Schultes 3809 (AMES!). Prance 7955 (MG!).; Para, Wullschlaegell s.n. (W!); Prance et al. P25739 (K! MG!); (MG!). Amazonas, Silva Adderley 43074 43170 (VEN!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 193; Hoehne, F.C. 1945. Flora Brasilica 12, 2: Orchidaceae: 18. 51. V. humblotii Rchb. f. in Gard. Chron. n.s. 23: t. 726 (1885). Type: Grande Comore, Humblot 1413 (holo. W!, iso. BM! K! P!). DISTRIBUTION : Comoros Islands. Another member of the V. roscheri group marks in the throat of the lip. COLLECTIONS : COMOROS: Mayotte, Labat 2905 (P, K!). SOURCES: Rolfe, R.A. 1905. Bot. Mag. t. 7996; DuPuy, Orchids of Madagascar: 207. 52. V. imperialis Kraenzl. in Notizbl. Bot. Gard. Berlin-Dahlem 1: 155, t.1 (1896). Type: Cameroon, Yaound, (holo, B). V. lujae De Wild., Belg. Colon. 10: 28 (1904). Luja s.n. (holo. BR). V. grandifolia Lindl. var. lujae (De Wild.) Geerinck in Bull. Soc. Roy. Bot. Belgique 107, 1: 58 (1974). V. imperialis Kraenzl. var. congolensis De Wild. in Rev. Hist. Nat. Appl. (Paris) 2: 188 (1921). Type: Mortehan 244 (holo. BR). DISTRIBUTION : Sierra Leone, Ivory Coast, Ghana, Nigeria, Cameroun, Democratic Republic of the Congo, Angola, Uganda, United Republic of Tanzania, Kenya. This common African species has the most white, heavily veined-blotched with deep rose to resembling some species of Costus, however, many specimens from East Africa have shorter bracts. It is

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla381closely related to V. ochyrae and to V. polylepis (see comments under the species). COLLECTIONS : CAMEROUN: Bates 1523 (BM!). GHANA: Westward 113, 226 (K!); Vigne 2412 (K!); Hall GC 35391 (K!). SIERRA LEONE: Assi 9041 (K!); Jaeger 40 (G 78889/184 185!); Jaeger 16566 (K!). NIGERIA: Chapman 4869 (K!). UGANDA: Eggeling 2062 (K!); Dmmer 2672 (K!), 2672A (BM!) Eggeling 1065 (K!); Ussher 83 (K!); Poulsen et al. 971 (K!); (BM!); van Someren E.A.H Moreau 429 (K!); Greenway 6664 (K!); Greenway 2928 (K!). ANGOLA: Rep. de Agricultura 14214 (K!); Gossweiller 13837 (BM!). SOURC E S : Rolfe, R.A. 1897. Flora Trop. Afr. 7: 3: 454; Portres, R. 1954. Le Genre Vanilla et ses V. lujae ); Summerhayes, V.S. 1968. Flora of West Tropical Africa 3(1): Orchidaceae: 205-206; Piers, 1969. Flora of Tropical East Africa. Orchidaceae 2: 259261; Geerinck, D. 1984. Flore D’Afrique Centrale Olszewski. 1999. Fl. de Cameroun: 290-291, pl. 103, A-E. 53. V. inodora Schiede in Linnaea 4: 574 (1829). Type: Mexico, Schiede s.n. (holo. BM!, iso. LE, W!). V. preussii Kranzl. in Notizbl. Bot. Gard. BerlinDahlem 7(67): 320 (1919). Type: Guatemala, Preuss 1445 (holo. B). V. pfaviana Rchb.f. in Gard. Chron. n.s. 20: 230 (1883). Type: Costa Rica, Pfau s.n. (holo. W!). DISTRIBUTION : Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica and perhaps Panama. This is the species of the membranaceous group with most northern distribution. It is distinguished by the high, thick cushion-like, axial callus of its emarginate lip. COLLECTIONS : MEXICO: Liebman 297 (K!); Liebman 284-296 (BM!); Liebman 7268 (BM!); Calzada 01031 (BM!). Veracruz, Nagel in Oestlund 2683 (AMES!); Purpus in Oestlund 4876 (AMES!); Purpus in Oestlund 5045 (AMES!); Purpus 8481 (AMES!). Puebla, Arsne 1682 (AMES!).; Guerrero: Nagel in Oestlund 1984 (AMES!), 1985 (AMES! BM!).; Chiapas: Matuda 17912 Gentle 1672 (AMES!). GUATEMALA: Steyermark 49171 (AMES!). HONDURAS: (AMES!). COSTA RICA: Skutch 2592 (AMES! K!); Lankester K331 (K!); von Wedel 1423 (BM!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 252-253 (mixed with V. mexicana) and 257 (as V. pfaviana). 54. V. insignis Type: Honduras, Edwards 407 (holo. AMES 40085!). DISTRIBUTION : Mexico, Guatemala, Belize, Honduras, reported by Dressler (1993) for Costa Rica. This common Mesoamerican species has rarely green tepals and a cream-colored lip bearing orangeochre retrorse papillae are characteristic. Sterile plants can be distinguished by their sulcate stems with minutely papillose surface. It is a member of the V. odorata group, but it has shorter leaves and more xerophytic habit. COLLECTIONS : MEXICO: Oaxaca, Nagel in Oestlund 4584 (AMES!).; Veracruz, Nagel in Oestlund 2682 (AMES!); Purpus in Oestlund 4866 (AMES!); Purpus in Oestlund 5961 (AMES!).; Quintana Roo, Nagel in Oestlund 4973 Gentle 1063 (AMES!). GUATEMALA: Steyermark 45233 (AMES!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Guide to the Orchids of Costa Rica and Panama. 55. V. kaniensis Schltr. in Repert. Sp. Nov. Regni Veg. Beih. 1: 32 (1911). Type: Papua New Guinea, Schlechter 17634 (holo. B; published drawing!). DISTRIBUTION : Bismarck Mountains, Papua New Guinea. Similar to V. kempteriana and V. wariensis, but with smaller (20-25 4-7 cm) leaves. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 221; Howcroft, N.H.S. 1984. A Guide to the monocotyledons of Papua New Guinea. R.J. Johns Forestry Department PNG, University of Technology. Papua New Guinea. SNP Ltd. p. 9.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.382 LANKESTERIANA56. V. kempteriana Schltr. in Repert. Sp. Nov. Regni Veg. Beih. 1: 32 (1911). Type: New Guinea, Schlechter 18045 (holo. B ; published drawing!; iso.G!). DISTRIBUTION : Papua New Guinea, between Lae and Madang. Distinguished from other species from the island by its rather broad leaves. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 220-221; Howcroft, N.H.S. 1984. A Guide to the monocotyledons of Papua New Guinea. R.J. Johns Forestry Department PNG, University of Technology. Papua New Guinea. SNP Ltd. p. 9. 57. V. kinabaluensis Carr in Gard. Bull. Straits Settlem.8: 176 (1935). Types: Borneo, Carr 3157 (syn. K!); Kadamaian (Tampassuk) River, at the entrance to Menetendok Gorge, c. 2,700 feet, March 1933, Clemens 26300 (syn. BM!); Dallas, 3,000 feet, Aug. 1931; Clemens 26725 (syn. BM! K!); Dallas, 3,000 feet, 15 Sept. 1931, Clemens (syn. K). DISTRIBUTION : Borneo (Sabah) and Peninsular Malaysia. Very similar to the Bornean J.J. lip with a median keel and without irregularly seriate warts. The column lacks the appendages found in V. . It is recognized by its extremely long open successively perhaps during much of the year. COLLECTIONS : MALAYSIA: Sabah, SFN-25273 (K!); (K!); Wood 909 (K!, and drawings); Lamb 1230/90 (K!); Lugas 1724 (K!); Burbidge s.n. (BM! K drawings!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 216; Holttum, R.E. 1964. Flora of Malaya. Orchids: 96-97. V. klabatensis Schltr. nomen nudum. V. latisegmenta V. guianensis V. laurentiana De Wild. = V. crenulata V. leprieurii Portres = V. hartii V. lindmaniana Kraenzl. = V. palmarum V. lujae De Wild. = V. imperialis V. lutea C.H. Wright ex. Griseb. = V. palmarum V. lutescens Moq. ex Dupuis= 58. V. madagascariensis Rolfe in J. Linn. Soc. Bot. 32: 476 (1896). Type: Madagascar, Bomatoe Bay, Bojer s.n. (holo. K!). DISTRIBUTION : Madagascar. Vanilla madagascariensis is very similar to V. phalaenopsis from the Seychelles and V. roscheri from pink to apricot throat to the lip, but they differ in the lip lobing. They have some of the most spectacular Vanilla. SOURCES 42B, 43C. V. majaijensis Blanco = ?V. planifolia V. marowynensis Pulle = V. appendiculata 59. V. martinezii Soto Arenas, ined. DISTRIBUTION : Guatemala, expected from Belize and Honduras. V. martinezii is member of the V. mexicana group. The most similar species is V. guianensis, however in V. guianensis the callus is formed by 5 basal keels and 3 rather prominent ones, but not or almost not verrucose, and the lip apex is trilobed. V. martinezii seems to be endemic of the wet lowlands around Lake Izabal. 60. V. methonica 97 (1854). Type: Colombia (New Granada), 6-8000ft., Warscewicz s.n. (holo. W19363!; iso. K-LINDL!). DISTRIBUTION : Colombia and Peru, apparently only on the eastern slopes of the cordillera. We have seen very few specimens referable to this species, and the type is poorly preserved. However, concept with modern specimens from Peru. membranaceous group), with small, broadly ovate in which the lateral lobes are oblong and rounded and the emarginate midlobe are characteristic. Like many others in the V. mexicana group, it has 3 longitudinal keels, converging on the lip apex, but the keels are thickened in the distal part and somewhat rugose. It

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla383seems to be more closely related to V. inodora, but it lacks the characteristic cushion-like, massive callus of that species. V. oroana Ecuador, has very large foliaceous bracts, the tepals are strongly contorted, and it has 5 apical keels on the acute lip; no other membranaceous vanilla is known from Andean South America. COLLECTIONS : COLOMBIA: Barragn in Lehmann HK1181 (K!); Goudot s.n. (W sketch!). PERU: Smith 3883 (MO!); Vargas 18697 (AMES!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 254-255. 61. V. mexicana Mill., Gard. Dict. (ed. 8), no. 1 (1768). Type: Haiti, Plumier s.n. (holo. Plumiers original drawing at P; reproduced in Plumier, Nov. Pl. Amer. Genera 25, pl. 28, 1703!). Epidendrum vanilla L., Sp. Pl. 2: 952 (1753). Type: “Volubilis siliquosa plantaginis folio” in Catesby, Nat. Hist. Carolina 2 (?3): app. t. 7. Jarvis, Taxon: 48(1): 47. 1999). V. aromatica Sw. in Nova Acta Regiae Soc. Sci. Upsal. 6: 66 (1799). Type: based on same type as Epidendrum vanilla L. V. anaromatica Griseb., Fl. Brit. W. Ind . 638 (1864). Type: Based upon Plumier, Ed. Burm., t. 188. V. epidendrum Mirb., Hist. Nat. Pl. 9: 249 (1802-6). V. vanilla (L.) Huth in Helios 11(9): 136 (1893). DISTRIBUTION : U.S.A. (Florida), Cuba, Puerto Rico, Hispaniola, Lesser Antilles, Trinidad and Tobago, perhaps N. South America (Venezuela). Reported repeatedly from Guianas and Brazil, but the examined material belongs to V. latisegmenta or other taxa. The taxonomic history of this species is very confusing. Mexican specimens referred to this species belong to V. inodora. Its lip is 3-lobed, the midlobe is well-developed and can be subacute to rounded. It is closely related to V. costaricensis with a truncate lip, to V. ovata and V. martinezii the latter with many warts near the base of the lip. Some authors have considered V. inodora as the valid name of this species. Although the type of V. inodora from the original area to establish that it is the species common in Mesoamerica that has a thick, longitudinal cushion-like callus and is widely known as V. pfaviana. The West Indies taxon with a callus formed by 3 longitudinal keels it is unknown in Mexico. COLLECTIONS : PUERTO RICO: Sintenis 1739 (K!); Sintenis 2456 (AMES!); Sintenis 2486 (G 7889/104, /107!); Sintenis 5469 (AMES! BM! G 7889/128! K!); Eggers 1076 (G 7889/ 178!); Eggers 1322 (K!); Kevorkian 6468 (AMES!). CUBA: Wright 3353 (AMES! K!); (AMES!); Clemente 2401 (AMES!); (AMES!); Schafer 3211 (AMES!), 3299 7982 (AMES!); JAMAICA: Purdie s.n. (K!); Morris 451 (K!); Wood s.n. (BM tracing!). TRINIDAD AND TOBAGO: Baker 14732 (K!); (K!); (AMES!); Broadway 3874 (K!); Crueger s.n. (K!); Freeman s.n. (AMES!); Simmonds 334 (AMES!); Britton 2945 (AMES!); (AMES!). GUADELOUPE: Smith 10350 (AMES!). DOMINICA: Immary 138 (K!); Ramage s.n. (K!); W.H. and B. T. Hodge 3671 (AMES!); Ernst 1462 (AMES!). ST. LUCIA: Proctor 17710 (AMES!). DOMINICAN REPUBLIC: Eggers 2086 (K!). HAITI: Eyerdam 395 (AMES!); Garca 42003 (AMES!). MARTINIQUE: Duss 2081 (AMES!). SOURCES Cuba I: 351 (as V. inodora); Portres, R. 1954. Le Genre Vanilla et ses Espces: 252-253 (mixed up and as V. inodora); Schultes, R.E. 1960. Native Orchids of Trinidad and Tobago: 40 (as Vanilla inodora); Kenny, J. 1988. Native Orchids of the Eastern Caribbean: Orqudeas Cubanas: 21 (as V. inodora); Dunsterville, Illustrated 1: 432-433 (as V. inodora); Foldats, E. (as V. inodora); Ackerman, J.D. 1995. An Orchid Flora of Puerto Rico and the Virgin Islands: 174; Cafferty Epidendrum vanilla. V. microcarpa H. Karst., Deut.sche Fl. 458 (1880-83). Type: Venezuela Poorly known species. V. montagnacii Portres in Bouriquet, Vanillier et Vanille 46: 282 (1954), nom. inval. Based upon:

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.384 LANKESTERIANAMadagascar, P.R. Montagnac 146 (not seen) This species has an entire lip with two subpapillose keels on its disc. It is possibly a semi-peloric form of V. decaryana or V. madagascariensis which are both found in the region. Portres failed to provide a Latin description of this name which is consequently invalid. COLLECTIONS : MADAGASCAR: N of Morondava, Bosser 18187 (P!); Abraham 78 (P!). 62. V. montana Ridl. in J. Fed. Malay States Mus. 6: 58 (1915). Type: Malaya, Perak, Gunong Kerbau, 4400 5500 ft, 19 March 1913, Robinson s.n. (holo. K!; iso.AMES 25073! BM!). DISTRIBUTION : Malaysia. Seidenfaden considered this species to be V. albida, but thought by us to be V. yersiniana. However, it differs from V. yersiniana by the much longer hairs at the lip apex (up to 4 mm long) and much broader leaves. It seems to be a rare plant and as far as we know it is only known from the type specimen. SOURCES : Ridley, H.N. 1924. The Flora of the Malay Peninsula: 201; Portres, R. 1954. Le Genre Vanilla et ses Espces: 222; Holttum, R.E. 1964. Flora of Malaya, ed. 3. Orchids: 97. 63. V. moonii Type: Sri Lanka, Thwaites 3204 (holo. K!; iso. AMES 71375! BM! G!). DISTRIBUTION : Endemic to Sri Lanka. This species has 2 rows of papillae at the apical lobe of the lip, in the way of many other Asian species. thick rachis is also characteristic. COLLECTIONS : SRI LANKA: Wight s.n. (K!); Champion (AMES! K-LINDL!); collector 1869 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 182; Jayaweera, D.M.A. 1981. Orchidaceae. A Revised Handbook to the Flora of Ceylon 2: 333. 64. V. nigerica Rendle, Cat. Pl. Oban 108 (1913). Type: S. Nigeria, Oban, 1911, Talbot 776 (holo. BM!). DISTRIBUTION : Nigeria and Cameroon. A species of the V. chalotii group. It is similar to V. seretii, both being characterized by their narrowly oblong leaves, verrucose stems and pretty, large, white of the V. chalotti group. Vanilla nigerica is different from V. seretii in its lip being provided with 2 gibbae at base, the coarsely hairy lip throat, and the recurved, not rolled midlobe of the lip. It is probably a rare plant. COLLECTION : CAMEROON: Bates 1263 (BM!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 172; Summerhayes, V.S. 1968. Flora of West Tropical Africa 3(1): Orchidaceae: 205-206. 65. 34: 291 (1998). Type: Cameroon, Jacques-Flix 4727 (holo. P!; iso. K!). DISTRIBUTION : Cameroon. A species closely related to V. imperialis, but distinguished by it clearly trilobed lip, with a semiorbicular midlobe larger than the larger lateral lobes and the margins undulate. The lamellae and trichomes of the lip are branched. COLLECTION : CAMEROON: Endenguele 113 (K!). 66. V. odorata C.Presl, Reliq. Haenk. 1(2): 101 (1830). Type: Ecuador, Haenke s.n. (holo. PR!; iso. PR!). V. ensifolia Rolfe in Bull. Misc. Inform. Kew 1892(65/66): 141 (1892).Type: Colombia, Goudot s.n. (holo. K!). V. uncinata Huber ex Hoehne in Arquiv. Inst. Biol. (S. Paulo) 8: 269 (1937). Type: Par, Brazil, Goeldi 7392 (holo. MG!). V. denticulata Pabst, 23rd Congr. Nac. Bot., Garanhuns: 109 (1972). Type: Brazil, Ferreira 37-67 (holo. HB). DISTRIBUTION : Mexico, Guatemala, Costa Rica, Panama, Colombia, Peru, Bolivia, Brazil. Vanilla odorata is widely distributed in continental tropical America, from Mexico to Bolivia. It is easily distinguished by its narrowly lanceolate, long acuminate leaves; although specimens grown in shaded spots may have broader leaves. Its fruits are strongly fragrant and are collected from wild plants in many areas of Latin America. It is occasionally grown in Oaxaca, Mexico, as source of vanilla fragrance. Vanilla denticulata and V. uncinata seem to dimensions falling within the known range of V. odorata, although the lip of V. uncinata is slightly emarginate. is also very closely

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla385and more abundant material from Amazonia and the Guyanas is necessary to be compared carefully with specimens of V. odorata from other areas. COLLECTIONS Whitefoord 1316 (BM!). COSTA RICA: (AMES!). GUATEMALA: Aguilar 164 (AMES!). PANAMA: Hunnewell 16433 (AMES!). COLOMBIA: Lehmann H.K. 1183 (K!). PERU: (AMES!); Williams 3161 (AMES!). BOLIVIA: White 1115, 1821 (AMES!); Spencer Moore 637a (BM!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Orchids of Peru I. Fieldiana: Botany 30: 43; Dodson, H.H. Gentry. 1985. La Flora de Jauneche: 110, t. 44A; Garay, L.A. 1978. Flora of Ecuador. Orchidaceae 67. V. organensis Rolfe in J. Linn. Soc. Bot. 32: 452 (1896). Types: Brazil, Organ Mountains, Gardner 632 (lecto. K-LINDL!, here designated; isolecto. AMES 71366! G! K! W 19645!) Miers (syn. BM! K!); Near Rio de Janeiro, Glaziou 11620 14320 (syn. K!). A membranaceous Vanilla which seems to be closely related and perhaps not distinct from V. parvifolia is very variable in other membranaceous vanillas. V. edwallii and V. bertoniensis are allied, but both have a long acuminate lip. SOURCES : Portres, R. 1954. Le Genre Vanilla et Orqudeas do Estado da Guanabara. Orqudea 28(26): 169-170. 68. V. oroana Dodson in Orquideologia 22(3): 210 (2003). Type: Ecuador, Prov. El Oro, 3.7 km N of (holo. AMES!; iso. K! (sterile) MO!). DISTRIBUTION : Known only from the type collection in western Ecuador. Vanilla oroana is a member of the membranaceous foliaceous bracts up to 110 65 mm. The large with papillae and the apex is 5-keeled. Its sepals and petals are green, and its lip white, with yellow-orange lamella on apical lobe. Its nearest relatives seem to be the members of the V. mexicana complex. V. ovalifolia Rolfe = V. zanzibarica 69. V. ovalis Blanco, Fl. Filip. 448 (1845). Type: Philippines, Mt Majaijai, Lilio, Azaola s.n. (not located). Epidendrum vanilla sensu Blanco, Fl. Filip. (ed. 2) 643: (1837), non L. V. majaijensis Blanco, loc.cit. 593 (1845). V. philipinensis Rolfe in J. Linn. Soc. Bot. 32: 459 (1896). Type: Philippine Islands, without precise locality, Cuming 2132 (holo. K!; iso. BM!). DISTRIBUTION : The Philippines (Luzon, Mindanao, Leyte, Samar and Sibuyan Islands). This is the only leafy Vanilla in the Phillippines. Its sepals and petals are greenish-cream and the lip is marked with purple lines. It is apparently related to V. platyphylla from Sulawesi and to the New Guinean species, but it is distinguished by its very hairy lip apex. COLLECTIONS : Cuming (K-LINDL!); Luzon, M. Perrotet s.n. (G 7889/103!); D.E. Elmer 16035 (BM! G 7889/110! K!); A.D.E. Elmer 9259 (BM! K!); H.G. Gutirrez 78082 (K!); A. Loher 5420 (K!), 5421 (K!) 5422 (K!); J.S. Serrato s.n. (BM! K!); Irosin 16035 (BM!); Merrill 665 (BM! K!); Sibuyan, D.A. Madulid 6875 (K!). SOURCES : Ames, O. 1908. Orch. 2: 47 (as V. philippinensis); Merrill, 1914. Species Blancoana: 459; Portres, R. 1954. Le Genre Vanilla et ses Espces: 182-183; Valmayor, H.L. 1984. Orchidiana Philippiniana 1: 322. 70. V. ovata Rolfe in J. Linn. Soc. Bot. 32: 451 (1896). Type: Guyana, Martin s.n. (holo. BM, tracing at K!). DISTRIBUTION : Venezuela and Guyana. This is perhaps not different from V. mexicana. We have not located the type specimen, and the rough tracing at K is inadequate to be sure. Illustrations V. mexicana from South America (e.g.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.386 LANKESTERIANArecognition. Another specimen illustrated by Hoehne V. ovata has very slender segments and narrow midlobe of the lip. We cannot be sure of the identity of the South American material until specimens have become available to us. COLLECTIONS J.A. Steyermark 87393 (G 7889/130!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Garay. 1959. Venezuelan Orchids Illustrated 1: 432; Hoehne, F.C.1945. Flora Brasilica t. 29. 71. V. palembanica Tijdschr. Ned. Indie 29: 243 (1867). Type: Sumatra, Teijsman (holo. L). DISTRIBUTION : Sumatra. Another scarcely known species that could be related to group. SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 167. 72. V. palmarum (Salz. ex Lindl.) Lindl., Gen. Sp. Orch. Pl. 436 (1840). Type: Brazil, Baha, Salzmann s.n. (holo. K-LINDL! iso. K! G 78889/134! LE!). Epidendrum palmarum Salzmann ex Lindl., Gen. Sp. Orchid. Pl. 436 (1840). V. lutea C.H.Wright ex Griseb., Cat. Pl. Cub. 267 (1866). Type: Cuba, Wright s.n. (holo. BM?). Kraenzl. in Kongl. Svenska Vetenskapsakad. Handl. 46(10): 17 (1911). Type: Brazil, Matto Grosso, near Palmeiras, Exp. I Regnell, Lindman 2481 (holo. S). DISTRIBUTION : Brazil, Venezuela, Guyana, Suriname, Bolivia, Peru. Mostly in the Amazonian basin. Vanilla palmarum is different from other American Vanilla species in lacking a penicillate callus on the lip, and in having its column united to the lip for only about a half of the column length. In other species the penicillate callus is necessary during the pollination, but V. palmarum seems to be autogamous. American vanillas. It forms infrutescences with many thick fruits with a conspicuous calyculus. Weak plants are sometimes confused with those of V. bicolor. It is repeatedly reported as growing on palms (Mauritia, Orbignya, Attelia, Scheelea, Syagrus), and can be very abundant. The mature fruits are apparently odour-less. Vanilla lindmaniana has been treated as an obscure, doubtful species. However, its habitat, habit, wooly column, simple ecallose lip, and concolorous yellow V. palmarum. Considering the published drawings, Vanilla palmarum is rather variable in the form of the lip. COLLECTIONS : SURINAME: Hostmann (K-LINDL!). Gardner 1419 (BM! K!). Maranhao, Vieira et al. 1545 (CEN 013782!). Tocantins, Walter et al. 405 (CEN 026289!). Bahia: Curran 402 (AMES!); Blanchet s.n. (G 7889/137,140!); Blanchet 139 (G 7889/138!); Vignier s.n. (G 7889/139!); Giacometti 878 (CEN 15182!). Goias, Walter et al. 4100 (CEN 32308, s.n.!), 4031 (CEN 32254!), 3698 (CEN 31844!), 2044 (CEN 024613!), 1274 973 (CEN 15626!). Mato Grosso, 26301 (CEN 3407, s.n.!); Argent et al. 6349 (K!). Pernambuco, Ridley et al. (BM!) E. Menezes 18 (K!). Para, (MG!); da Silva et al. 1959 (MG!). Amazonas, Ribeiro et al. 1333, 1963 (K!). Acre, Prance et al. 7955 (K!). BOLIVIA: White 935 (AMES!), 1512 (AMES!), 2332 2366 (AMES!); White s.n. (AMES 48244!) SOURCES : Martius, F.P., 1896. Fl. Brasiliensis 3: pt. 4: 152. 1896; Portres, R. 1954. Le Genre Vanilla Venezuelan Orchids Illustrated 6: 438-439. Dodson, V. chamissonis). V. parishii Rchb. f. = V. aphylla 73. V. parvifolia Barb. Rodr., Orch. Nov. 2: 271, pl. 36 (1881). Type: Brazil, Paran Province, near Curitiba, Dec., Rodrigues s.n. (holo. RJ) V. perexilis Bertoni in An. Ci. Parag. ser. 1. 8: 8 (1910). Type: Paraguay, Puerto Bertoni, Bertoni 3631 (not seen). DISTRIBUTION : Paraguay and Brazil (Paran). Vanilla parvifolia has a number of closely related species that must be studied carefully when more material become available. It is distinguished from the closely related V. angustipetala (syn. V. rojasiana), the papillae or warts on the disc.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla387Johnson (unpublished manuscript) stated that V. perexilisr orm of V. parvifolia. SOURCES : Martius, F.P. 1896. Fl. Brasiliensis 3: pt. 4: 151, t. 33. 1896; Portres, R. 1954. Le Genre Vanilla (unpublished). Las especies argentinas del gnero Vanilla. Dressler = V. trigonocarpa 74. V. penicillata Ill. 3: 324 (1965). Type: Venezuela, Ro Carrao, Dunsterville 787 (holo. AMES). DISTRIBUTION : Venezuela and Brazil. Vanilla found in the South America. It was described from a specimen with immature buds. Flowering material has larger with greenish-yellow sepals, white petals and a white lip with a yellow center (DS = 57 11 mm; LS = 57 12 mm, P = 57 11 mm, L = 53 32 mm, column = 42 mm long; ovary = 30 3 mm). It differs from purplish markings, the very long trichomes on the apex of the lip (up to 1 cm long), and the laceratethe very long column suggest that it is probably not V. ribeiroi, which also has reduced, xerophytic leaves. Vanilla penicillata grows in white sand areas and its stems are less than 5 mm thick. COLLECTIONS Dunsterville 786 (K drawing!), 1372 (K published drawing!); Steyermark (K!). SOURCES : Foldats, E. 1969. Flora de Venezuela. V. perexilis Bertoni = V. parvifolia 75. V. perrieri Schltr. in Repert. Sp. Nov. Regni Veg. 33: 114 (1924). Type: Madagascar, Ankarafantsiba, H. Perrier 1851 (holo. P!). DISTRIBUTION : Endemic to Madagascar. Vanilla is similar to the V. madagascariensis with a dark brownish throat of the lip. Its closest relative is V. humblotii. SOURCE Hermans. 1999. The Orchids of Madagascar: 207-208, pl. 42A. V. pfaviana Rchb. f. = V. inodora 76. V. phaeantha Rchb. f. in Flora 48: 274 (1865). Type: Cuba, C. Wright 3351 (holo. W; iso. AMES 71001, mixed with cf. V. poitaei! BM (another specimen is V. cf. poiteai)! G 7889/109! K!). Vanilla planifolia var. macrantha Grisebach, Cat. Pl. Cub. 267 (1866). Type: Cuba, Wright 3351 (holo. BM?). DISTRIBUTION : U.S.A. (Florida), Cuba, Jamaica, Lesser (Yucatn). A common leafy species in the Caribbean area. The oblong-elliptic, xerophytic, leaves are usually with greenish tepals, and the lip is white with yellow stripes. It is related to V. bahiana, from Brazil. From the V. planifolia and V. odorata complexes can be distinguished by its not warty disc. COLLECTIONS : U.S.A.: Florida: Eaton 1129 (BM! K!). CUBA: (AMES!); Singleton 862 (AMES!); Singleton s.n. (AMES!); (AMES!); Jack 4775, 5474 (AMES!); Grey (AMES!); (AMES!); (AMES!). Rorer s.n. (AMES!); (AMES!); Baker 14733 (K!). JAMAICA: Harris s.n. (BM!); Fawcett 2 (K!). ST. VINCENT: Guilding s.n. (K!); 1336 (K!). SOURCES Cuba 1: 351; Portres, R. 1954 Le Genre Vanilla et ses Espces: 242-243; Schultes, R.E. 1960. Native Orchids of Trinidad and Tobago: 40; Kenny, J. 1988. Native Orchids of the Eastern Caribbean: 23; Britton, las Orqudeas Cubanas: 22. Luer, C.L. 1975. Native Orchids of North America: 70, pl. 18. 77. V. phalaenopsis Rchb. f. ex van Houtte, Fl. Serres 17: 97, t. 1769-1770 (1867-1868). Type: Seychelles, Horne 607 (holo. W! iso K!). DISTRIBUTION : Seychelles.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.388 LANKESTERIANA This species is scarcely different from V. roscheri from continental Africa and the nearby islands. The appearence is exactly the same, however the base of the lip is said to be much more pubescent than in V. madagascariensis. COLLECTIONS : SEYCHELLES: Jeffrey s.n. (K photo!); Schlieben 16692 (K!); Thomasset s.n. (K!); Fosberg 51988 (K!); (K!); Zelia 375 (K!). SOURCE : Reichenbach, H.G. 1877. Orchidaea Roezlianae Novae. Linnaea 41: 65-67. V. philippinensis Rolfe = V. ovalis V. pierrei Gagnep. = ?V. siamensis V. pilifera Holttum = V. borneensis V. pittieri Schltr. = V. pompona 78. V. planifolia G. Jackson in Andrews, Bot. Repos. 8: t. 538 (1808). Type: West Indies, cult. Greville (illustration cited here, lecto. selected by Garay, Fl. Less. Antill. 44. 1974). Epidendrum rubrum Lam., Encycl. 1, 1: 178 (1783), nomen conf. V. aromatica Willd. Sp. Pl. (ed. 4): 121 (1804), nomen. Myrobroma fragrans Salisb., Parad. Lond. 2(1): t. 82 (1807). Type: Illustration cited here. Blume, Bijdr. 422 (1825). Type: Cult. Buitenzorg BG (holo. not seen). V. sylvestris Schiede in Linnaea 4: 573 (1829). Type: Mexico, Schiede s.n. (holo. BM!). V. sativa Schiede in Linnaea 4: 573 (1829). Type: Misantla, Schiede s.n. (iso. BM!). V. duckei Huber, Bol. Mus. Paraense Hist. Nat. 5(2): 327 (1909). Types: Brazil, Amazonas, A. Ducke 3079 3489 (syn. MG). V. domestica Druce, Rep. Bot. Exch. Club 3: 425. 1913 (1914). V. rubra (Lam.) Urban in Repert. Sp. Nov. Regni Veg. Beih 5: 157 (1920), nomen conf. V. fragrans (Salisb.) Ames, Sched. Orch. 7: 36 (1924). V. bampsiana D. Geerinck in Bull. Jard. Bot. Belg. Leopold II, Sept. 1930, Lebrun 1459 (holo. BR!; iso. K!). V. majaijensis Blanco, Fl. Filip. (ed. 2) 593 (1845). Type. Philippines, Mt. Majaijai, Azaola s.n. (holo. not located). DISTRIBUTION : Native of Mexico, Guatemala, Belize, Honduras, Costa Rica, and Panama. Apparently New Guinea, and perhaps in many other tropical areas. This species is the cultivated source of the vanilla essence. Its natural range probably ran from Central Veracruz in Mexico to Costa Rica or Panama, but it is now found in many tropical areas as an escape. All the material we have seen from Africa, the West Indies, South America (including Ecuador), and most of Costa Rica and Panama, is indistinguishable from the cv. “Mansa” from Veracruz, and different from wild specimens from Chiapas, Belize and Honduras which have characteristically broad, elliptic leaves. We have not examined the syntypes of V. duckei Huber, but follow Hoehne (1954) who considered it a synonym of V. planifolia. The material of V. carinata Rolfe is very inadequate, and we are not completely V. planifolia. This species has several allies with which it can be confused, particularly V. hartii, V. insignis, V. odorata and V. tahitiensis (see comments under those species). COLL E CRTIONS : MEXICO: Pavn (actually Sess and s.n. (G 7889/150!); without collector (G 7889/106!); Schott s.n. (BM!); Exploradora 7725 (AMES!); Gonzlez in Oestlund 5790 (AMES!); Nagel in Oestlund 5001 (AMES!); Nagel in Oestlund 5050 (AMES!). GUATEMALA: Lehmann 1436 (BM!); Standley 25064 (AMES!); Johnson 1178 Gentle 2894 (K!); Schipp S-971 (AMES!). HONDURAS: Standley 55583 (AMES!); (AMES!). COSTA RICA: Brenes 690 (F!); von Wedel 1529 (BM!); (AMES!). PANAMA: Standley 29103 (AMES! sterile); Standley 29731 (AMES! sterile); Proctor 287 (AMES!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses La Flora de Jauneche: 110, pl. 44-B; Castillo Mayda, Rico and the Virgin Islands: 54, pl. 11; Ackerman, J.D. 1995. An Orchid Flora of Puerto Rico and the Virgin Islands: 176, fug. 96.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla389V. planifolia var. gigantea Hoehne = V. hostmanii V. planifolia var. macrantha Griseb. = V. phaeantha SOURCES of Jamaica.: 207-208. Foldats, E. 1969. Flora de may be based on a different species); Dodson, C.H. 44-B. 79. V. platyphylla Schltr. in Repert. Sp. Nov. Regni Veg. 10: 7 (1911). Type: Sulawesi, Gunong Klabat, c. 400 m, 13 Dec. 1909, Schlechter 20556 (holo. B; iso. K!). DISTRIBUTION : Endemic to Sulawesi. This poorly known species has the vegetative aspect of V. kinabaluensis the large, thick capsule and the form of the buds are reminiscent of V. kinabaluensis. An illustration of a SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 213; Mansfeld, R. 1934. Repert. Sp. Nov. Regni Veg. 74: t. 14. V. pleei Portres in Bull. Soc. Bot. France 58: 94 (1951). Type: Ile de la Martinique, Ple 621 (holo. P). was based on a mixed collection apparently containing the leaves of V. mexicana V. pompona or of an allied species. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses of the Lesser Antilles. Orchidaceae: 46. 80. V. poiteai Rchb. f. in Linnaea 41: 66 (1877). Type: Santo Domingo, Poiteau s.n. (holo. W 19346!). V. aphylla sensu Eggers in Vidensk. Meddel Dansk Naturhist. Foren. Kjoebenhavn 21 (1889), non Blume (1825). Type: Santo Domingo, Llanos de Rafael, 200 m, 7 June 1887, Eggers 1958 (iso. K! sterile; G 7889/172! sterile). V. eggersii Rolfe in J. Linn. Soc. Bot. 32: 472 (1896). Based on V. aphylla Eggers. V. correllii 33(2): 192 (1981). Type: Bahama Islands, Grand Bahama, Freeport Hammock, off Coral Road, (holo. NY; iso. FTG). DISTRIBUTION : Bahamas, Cuba, Hispaniola, and Puerto Rico. Vanilla, in which small, hooked leaves are present at each node. It has an irregular, crisped and maroon margin. The identity of this species has been recently solved by Ackerman COLLECTIONS : DOMINICAN REPUBLIC: Fuertes 912 (BM! K!); Wright 3351 (BM! AMES!, K! sterile; formerly mixed with the isotype of V. phaeantha). PUERTO RICO: Garber 20 (K! fruits only). SOURCE : Ackerman, J.D. 1995. An Orchid Flora of 81. V. polylepis Mwinilunga District, River Matonchi, 23-24 Oct. 1937, Milne-Redhead 2930 (holo. K!; iso. K!). V. schliebenii 1937: 86: 42, 43, 58 (1937), nomen. Based on: Tanzania, Schlieben 1422 (G DISTRIBUTION Uganda. A species closely related to V. imperialis, differing by its notched lip, shorter, broader tepals, lip with hairs), and a purple-violet blotch on lip, rather than a lip marked with streaks. COLLECTIONS : D.R. CONGO: Seighita 3570 (BR, K!); Rogers 148 (BM! K!); van Hirschberg 28 (K!). KENYA: Verdcourt 1011 (K!); Piers 75 (K!, drawing),; 19527 Burtt 18136 (BM!); Petterson 3-a MilneReadhead 4265 (K!); Richards 2026, 15355, 20509 (K!); Morze 126A, 126B (K!); Bredo in Moreau 864, 864A, 864B, 867 (K!); Fanshawe 2545 (K!),; 9660 (K!); Stevenson 1653 (K!); Holmes 0137 (K!); Burtt 6338 (K!); Harder et al. 2044 (K!). MALAWI: La Croix et al. 880 (K!); Pawek 1020 Ball 844 (K!); Chase s.n., 3896 (K! BM!),; 4307 (K! BM!),; 4680 (K!, pictures; BM!); Renz et al. 11344 (K!); Flowers 2667 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Orch. Trop. Afr.: 114, t. 45; Hunt, P.F., 1974. K.B. 29:

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.390 LANKESTERIANA425; Stewart, J. 1974. Upland Kenya Wild Flowers: 756; Williamson, G. 1977. Orchids of South Central Rwanda-Burundi): 243-244; Stewart, J. 1996. Orchids 1999. Fl. de Cameroun: 288, pl. 102, I-M. 82. V. pompona Schiede in Linnaea 4: 573 (1829). Type: Mexico, Schiede 1043 (lecto. BM!, designated by Garay, Fl. Ecuador Orch. 1:52. 1978; isolecto. K! LE! P! W!). V. pittieri Schltr. in Repert. Spec. Nov. Regni Veg. 3: 106 (1906). Type: Costa Rica, Pittier 6600 (holo. B; trace at AMES 24329!). DISTRIBUTION : Mexico, Nicaragua, Costa Rica, and probably W Colombia and Ecuador. Vanilla pompona is a variable, widely distributed species. We consider it to be a species complex. apparently isolated from the rest of the populations of the species because there are no known populations in Chiapas, Guatemala, Belize, or El Salvador. It reappears only in Central Honduras and Nicaragua. COLLECTIONS : MEXICO: without precise locality, Sall (BM!). Michoacan, Hinton et al. 13815 (AMES!); 15910 (AMES!). Guerrero, Langlass 941 (AMES! G 7889/153! 154! K!); Hinton 11387 (K!). OAXACA: Cuming s.n. (G 78889/155!); Nagel in Oestlund 4574 (AMES!); Copalita, 6102 (AMES!); Veracruz, Bourgeau 2232 (K!); Nagel in Oestlund 2684 (AMES!). COSTA RICA: Alfaro 269 (AMES!). COLOMBIA: Smith 2370 (K!) 2371 (AMES! K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. pittieri); Garay, L.A. 1978. Flora of Ecuador. Orchidaceae McVaugh, R. 1985. Flora Novo-Galiciana 16: An Orchid Flora of Puerto Rico and the Virgin Islands: 174-176; Pupulin, F. 1998. Orchid Florula of Parque Nacional Manuel Antonio, Quepos, Costa Rica. Rev. Biol. T rop. 46(4): 1029. V. ponapensis = Pseudovanilla ponapensis V. porteresianaV. hostmanii V. preussii Kraenzl. = V. inodora 83. V. pseudopompona Soto Arenas, ined. DISTRIBUTION : Panama. This species is very near to V. pompona, but it has larger penicillate callus. COLLECTIONS : PANAMA: Seeman 1159 (BM! K-L!); Allen 4428 (G 7889/121!); (AMES! G7889/151!); Powell 137 (AMES! BM!); Powell 3507 (AMES!); Maxon 6779 (AMES!); Standley 29919 (AMES!); Bangham 458 (AMES!); Piper s.n. (AMES!). V. pterosperma Wall. ex Lindl. = Galeola hydra V. purusara Barb. Rodr. ex Hoehne = V. appendiculata 84. (J.J. Sm.) J.J.Sm. in Bull. Jard. Bot. Buitenzorg ser. 3, 2: 22 (1920). Based on same type as V. ramosa J.J.Sm. V. ramosa J.J.Sm. in Repert. Sp. Nov. Regni Veg. 11: 130 (1912), non Rolfe. Type: New Guinea, Papua (Dutch New Guinea), Jayapura (Hollandia), Gjellerup 706 (holo. L). DISTRIBUTION : New Guinea: Irian Jaya and Papua New Guinea, in the Southern Highlands Province and on Normanby Island. The small leaves (100-130 25-40 mm) and the with the V. kinabaluensis complex. We have examined specimens cultivated in the National Botanic Gardens, Papua New Guinea which correspond well with the published drawings and share with other New Guinean species the very thick ovaries COLLECTION : PAPUA NEW GUINEA: Streimann LAE 52525 (K!). SOURCES : Smith, J.J. 1915. Die Orchideen von Niederlndisch-Neu-Guinea: 187, t. 62, 100; Portres, R. 1954. Le Genre Vanilla et ses Espces: 210; Howcroft, N.H.S. 1984. A Guide to the monocotyledons of Papua New Guinea. R.J. Johns

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla391 Forestry Department PNG, University of Technology. Papua New Guinea. SNP Ltd. p. 9. 85. V. ramosa Rolfe in J. Linn. Soc. Bot. 32: 457 (1896). Types: West Tropical Africa, Niger, about Angiama, Barter 20134 (lecto. K! selected by Geerinck, 1984); interior of Western Lagos, Rowland s.n. (syn. K!). V. ovalifolia Rolfe in J. Linn. Soc. Bot. 32: 457 (1896). Type: West Tropical Africa, Old Calabar, Thomson 132 (holo. K!). V. laurentiana De Wild, Not. Pl. Utiles Congo, 1: 327 (1903). van Rysselberghe 5 (holo. BR, iso. K!). V. africana Lindl. var. laurentiana (De Wild.) Portres in Bull. Soc. Bot. France 98: 126 (1951). V. laurentiana De Wild. var. gilletii De Wild., loc. Kimpasa, Collet 3362 (holo. BR; iso. K!). V. africana Lindl. subsp. ramosa Olszewski in Fl. Cameroun 34: 286 (1988). DISTRIBUTION : Ghana, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Democratic Republic of the Congo and Central African Republic. The reports from Eastern Africa are here considered as V. zanzibarica. Vanilla ramosa is a member of the V. africana branched, the oblong-elliptic leaves are much broader, and the midlobe of the lip is oblong-elliptic and apiculate-acuminate. In living condition the lip is V. crenulata which also has large, broad lower bracts in the simple raceme. Some authors (e.g. Geerinck, 1984) have treated V. ramosa as synonym of V. crenulata, but it differs in the conduplicate-gibbose lip of V. crenulata, in its broadly elliptic to suborbicular measurements. COLLECTIONS : GHANA: Bowling 36647 (K!); Irvine 2197 (K!); Andoh FH 5257 (K!). NIGERIA: Talbot s.n. (K!); Kennedy 220, 2081 (K!). CAMEROON: Bates 1117 (BM!); Zenker 1127 (BM! K!) (BM! K!); Thomas 5922 (K!); FHI 35648 (K!); Leeuwenberg 5465 (K!). GABON: Le Testu 863 (BM!), 7723 (BM! K!), 9015 (BM! K!); Tisserant 1575 (BM!). D.R. CONGO: De Giorgi 1715 (AMES! BM! K!); Louis 8577 (BM!); Vermoesen 1869 (K!); Lebrun 1948 (K!); Germain 4520 (K!); Lebrun 2493 (K!). EQUATORIAL GUINEA: Tessmann 224 (K!). WITHOUT LOCALITY: Kew spirit collection 45787 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses 242; Summerhayes, V.S. 1968. Flora of West Tropical Africa 3(1): Orchidaceae: 205-206; Szlachetko, D.L. V. ramosa J.J. Sm. = 86. V. ribeiroi Hoehne, Comm. Lin. Telegr., Bot. 1: 28 (1910). Type: Brazil, Matto Grosso, by Rio Jaur, near Praxedes, Nov. 1908, Ribeiro in Hoehne 997 (holo. RJ 2512!). DISTRIBUTION : Brazil, Guyana, and Amazonian Colombia. with conspicuous apical, retrorse, yellow trichomes. The lip covered with hairs is reminiscent of that of the Central American V. helleri, which has a more trilobed lip and broader leaves; in V. ribeiroi the leaves are very short and narrow. COLLECTIONS : COLOMBIA: 15961 (AMES!). GUYANA: Jansen-Jacobs et al. 2524 (K!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 185-186 V. ronoensis Hayata = V. somai 87. V. roscheri Rchb. f. in Linnaea 41: 65 (1877). Roscher s.n. (holo. W 19388! W 19389! drawings; K! tracing of W specimen). DISTRIBUTION : Ethiopia, Kenya, Somalia, Tanzania Africa. lip, petals which are very broad and a lip with a somewhat undulate margin. It is very closely related to V. madagascariensis and V. phalaenopsis from the

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.392 LANKESTERIANAcorrect name for the species is V. phalaenopsis, the oldest epithet. Vanilla roscheri is common in tropical deciduous coastal forests of Kenya and Tanzania, and it is occasionally cultivated as ornamental. It has been only recently recorded from near Lake Sibaya in South Africa. COLLECTIONS : KENYA: Bally 8144 (K! K photos!), B10462 (G 7889! K!), 8540 (K!); Verdcourt 1195, 3223 (both K!); Brenan et al. 14549 (K!); Evans 70/781 (K!); (K!); Napper 1468 (K!); Tweedie 1217 (K!), 1260 (K!); (K!); Graham 1605 (K!); Greenway s.n. (K!), 9487 (K!); 10125 (K photo!); Greenway 12455 (K!), Moreau 134 (K!); 134A (K!); Archbold 2786 (K!); Richards 21992 (K!); Franzier 1185 (K!); Vaugham 2530, 2703, 2785 (all K!); Greenway 4125 (K!); Tanner 2453 (K!); Grey-Wilson 10013 (K!); (K!); Curle 442 (BM!); Pemba, Lambu s.n. (K!); Vaughan 937 Kirk 3/701 (K!, 3/78 K! drawing); Lyne 4 (K!); Greenway 1315 (K!), 2669 (K!); Wigg s.n. (K!); Burt 2626, 2628 (K!); Vaugham 45 (BM!), 1662, 2153, 2173 (all K!). SOUTH AFRICA: (K! NH), 338 (K!). Janson 7492 (K!); Danitree s.n. (K!); (K!); Ball 934 (K!); (K!); Grandvaux B 7724 (K!); Ball 1000 (K!); Watmough 351 (K!); Greenamdijk (K!); W.E. Taylor s.n. (BM!); Scott Elliot 6132 (BM!). SOURCES : Rolfe, R.A. 1897. Fl. Trop. Afr. 7: 179; 24: 41; Cribb, P. 1984. Flora of Tropical East Africa. 1999. Orchids of Southern Africa: 350-351. V. rubiginosa Griff. = ?Galeola hydra V. rubra (Lam.) Urb. = ?V. planifolia 88. V. ruiziana Klotzsch in Bot. 563 (1846). Type: Peru, Chicoplaya, (holo. B ; iso. AMES 38626! and photo, G 7889/ 167! F!; tracing K! BM! G 158!). V. weberbaueriana Kraenzl. in Bot. Jahrb. Syst. 37: 395 (1906). Type: Peru, Prov. Tarma, Dep. Junin, near la Merced, Chanchamayotal, 1000 m, M. (holo. B, AMES 38631 photo! iso. MOL!). DISTRIBUTION : Peru and Bolivia. The identity of V. ruiziana has remained obscure since its publication in 1846. The holotype specimen, lost when the Berlin Herbarium was destroyed, was sterile, having stout stems and acute to acuminate, lanceolate-oblong leaves. However, at least a duplicate of the same collection ( Epidendrum lanceolatum, Pavn handscript, and annotated “M. Pavn”, perhaps in Reichenbach’s hand) has been found in Geneva preserved, but it is possible to appreciate its granulose sepals and ovary surface, the thickened veins of the lip and the acute, strongly subcalyptrate sepals. This combination of characters is exclusive to the modern specimens from Peru and Bolivia. The type specimen of V. weberbaueriana, although bearing only young buds, also belongs here. Leaf shape differences between the types of V. ruiziana and V. weberbaueriana seem striking, but there are intermediate specimens connecting both extremes. The vegetative parts of V. ruiziana seem to be taken from a shaded spot. Portres (1954) assigned V. weberbaueriana to subsection Membranacea (= subsect. Vanilla). The study of more abundant material shows that V. ruiziana is closely related to V. hostmanii, differing mostly in the sepals conspicuously calyptrate and a larger, ovate midlobe to the lip which bears 1-3 elevated, axial keels near the apex,. COLLECTIONS : BOLIVIA: White 2326 (AMES!). PERU: (AMES!); Rubio Kayap 223 (MO 2922173!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 233 and 253-254; Schweinfurth, C. 1958. Orchids of Peru I. Fieldiana: Botany 30: 44. V. sativa Schiede = V. planifolia 89. V. savannarum Britt. in Mem. Torr. Bot. Club 16: 61 (1920). Type: Cuba, near Camagey, Cowell 13210 (not seen)

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla393 DISTRIBUTION : Cuba. buds show a very close relationship with the South American V. palmarum , and these species must be compared carefully to determine if they are distinct. With Vanilla palmarum , it shares the similar habit and habitat, the lack of a penicillate callus, the presence of 5-6 axial, shortly pubescent lines on the distal half of the lip, the long lanuginose area beneath the stigma, and the lip fused to the column for only about a half of the column length. The densely-fruited raceme and the pods with a well-developed calyculus suggest self-pollination. COLLECTIONS : CUBA: Gonzles 288 (AMES!); (AMES!); Schaffer 615 (para. AMES!), 1072 (para. AMES! K!); Ekman 19043. SOURCES Alain. V. schliebenii Summerh. = V. polylepis 90. V. schwackeana Hoehne in Arq. Bot. Estado So Paulo 1: 125. t. 135 (1944). Type: Brazil, Minas Gerais, Schwacke 11106 (holo. RJ 37015!). This species seems to be related to V. cristagalli, but the ornamentation on the lip is much reduced to warty, apical papillae. Hoehne (1944) indicated that the specimen could be a mixture of parts of different species, the leaves being very long acuminate, and thin. Specimens collected in Amazonia (J.B.F. da Silva 590 those of the type of V. schwackeana, but with thick, coriaceous, oblong-elliptic leaves with recurved margins, suggesting that Hoehne’s doubts about the mixture nature of the type specimen had substance. COLLECTIONS da Silva 590 (MG!), 591 (AMO! sterile). 91. V. seranica J.J. Sm. in Bull. Jard. Bot. Buitenzorg ser. 3, 10: 93 (1928). Types: Moluccas, Ceram, Rutten 1800 1880 (syn. L). DISTRIBUTION : E Indonesia, Moluccas. Said to be allied to and V. klabatensis. SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 215. 92. V. seretii De Wild. in Bull. Jard. Bot. tat 5(2): 180 Seret 756 (syn. BR); near Arebi, 1906, Seret 714; Nala (syn. BR); 1911, Boone 111 (syn. BR); Libenge, 1913, Mestdagh 59 (syn. BR; isosyn. BM!). V. tisserantii Portres in Bull. Soc. Bot. France 98: 93 (1951). Type: Central African Republic (French Equatorial Africa), Bukoko, Oubangui, SW of Bangui, 18 Aug. 1947, Tisserant 151 (holo. P; iso. BM! G!). DISTRIBUTION : Democratic Republic of the Congo and Central African Republic. This is another member of the V. chalotii group, in which the lip is deeply saccate. It is distinguished from V. nigerica by its lip in which the throat has few hairs and lacks a gibbous base. COLLECTION : D.R. Congo: H. Callens 906 (K!). SOURCES : Portres, R. 1954. Le Genre Vanilla et ses V. tisserantii); Szlachetko, 102, A-H. Vanilla shenzhenicaV. somai 93. V. siamensis Rolfe ex Downie in Bull. Misc. Inf. Kew 1925(10): 410-411 (1925). Type: Thailand, Doi Sutep, 690 m, Kerr 95 (holo. K!; iso. C!). V. pierrei Gagnep. in Bull. Mus. Hist. Nat. (Paris) 3: 686 (1931). Types: Cambodia, Mts Kamchay, Prov. Kampot, Pierre 1560 (syn. P); Cambodia (Cochinchina), Mts Binh-dinh, Pierre s.n. (syn. P). DISTRIBUTION : Thailand, Kampuchea (Cambodia), Vietnam, China (Yunnan). This species is characterized by its large, broadly elliptic or ovate, shortly apiculate leaves. The large is similar to and V. kinabaluensis. We here consider V. pierrei was described from immature buds, but the general appearence is the same as V. siamensis. However, we would like to compare better material of V. pierrei with that of V. siamensis. Vanilla siamensis is a montane species found from 700 to 1300 m elevation. COLLECTIONS : THAILAND: R. Geesink et al. 6755 (K!), 6698 (K!); without collector 38 (K! sterile).

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.394 LANKESTERIANA SOURCES : Ga Genrale de LIndo-Chine 2. Orchidaces: 582-584, Vanilla et (as V. pierrei The Orchids of Thailand: a preliminary list, part 1: orkhidnykh: 112 (as V. pierrei); Averyanov, L.V. 1994. V. siamensis Orchids of China. Science Press, Beijing: 411. 94. V. somai Hayata, Ic. Pl. Formosan. 6: 88, t. 14 (1916). Type: TAIW AN , between Urai and Agisku, May 1916, Hayata s.n. (holo. TI). sensu Hayata in J. Coll. Sci. Imp. Univ. Tokyo 30, 1: 343 (1911), non Rchb. f. (1854). Rchb. f. var. formosana Ito, Icon. Pl. Japan. 1(4): 1 t. 1 (1911). Vanilla ronoensis Hayata, Icon. Pl. Formosan. 9: 114 (1920). Type: Taiwan, between Rono and Kosenpo, April 1917, Hayata s.n. (holo. TI). Rchb. f. var. ronoensis (Hayata) S.S. Ying, Col. Illustr. Indig. Orch. Taiwan, 1(2): 509 (1977). Vanilla albida sensu H. J. Su in Huang, T.S. et al. (eds), Flora of Taiwan 5: 1065 (2000), non Blume (1823). Vanilla shenzhenica Phytotax. Sinica 45, 3: 301 (2007). Type: China, Guangdong, Shenzhen, Longgang, Meishajian, 22 Feb 2005, Liu 3025 (holo. NOCC). DISTRIBUTION : Taiwan, Vietnam, and China. structure is similar to that of V. annamica, but in that COLLECTIONS : CHINA: Guangdong, Liu 3025 (NOCC!). TAIWAN: Price 302 (K!), 302 bis (K!); Henry 479 (K!). SOURCES : Hayata, B. 1911. J. Col. Sci. Univ. Tokyo 30(1): 342 (as ); Portres, R. 1954. Le Genre Vanilla et ses Espces: 183 (as V. ronoensis) and 184pl. 50: 54-1 (as ); Ying, S.S. 1977. Coloured Illustrations of Indigenous Orchids of Taiwan 1: 338340, pl. 339 (as var. formosana); Chow Cheng. 1989. Formosan Orchids. Epiphytes: 204-206 (as V. albida); Tsan-Piao Lin. 1975. Native Orchids of Taiwan: 248-249. V. speciosa Boxall ex Neves, in Blanco, Fl. Philipp. de. III. Nov. App. 248, nomen. 95. V. sprucei Rolfe in J. Linn. Soc. Bot. 32: 461 (1896). Type: Colombia, R. Vaups, Nov. 1852, Spruce 2727 (holo. K-LINDL!). DISTRIBUTION : Amazonian Colombia. This species is rather mesophytic, the type having slender stems, drying almost black, and thin leaves covered with epiphylls. The parallel veins and transverse secondary veins of the leaves resemble those of Epistephium leaves. The raceme is lax with about 4 to 8 (perhaps more) successive whitishlong tubular lip with a conspicuous penicillate callus. is in anthesis, perhaps in the way described for V. , to which might be related. Cabrera 12497 very odd for American species. A calyculus is scarcely present. COLLECTION : COLOMBIA: 12497 (AMES!). SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 247. 96. V. sumatrana J.J. Sm. in Bull. Jard. Bot. Buitenzorg, ser. 3, 2: 22 (1920). Type: SUMATRA , Benkoelen, Bukit Barisan, Rimbo-Pengadang, 1000 m, June 1916, Jacobson (L not seen). DISTRIBUTION : Sumatra. A little known species which, in the protologue, was compared with Rchb. f., but differs in the narrowly spathulate sepals and petals, the small the series of transverse lamellae beneath the hair cushion, and the different anther. It has a reddish lip. It is found at higher elevations, more than 1000 m, than .

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla395 SOURCES : Carr, C.E. 1935. Two Collections of Orchids from British North Borneo 1. Gard. Bull. Straits Settlements: 8: 176; Portres, R. 1954. Le Genre Vanilla V. surinamensis Rchb. f. = V. guianensis 97. V. tahitensis J.W. Moore in Bull. Bernice P. Bishop Mus. 102: 25 (1933). Type: Society Islands, Raiatea, Moore 294 (holo. BISH!). V. tiarei 201 (1915), nomen nudum. V. hirsuta 1(1): 47 (1996). Type: Papua New Guinea, Clements 6742 (holo. CANB; iso. NCBG). V. tahitensis var. haapape Portres, Bull. Soc. Bot. France 98:127 (1951), nomen nudum. Based upon: cult.. Tahiti, without further provenance. V. tahitensis var. potiti Portres, Bull. Soc. Bot. France 98: 127 (1951), nomen nudum. Type: cult. Tahiti, 1926, without further provenance. Vanilla tahitiensis is the second most important vanilla for production of aromatic pods. It is closely related to V. planifolia and V. odorata and a member of the American clade of the genus with penicillate is striking, and it is very probable that it is a hybrid distributed originally from the Philippines and taken to its present area by man. A good drawing of it is provided by Howcroft (1984). The Tahitian vanilla is a probably a hybrid between Vanilla odorata and V. planifolia. Five varieties are listed by Portres (1959). COLLECCTIONS : FRENCH POLYNESIA: Raiatea, St. John 17310 (K!); Mongareva Isl., St. John 14459 (K drawing!). SOURCES : Howcroft, N.H.S. 1984. A Guide to the monocotyledons of Papua New Guinea. R.J. Johns Forestry Department PNG, University of Technology: Vanilla et ses Espces: 239-242, Vanilla taiwaniana S.S. Ying. See in excluded species. Vanilla tiarei V. tahitiensis V. tisserantii Portres = V. seretii V. tolepephora Ridl. = ? 98. V. trigonocarpa Hoehne in Arq. Bot. Estado So Paulo, n.s., 1(6): 126 (1944). Type: Brazil, Par, outskirts of Belem do Par, 23 Nov. 1926, Ducke s.n. (holo. RJ 19445!). R.L. Dressler in Orquideologa 13(3): 229 (1979). Type: Panama, Dressler 5290 (holo. US). DISTRIBUTION : Costa Rica, Panama, Colombia and Brazil. A very distinct Vanilla known from wet forests in Central America, Colombia, and from the delta of the Amazon River. The main difference between V. trigonocarpa and is the more trilobed lip of V. trigonocarpa. The plant seems to be related to V. espondae from Colombia, and perhaps to V. sprucei. COLLECTION : COSTA RICA: C24.1 (AMES!). 99. V. utteridgei J.J.Wood in Orchid Rev. 110: 232 (2002). Type: New Guinea, Papua, Mimika Regency, Kuala Kencana, Utteridge 456 (holo. K). DISTRIBUTION . New Guinea: S Papua only. Closely allied to V. kempteriana, differing in its pubescent column and the two lateral calli in front of the penicillate callus. V. uncinataV. odorata V. vanilla (L.) H. Karst. = ?V. mexicana V. vanilla (L.) Huth = ?V. mexicana 100 V. vellozii Rolfe in J. Linn. Soc. Bot. 32: 467 (1896). Type: Brazil, Glaziou 14302 (lecto. K! designated here). V. chamissoins var. brevifolia Cogn., Fl. Bras. 3(4): 149 (1893). Based on Epidendrum vanilla Vellozo, F.Flum. Ic. 9: t. 1 (1827). V. argentina Hicken in Anales Soc. Ci. Argent: 235. 1916; Vanilla Argent. 3 (1917). Types: Argentina, Formosa, Pilcomayo, Cceres (not seen; 3 immature fruits only); Gran Chaco, J. Domnguez 307, 25-IX-1915, Inst. Darwiniano DISTRIBUTION : Brazil, Paraguay, Argentina. This species has been confused with V. chamissonis but it is easily distinguished by its shorter segments, especially its very broad, subentire lip. V. chamissonis has a very elongate midlobe to the lip

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.396 LANKESTERIANAand longer tepals. It seems to be common in coastal areas. However, they are probably closely related, sharing with V. calyculata the conspicuous calyculus, COLLECTIONS Dusn 16125 (AMES!). Rio de Janeiro, Glaziou 11621 (K! without van der Berg s.n. (AMO slides!). PARAGUAY: Fiebrig 564 (K! AMES! G7 889/113!). 101. V. verrucosa Hauman in An. Mus. Nac. Buenos Aires 29: 365-366 (1917). Type: ARGENTINA , Misiones, Piray, March 1901, Kermes 127 (holo. Herb. Min. Agr. 6358; not seen) V. verrucosa is distinguished from its close relatives by its very small dimensions and the triangular acuminate midlobe of the lip. Andrs Johnson (pers. com.) indicates that the type specimen is sterile and that the species has not been recollected. He stated that is not improbable that V. verrucosa is an atypical form of V. rojasiana Hoehne (=V. angustipetala Schltr.). We suspect that the original description might have been based on an immature bud. The small members of the membranaceous group found in Brazil and Argentina are a taxonomically preservation or condition of the type specimens, and the similarity of the described taxa. Vanilla angustipetala (syn. V. rojasiana) and V. parvifolia in V. parvifolia and rows of warts on the lip in V. angustipetala. The plant illustrated in t. 22 of Hoehne’s Flora Brasilica V. parvifolia, shows midlobe of the lip, approaching to V. edwallii or V. organensis. SOURCE : Portres, R. 1954. Le Genre Vanilla et ses Espces: 188. Blume = V. planifolia 102. V. walkeriae Wight, Icon. Pl. Ind. Orient. t. 932 (1845). Type: SRI LANKA , Walker s.n. (holo. K!). DISTRIBUTION : India and Sri-Lanka. This species has been repeatedly confused with V. wightii. V. walkeriae species V. phalaenopsis and V. madagascariensis, with longitudinal rows of hairs on the lip disc. It has been confused with V. wightii. Part of the confusion can be on the type sheet of V. walkeriae at Kew. COLLECTIONS : SRI LANKA: Champion s.n. (K-LINDL!); (K!); Mueller-Dombois et al. s.n. 69042619 (K!); Thwaites s.n. 2964 (K!); Cramer 4664 (K!). INDIA: Fisher 1011a (K!); Wight s.n. (K!). SOURCES : Trimen, H. 1898. Handb. Fl. Ceylon 4: 220; Seidenfaden, G. 1983. The Flora of the Tamilnadu Carnatic. Orchidaceae: 1570-1571, pls. 698 and 699; Pradhan, U.C. 1976. Indian Orchids: Guide to 1981. Orchidaceae. A Revised Handbook to the Flora of Ceylon 2: 331. 103. V. wariensis Schltr. in Repert. Sp. Nov. Regni Veg. Beih. 1: 30 (1911). Type: PAPUA NEW GUINEA , Schlechter 19928 (holo. B). DISTRIBUTION : Endemic to Papua New Guinea. O’Byrne (1994) mentioned that it is common to the North of the Owen Stanley Range. It can be distinguished from other New Guinean vanillas by its very large leaves 27-35 5-7 cm. SOURCES : Portres, R. 1954. Le Genre Vanilla et ses Espces: 212; Howcroft, N.H.S. 1984. A Guide to the monocotyledons of Papua New Guinea. R.J. Johns Forestry Department PNG, University of Tchnology. p. 126; O’Byrne, P. 1994. Lowland Orchids of Papua New Guinea. SNP Ltd. p. 9. V. weberbaueriana = V. ruiziana 104. V. wightii Lindl. ex White, Icon. Pl. Ind. Orient. sub t. 931 (1844). Type: S INDIA , Wight 2091 (holo. K-LINDL! iso. K!). V. aphylla Wight., Icon. Pl. Ind. Orient. t. 931 (1845), non Blume (1825). Based upon Travancore, Trevandrum, Wight s.n. (K). V. wightiana Lindl. in Hook.f., Fl. Brit. India 6: 90 (1890), sphalm. pro V. wightii. DISTRIBUTION : India (Deccan) and Sri Lanka. This species has been confused in the literature. The type collections at K consist of different, separated

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOTO ARENAS & CRIBB Vanilla397fragments on t hree sheets, some fragments seem to be referable to Wight’s concept of V. wightii , and others to V. walkeriae . Vanilla wightii is closely related to V. aphylla and V. calopogon , while V. walkeriae is a member of the V. madagascariensis group. In V. wightii has two rows of hairs from the base to the sides of the crest, which is like a large penicillate callus continuous with an apical strip of hairs. A fragment in each one with a thick fruit and many protuberant scars, lacking bracts, typical of V. walkeriae , in which the bracts are deciduous. Seidenfaden (1983) illustrated a specimen of V. walkeriae with narrow segments, but lacking the hirsute lip apex characteristic of the V. aphylla complex. Pradhan (1976) also illustrated V. walkeriae rather than the true V. wightii . A good illustration of V. wightii can be found in Abraham and Vatsala (1981). Although both V. wightii and V. walkeriae belong to different lineages of the genus. Vanilla walkeriae resembles closely V. roscheri COLLECTIONS : WITHOUT LOCALITY: cult. Hort. Bot. Reg. Kew a. 191 (K!, in part). SRI LANKA: C.V. 2964 (K! watercolor). SOURCES to Orchids: 500-502. Seidenfaden, G. 1983. The Flora of the Tamilnadu Carnatic. Orchidaceae: 1570, pl. 698; Pradhan, U.C. 1976. Indian Orchids: Guide to V. wrightii Rchb. f. = V. bicolor 105. V. yersiniana Mus. Hist. Nat. sr. 2, 36: 162 (1964). Type: VIETNAM (Annam), road from Yersin to Hon ba, Suoi Dao, cult. Suoi Dao, May 1963, de Sigaldi 362 (syn. P). DISTRIBUTION : Thailand, Vietnam. A common species in Indochina which is readily distinguished by its lanceolate, long-acuminate papillae on the lip apex. The narrow, elongate funnelAsian species, except its close allies V. montana, V. albida and V. andamanica. Good photographs and illustrations can be found in Seidenfaden and Wood V. albida, a different species from Java. V. albida has greener tepals and different papillae at the apex of the lip (see pictures in Comber, 1990). COLLECTIONS : THAILAND : (AAU! K!); Kerr 0805 (K!), 0805A (K!), s.n. (K!); Hull 671 (K! sterile). SOURCES The Orchids of Thailand: a preliminary list, part 1: 60 (as V. sp. Kerr 805 and 805A); Seidenfaden, G, . 1978. Orchid Genera in Thailand VI. Dansk Bot. V. albida Peninsular Malaysia and Singapore pages?? Comber, J.B. 1990. Orchids of Java: 76; Averyanov, L.V., orchid checklist of Cao Bang Province. Lindleyana 15(3): 163 (as V. albida). 106. V. zanzibarica Rolfe in Bull. Misc. Inform. Kew 1906: 116 (1906). Type: ZANZIBAR , on the Wanda, near Dunga, Lyne s.n. (holo. K!). DISTRIBUTION : ?Uganda, Kenya, Tanzania, including Vanilla zanzibarica has been treated as a synonym of V. ramosa (e.g. Cribb, 1984; Stewart, 1996). However, the specimens from East Africa usually have much broader leaves, the midlobe of the lip is broader, obtuse, apiculate and conduplicate, and the total length of the lip is longer (up to 25 mm long). The conduplicate lip is more similar to that of V. crenulata, although not so high and gibbose, but V. zanzibarica lacks the foliaceous bracts of V. crenulata. Leaf shape and size approach those of V. ramosa are much larger. Vanilla zanzibarica seems to be geographically separated from the other members of this complex, being restricted to a small area in the the Shimba Hills in Kenya. COLLECTIONS Moreau 408 (K!); Holtz 159 (K!); Greenway in Moreau 408A (K!); Greenway 4684, 7043 (K!); Mcke s.n. (K!); Zimmermann s.n. (K!); Borhidi et al. 86353 (K!).; Vaughan 1254 (K!); Williams 64 (K!); Fawcras 3121 (K!); Greenway 1297, 2606 (K!); Toms s.n. (K!). Pemba, Soaper H45/30 (K!); Williams 110 (K!). KENYA: Luke

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.398 LANKESTERIANA3847 (K!); (K!); Glover 583 (K!). SOURCES: Portres, R. 1954. Le Genre Vanilla et ses Espces: 200-201; Cribb, P. 1984. Flora of Tropical East Africa. Orchidaceae 2: 261, 263 (as V. ramosa); Stewart, J. 1996. Orchids of Kenya: 89-90 (as V. ramosa). EXCLUDED AND POORL Y UNDERSTOOD SPECIES Vanilla carinata Rolfe in J. Linn. Soc. Bot. 32: 468 (1896). Type: BRAZIL . Organ Mountains, Miers s.n. (holo. K!). The type of this species consists of a single sheet up to 7 cm long) and no leaves or stems. The rachis bears two ovaries with a conspicuous calyculus, and the lip seems to be longer than sepals. It could be V. chamissonis or perhaps V. vellozii, but the material are inclined to think that the “strongly keeled” lip apex described by Rolfe is an artifact due to the poor Hoehne (1945) listed it as a synonym of V. planifolia. Vanilla taiwaniana S.S. Ying in Quart. J. Chinese Forest. 20, 2: 55 (1987). Type: Taiwan, Taichuang, July 1984, F.S. Ho s.n. (holo. not located). A poorly known species. The short description suggests that it could be a synonym of Erythrorchis ochobiensis (Hayata) Garay, but with its green colour and developed leaves it may belong to a Pseudovanilla. ACKNOWLEDGMENTS : We thank the curators of the following herbaria for the loans or permitts to examine their collections: AMES AMO, BM, BR, CAS, CEN, CR, ENCB, G, HAC, HAJB, HB, K, LE, MA, MEXU; MG, MO, NY, P, RJ, US, UVAL, W, XAL. We would like to thank J. Ackerman, the late J.B. Comber, Andrs Johnson, G. Salazar, A. Toscano de Brito and Bill Goldner who offered important information, opinion or material. LITERATURE CITED Comber, J.B. 1990. Orchids of Java. Bentham-Moxom Trust, Royal Botanic Gardens, Kew. pp. 74-76. Christenson, E. 1995. The long-lost Vanilla bicolor. Amer. Orch. Soc. Bull . 64(8): 844-847. Correll, D.S. 1951. Orchids of North America north of Mexico. Chronica Botanica, Waltham, Mass. pp. 155163. Vanilla . In Polhill R. (ed.), Flora of Tropical East Africa Orchidaceae 2: pp. 258-263. Orchids Illustrated. 1-6. A. Deutsch, London. Hoehne, F.C. 1945. Flora Brasilica 12, 2. Orchidaceas. Secretara de Agricultura, Industria e Comrcio de Sao Paulo, Brasil, pp. 3-36, t. 1-29. Klotzsch, J.F. 1845. Ueber die Arten der Gattung Vanilla . Orchidaceae. 1. Balkema, Rotterdam. Luer, C. 1972. The Native Orchids of Florida. New York Botanical Garden, pp. 68-69. Porters, R. 1954. Le genere Vanilla et ses espces: pp. 94290. In G. Bouriquet, ed., Le Vanillier et la Vanille dans le Monde. Enc. Biol. 46. Ed. Paul Lechevalier, Paris. Rolfe, R.A. 1896. A revision of the genus Vanilla . J. Linn. Soc. 32: 439-478. of euglossine orchid-bees (Apidae: Euglossini) in Panama. Oecologia (Berlin) 73: 321-333. Dressler, R.L. 1981. The Orchids. Natural History and . Press, Cambridge, 332 p. Family. Dioscorides Press. Portland, Oregon, 314 p. Seidenfaden, G. 1992. Orchids of Indochina. Opera Botanica 114: 71-72. Vanilla (Orchidaceae) and the evolution of pollination systems in the genus. Stewart, J. 1996. Orchids of Kenya. St .Paul Bibliographies, Winchester. Summerhayes, V.S. 1968. Flora of West Tropical Africa. Orchidaceae, ed. 2. Crown Agents, London, pp. 205-206. Orchidaceae 1: 279-294. pollination and evolution. Univiversity of Miami Press, Coral Gables, Florida, 23, 77, 125.

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COMPARETTIA SOTOANA (ORCHIDACEAE: ONCIDIINAE), A NEW ECUADORIAN SPECIES FRANCO PUPULIN 1 & GILBERTO MERINO 31 Jardn Botnico Lankester, Universidad de Costa Rica, P. O. Box 1031-7050, Cartago, Costa Rica.2 Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts, U.S.A.3 ngel Andreetta Research Center on Andean Orchids, Universidad Alfredo Prez Guerrero, Ecuador4 Corresponding author: fpupulin@cariari.ucr.ac.cr399. 2010. LANKESTERIANA 9(3): Some of the genera of the subtribe Oncidiinae, belonging to the so-called Rodriguezia clade, variously treated as the Subtribes Comparettiinae Schltr. (i.e., Senghas, 1995), had a complex taxonomic history, with many species moving forth and back to alternative genera. This has been particularly true for the genera close to Scelochilus Schltr., including Neokoheleria Schltr., Scelochiloides Scelochilopsis natural relationships through nomenclature was done by Dodson and Chase (in who reduced Neokoheleria under Scelochilus. The generic circumscription of Scelochiloides has been also subjected to different interpretations, with species of this genus assigned to both Neokoheleria (Senghas 2001), and Scelochilus (Kniger 2008). Vegetative architecture is amply variable among members of the highly derived Rodriguezia-clade. Many of the genera as traditionally circumscribed organization, often with retention of paedomorphic traits in the adult psygmoid habit and reduction of pseudobulb size, and leaf shape. Within the clade, the genera Ionopsis Kunth, Neokoehleria, Rodriguezia Scelochilus, include species with features have been emphasized to circumscribe genera in the group. Neokoehleria was basically distinguished from Scelochilus by the shape of the sepaline spur (clubshaped vs. conical), and Scelochiloides the presence of a single nectariferous horn at the base of the lip (vs. 2 in Scelochilus), while Scelochilopsis was characterized by the lack of spur at the base of the lip (Dodson 1998). Eventually, the molecular ABSTRACT . A new species of Comparettia , C. sotoana, is described and illustrated from the province of MoronaSantiago, in southeastern Ecuador. Comparettia sotoana is compared with Comparettia langkastii , from lip with a cuspidate midlobe (vs. pandurate with a deeply retuse, apiculate midlobe) provided with digitateconical basal processes, uncinate at apex and expanded into rounded knobs (vs. straight, acuminate), and lacking apical calli (present in C. langkastii ). RESUMEN . Se describe e ilustra una nueva especie de Comparettia , C. sotoana, de la provincia de MoronaSantiago en la regin suroriental de Ecuador. Comparettia sotoana se compara con Comparettia langkastii , de con un lbulo mediano cuspidado (vs. pandurado con lbulo mediano profundamente retuso y apiculado), globosas (vs. rectos, agudos) y carente de callos apicales (presentes en C. langkastii ). KEY WORDS: Orchidaceae, Oncidiinae, Comparettia , C. sotoana, new species, Ecuador

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studies carried out by Chase, Williams and co-workers (Williams et al. 2001a, 2001b, Sosa et al. 2001, Chase 2009), aimed to reconstruct a phylogenetic scheme of relationships within the Oncidiinae, revealed that the genera centered around Comparettia are all embedded within Scelochilus. Morphologically, taxa in this group only differ in subtle details of their nectar horns and the relative length of the spur that embrace them. In order Chase et al. (2008) combined all the species of this group provided with short spur, previously treated as the genera Chaenanthe Lindl., Diadenium Endl., Neokoehleria, Senghas, Scelochilopsis, Scelochilus, and Stigmatorthos Chase, into the single genus Comparettia. During the ongoing work aimed to the documentation of orchid species in the collection of Ecuagenera, to be accessed to the living collection and ancillary collections of the Andean Orchid Research Center (CIOA by its acronym in Spanish), University Alfredo Prez Guerrero, we discovered a new species of Comparettia that we describe here as new to science: Comparettia sotoanasp. nov. TYPE: ECUADOR . Morona Santiago: Tumbez, 1200 m, collected by G. Merino, 15 February 1993, cultivated by Ecuagenera at Gualaceo, accession No. CIOAF. Pupulin 7063 (holotype, QCA-Spirit). Fig. 1. Species Comparettiae langkastii (Senghas) M.W. labello ovato-subpandurato lobo medio cuspidato, processibus basalibus digitato-conicis apicaliter uncinatis globosis instructo, callibus apicalibus destituto recedit. Epiphytic, erect, caespitose herb. Roots slender, ca. 1 mm in diameter, produced from the rhizome under the pseudobulb. Pseudobulbs small, 4 mm in diameter 5 mm long elliptic, complanate, monophyllous at apex (or with a second, apical, small, the bases of 2 foliaceous sheaths, the sheaths 6 mm long, articulated with the leaf. Leaves laterally acute, minutely mucronate, thick-coriaceous, green suffused with purple, 3.5.0 x 0.4.8 cm, the larger ones provided with a subcylindric petiole to 5 mm long. lateral, arising from the subumbellate raceme to 3 cm long; peduncle erect, terete, to 2 cm long, with 1 strongly conduplicate, bracts lanceolate, glumaceous, 4.0.0 x 1.5 mm. Ovary terete, to 8 mm long including the pedicel. Flowers not completely spreading, campanulate, the sepals pale yellow, the petals greenish white suffused with orange, the lip white spotted orange, the column pale yellow, the anther white. Dorsal sepal ellipticoblanceolate, obtuse, concave, reclined over the column, 4.6 x 2.5 mm. Lateral sepals connate for about 3 mm into an elliptic, concave synsepal, 5.8 x 3.0 mm, the free apexes obtuse to subacute, extending at the base into a sepaline, saccate spur ca. 2 mm long. Petals elliptic, asymmetrical, rounded, apiculate, slightly concave, 5 x 3 mm. Lip 3-lobed, ovatesubpandurate, canaliculated, the lateral lobes elliptic, rounded, suberect; the midlobe cuspidate, slightly conduplicate; the base of the lip protruding toward the rear into two digitate-conical processes, uncinate at apex and expanded into rounded knobs, 2 mm long; disc with 2 low, subquadrate, diverging calli. Column straight, ascending, subterete, thicker toward the apex, 3.7 mm long, provided with broad elliptical wings; stigma rounded. Anther cap cucullate, subquadrate, 2-celled. Pollinia 2, elliptic-obovate, complanate, on an ovate viscidium. Fruit not seen. EPONYMY : dedicated to Miguel Angel Soto Arenas, for his outstanding contributions to Neotropical orchidology. DISTRIBUTION AND ECOLOGY : Know exclusively from Ecuador. Populations of C. sotoana grow epiphytically in open shade in wet forests in the province of Morona Santiago at approximately 1200 meters of elevation, where they are apparently restricted to the terminal branchlets of secondary vegetation. Plants have been C. sotoana, as an unnamed species of Scelochilus, LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.400 LANKESTERIANA

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FIGURE 1. Comparettia sotoana column and lip, lateral view. E — Pollinarium, three views. F — Anther cap, ventral view. Drawn by F. Pupulin from the holotype.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN & MERINO — Comparettia sotoana401

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on the basis of an Ecuadorian collection by A. Hirtz (8134, probably at RPSC, now at MO). Comparettia sotoana belongs to a small group of species, mostly previously included in Neokoehleria, known from the Amazonian basin and the eastern slopes of the Andes from Colombia to Peru and Bolivia. Among them, C. sotoana is most similar to C. langkastii endemic to Ecuador. It differs from C. langkastii in several characters, including the shape of pseudobulb (elliptical-subcylindrical vs. rounded-complanate) and triquetrous), the subumbellate (vs. distichous), severalmm vs. >5 mm long), straight (vs. curved) sepaline spur, and the ovate-subpandurate lip with a cuspidate midlobe (vs. pandurate with a deeply retuse, apiculate midlobe) lacking apical calli (present in C. langkastii). Furthermore, the nectariferous horns at the base of the lip of C. sotoana are digitate-conical, uncinate at apex and expanded into rounded knobs, while in C. langkastii they are straight and apically acuminate. ACKNOWLEDGEMENTS . We are grateful to Jos “Pepe” Portilla and his family for allowing us free use of the plants in the collection of Ecuagenera and for giving us any kind of facilities at Gualaceo. To the Ministry of Environment of Ecuador for extending the management permits of the living collections where the species treated here have been documented. To the Foundation for Orchid Research and Conservation, Vancouver Orchid Society, Canada, for its kind sponsoring of the AORC. The present paper is part of by the Vice-Presidency of Research, University of Costa Rica. LITERATURE CITED plants and gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45. Whitten. 2008. Taxonomic transfers in Oncidiinae to accord with Genera Orchidacearum, vol. 5. Lindleyana (in Orchids) 21(3): 20. Chase. M.W. 2009. Sultribe Oncidiinae. In Pridgeon, A. Genera Orchidacearum, vol. 5. Epiendroideae. Oxford University Press. Dodson, C. H. 1998. Miscellaneous new orchid genera and species. Orquideologia 21(1): 61. Dodson, C. H. 2004. Native Ecuadorian Orchids. Volume 5: Rodriguezia-Zygosepalum. Dodson Publishing, Sarasota, Florida. Kniger, W. 2008. Neue arten der gattungen Oncidium und Scelochilus /New species of the genera Oncidium and Scelochilus 16: 376-391. Senghas, K. 1995. Tribus Oncidieae. Subtribusgruppe (Tribella) Bipolliniata. 72. Ionopsidinae. Pp. 1970– 1973 in Parey, Berlin. Senghas, K. 2001. Nachtrge 19. Tribus Oncidieae. Pp. 2784— 2810 in: Schlechter, Die Orchideen, 3. Bd.1c. Paul Parey, Berlin. H. Williams. 2001. Phylogenetic position of Dignathe (Orchidaceae: Oncidiinae): Evidence from nulcear ITS ribosomal DNA sequence. Lindleyana 16: 94. Szlachetko, D. L. 1995. Systema Orchidalium. Fragm. Flor. Geobot., suppl. 3: 1-152. 2001a. Molecular systematics of the Oncidiinae base don evidence from four DNA sequence regions: expanded circumsciptions of Cyrtochilum, Erycina, Otoglossum, and Trichocentrum, and a new genus (Orchidaceae). Lindleyana 16: 113. Phylogenetic positions of Miltoniopsis, Caucaea, a new genus, Cyrtochiloides , and Oncidium phymatochilum (Orchidaceae: Oncdiinae) base don nuclear and plastid DNA). Lindleyana 16: 272.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.402 LANKESTERIANA

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CONFUSION IN EPIDENDRUM BRENESII SCHLTR., AND A NEW COSTA RICAN SPECIES: EPIDENDRUM SOTOANUM (ORCHIDACEAE) ADAM P. KARREMANS 1,3 & ERIC HGSATER 21 Jardn Botnico Lankester, Universidad de Costa Rica, Cartago, Costa Rica 23 Corresponding author: akarremans@gmail.com403. 2010. LANKESTERIANA 9(3): A couple of years ago, while collecting above his companions of the Lankester Botanical Gardens, collected a plant which resembled Epidendrum brenesii Schltr., because of its characteristic vertical growth habit, with successive lateral growths produced from the middle of the previous growth, and roots produced usually only from the base of the primary stem. After expected deep purple, velvety lip, but rather a larger, greenish yellow, glabrous lip, deeply cleft at the apex (Fig. 1). Hgsater, upon seeing the images, immediately remembered a manuscript sent to him in 1983 by Kerry Barringer and Eric Christenson, proposing a yellowEpidendrum as new, but which he considered at the time to corresponded to E. brenesii. The matter slept for 16 years in a drawer. Investigating the case further, all the material under E. brenesii was reevaluated to determine the presence of possible confused specimens. We that Luis Snchez, of AMO had also collected a images of the original material pressed by Barringer and Christenson, and several Skutch and Standley specimens, corresponding to the eastern part of the range of what went for E. brenesii , were all marked upper basin of the Ro Reventazn. More studied specimens collected further west all showed yellowIt was obvious that a detailed re-evaluation of the original description and tracings of the type of Epidendrum brenesii , lodged at AMES, was necessary (The original type material was destroyed in Berlin during the war). It immediately became evident that both the illustration and the description a relatively narrow, reniform, short-emarginate lip that corresponds to the purple-colored species. On the lip with a very wide sinus; the lines in the middle of ABSTRACT . Two species have been confused under the name Epidendrum brenesii Schltr., both were used in E. brenesii, a specimen collected by Brenes close to the original locus typicus is therefore used to neotypify Epidendrum sotoanum honor of the late Mexican botanist Miguel ngel Soto Arenas. KEY WORDS: Epidendrum brenesii , Epidendrum sotoanum, Costa Rica, Miguel ngel Soto Arenas FIGURE Epidendrum brenesii Schltr. (Bogarn 1513) and B – Epidendrum sotoanumBogarn 3920). Scale bar = 1 cm. Photo by F. Pupulin (A) and D. Bogarn (B).

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the lip suggest a pair of keels running down the disc to the apex; features which correspond to the yellowcolored species. The drawing of the plant habit clearly illustrates the typical architecture of this group within Epidendrum , but is not useful to distinguish between the two entities (Fig. 2). are light green, tinged with violet, which is what has traditionally been called E. brenesii. However, the text indicates that the lip is widely reniform, nearly twice as wide as it is long (15 x 8 mm), that it is widely 1923). Those features correspond to the yellowIt is thus evident to us that Schlechter made a composite description and the unpublished illustration of the floral parts is also a composite. It could have happened when the floral parts were put together for the purpose of publishing the floral analysis; the view of one flower got mounted with the floral segments of another. Later, when Santiago E. brenesii in Icones Orchidacearum, though the illustration was prepared from one single specimen of purple-colored flowers, they also prepared a composite description covering all the material then thought to belong to a single species, with either yellow or purple flowers. Another distinguishing feature is the surface of the lip in the purple-colored species, which is clearly microscope to be short-setose, densely covered by has a glabrous appearance, the surface covered with low, rounded papillae. Upon mapping all the collection localities, the purple-colored species appears to be endemic to the Cordillera Central, from the upper ridge south, while the yellow-colored species ranges along the Atlantic slope east as far as the upper basin of the Ro Reventazn. Both species are present in the region of Varablanca. After viewing all available evidence, we hereby with a neotype, using another specimen collected by Brenes near the type locality and located in the National Herbarium in San Jos, Costa Rica: FIGURE 2. Drawing of the holotype of Epidendrum brenesii (AMES), from Brenes 118. Courtesy of the Orchid Herbarium of Oakes Ames, Harvard University Herbaria.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.404 LANKESTERIANA

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Epidendrum brenesii Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19: 209. 1923. TYPE: COSTA RICA . Alajuela: San Pedro de San Ramn, 1050 m, November 1921, A. M. Brenes 118. Holotype: B (destroyed). Lectotype (designated by E. in S., Icon. Orch. (Mexico City) 8: pl. 814. 2006.): tracing Standley 36573, AMES 29339! (Cited by Barringer, 1986.), here rejected. Neotype (here designated): COSTA RICA: Alajuela: La Palma de San Ramn, 1250 m, 14 November 1922, Brenes 449(154), CR! Epiphytic, monopodial, erect herb , 30-70 cm tall. Roots basal from the initial stem and sometimes from an upper stem, 1.5-3 mm thick. Stems simple, canelike, terete, straight, the new stem originating from a middle internode of the previous stem, gradually becoming thinner and shorter, 5-30 x 0.2-0.5 cm. Leaves 3-5, aggregate along the apical half of the stem, alternate, articulate, erect-spreading, unequal in size, the basal one generally smaller, sub-coriaceous; with time, light green tinged with purple, 11-17 mm long; blade oblong, acute, margin entire, 2-8 x 1.32.5 cm. racemose to rarely paniculate, arching, 2-4 cm long; peduncle thin, laterally compressed, straight, light green tinged purple, 1.0-1.5 cm long. Floral bracts about half as long as the ovary, triangular, acuminate, tinged purple, 4-5 mm long. Flowers 8-12[40], successive, though several may be open at one time, resupinate, light green with purple tinge to deep purple with the column green; slight “banana-peal” fragrance at sundown. Ovary tinged purple, 8-13 mm long. Sepals partly spreading, free, obovate-elliptic, acute, slightly concave, glabrous, 3-veined, margin entire, spreading, dorsal sepal 12-13 x 6.0-6.5 mm, laterals oblique, 14.5-15.0 x 7.5 mm. Petals partly spreading, free, linear-oblanceolate, acute, 3-veined, margin entire, spreading, 12 x 2.5-3 mm. Lip united to the column, entire, reniform, slightly convex, velutinous, short setose, densely covered by short, pointed trichomes, emarginate, the apex sometimes slightly 4-lobed, margin entire, 8-12 x 15-19 mm; bicallose, the calli formed by a pair of short, laterally compressed thickenings; disc with a low, wide keel that reaches the apical sinus. Column thin at the base, gradually widened towards the apex, straight, 6.5 mm long; clinandrium-hood reduced, the margin erect, entire; rostellum apical, slit. Anther subreniforme, 4-celled. Pollinia 4, obovoid, laterally compressed, caudicles soft and granulose, as long as the pollinia. Nectary penetrating somewhat the ovary, smooth. MATERIAL STUDIED : COSTA RICA . Alajuela: La Palma de San Ramn, 1250 m, 8 III 1930, Brenes 11909a, NY! Valverde Vega, Bajos del Toro, Hacienda Ro Toro, Reserva Biolgica Privada Bosque de Paz, 1500-2000 m, cult. XII 2007, Karremans 2188, JBL-Spirit! Digital image, AMO! JBL! Sarapiqu, 8 II 1966, Lankester s.n. sub Rodriguez 366, USJ! Vara Blanca y Cariblanco, col. 19 III 1982, pressed cult. 23 I 1983, Hgsater 6822, AMO! Varablancacult. 22 II 1983, Hgsater 6824, AMO! (Illustration voucher, Fig. 3). Vara Blanca, 2 km noreste de Alto del Roble, faldas al norte del Cerro Chompipe, Refugio de Vida Silvestre Cerro Dantas, mrgenes de la Quebrada Grande, 10.5”N 84.4”W, 1981 m, 14 IV 2005, Bogarin 1513, JBL-Spirit! Digital image, AMO! JBL! Vara blanca, cerca del cruce a San Miguel, 1000 m, 28 X 1995, Morales 4966, INB! OTHER RECORDS : COSTA RICA . Alajuela: Valverde Vega, Bajos del Toro, Karremans s.n., digital photo AMO! JBL! Sarapiqu, 11 II 1966, Lankester s.n. sub Rodriguez 366, xerox of illustration, AMO! DISTRIBUTION AND ECOLOGY : Endemic to central Costa Rica, known only from the upper ridge of the Cordillera Central and south towards San Ramn, in humid premontane forest at 1000-2000 m altitude. Flowering from October to April. new: Epidendrum sotoanum sp. nov. TYPE: COSTA RICA : Cartago; Finca Kiri, camino a la Reserva Natural de Tapant, 1300-1400 m, col. 25 IX 1996, pressed 12 VI 1998, L. Snchez 401, J. Garca . Holotype: USJ! Clonotype: Ibid. pressed 19 V 1997, AMO!LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. KARREMANS & HGSATER. — Confusion in Epidendrum brenesii405

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FIGURE 3. Epidendrum brenesii Schltr. A – Flower. B – Habit. C – Column and lip, lateral view. D – Dissected perianth. E – Column, ventral view. F – Pollinarium. Drawn by E. Hgsater from Hgsater 6824 (AMO).LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.406 LANKESTERIANA

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maculatis, labello latiore profunde et late emarginato recedit. Plant epiphytic, monopodial, erect, branching, up to 50 cm tall. Roots basal, from the basal stem only, Stems each new stem produced from a subapical internode of the previous stem, below the leaves. Leaves few (3 in the holotype), aggregate towards the apex of the stem; sheath tubular, rugose; blades obovate-elliptic, obtuse, variable in size, the largest up to 5-10 x 1.32.5 cm. from each stem; peduncle laterally compressed and somewhat ancipitose, 2 cm long. Floral bracts half as long as the ovary, acute, 0.6-1 cm long. Flowers 4-15, greenish brown or yellow, column green, darker at the base. Ovary terete, smooth. Sepals partly spreading, obovate-elliptic, obtuse, 3-4-veined, margin entire, spreading; dorsal sepal 12-16 x 4.0-7.5 mm, lateral sepals oblique, 13-18 x 7-9 mm. Petals partly spreading, narrowly-obovate, obtuse, 3-veined, margin entire, spreading, 11-15 x 2-4 mm. Lip strongly bilobed, widely reniform, deeply emarginate, slightly convex in but with low rounded papilla when observed under a microscope, 10-17 x 20-30 mm; lateral lobes obscurely bilobed, sub-orbicular, sometimes emarginate; margin entire; bicallose, the calli thickened at the base and ending in a pair of low keels. Column somewhat arching downwards, short, thickened towards the apex, sub-terete, 7-8 x 5 mm above the middle, with apical rounded wings; clinandrium-hood erect, very short, margin entire; rostellum near the apex of the column, slit. Anther obovate, 4-celled, 2.5 mm wide. Pollinia 4, obovoid, laterally compressed. Nectary short, without penetrating the ovary, smooth. MATERIAL STUDIED : C OSTA RICA . Cartago: Cartago, San camino a Alto Beln, entre Sombrero y Quebrada Patarr, 9.3”N 83.1”W, 1430-1620 m, y rboles en zonas abiertas, 23 V 2007, Bogarn ; JBL-Spirit! Digital image, AMO! JBL! (Illustration voucher, Fig. 4). Ibid. Bogarn et al. 3920, JBLSpirit! Digital image, AMO! JBL! Orosi valley, in Rio Sombrero, 1500 m, 9’N, 831’W, 6 VI 1983, , F! Digital image and illustration, AMO! Heredia: San Rafael, Ro Tibas, toma de agua de Concepcin de San Rafael, 1700 m, 2 XII 1995, Lpiz 160, INB! Vara Blanca de Sarapiqu, between Pos and Barva volcanoes, 1770 m, II 1938, Skutch 3510, AMES! Ibid. 1950 m, II 1938, Skutch 3526, AMES! K! MO! Yerba Buena, NE of San Isidro, 2000 m, 22 II 1926, Standley 49987, AMES! US! Vara Blanca, 1400-1700 m, 27 VI 1978, Todzia 332, CR! San Jos: La Hondura, 1300-1700 m, 2 III 1924, Standley 36573, AMES! US! OTHER RECORDS : COSTA RICA . Alajuela: San Pedro de San Ramn, 1050 m, XI 1921, Brenes 118*, B (destroyed), Standley 36573, AMES 29339! (Cited as type of E. brenesii by Barringer, 1986 and designated as lectotype of by camino a Alto Beln, entre Sombrero y Quebrada Patarr, 9.3”N 83.1”W, 1430-1620 m, y rboles en zonas abiertas, 23 V 2007, D. Bogarn Digital image, AMO! JBL! DISTRIBUTION AND ECOLOGY : Endemic to Costa Rica, on the Cordillera Central, along the Atlantic slope, and east to the upper basin of the Ro Reventazn. Epiphytic in secondary humid premontane forest, at 1500-2000 m elevation. Flowering in DecemberJune. Epidendrum sotoanum belongs to the Arbuscula Group, Incomptum Subgroup, which is characterized by the successive lateral growths produced from the middle of the previous growth, few leaves which aggregate towards the apex of the stems, a short apical * The illustration and original description of E. brenesii E. brenesii ; the illustration of the plant habit is indistinguishable. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. KARREMANS & HGSATER. — Confusion in Epidendrum brenesii407

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FIGURE 4. Epidendrum sotoanum from Bogarn 3919 (JBL-Spirit). LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.408 LANKESTERIANA

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species is recognized by the large greenish yellow or papillose), very large, deeply emarginate lip, and the apical rounded wings of the column. It is similar and has been confused with Epidendrum brenesii Schltr. which is known from upper ridge of the Cordillera Central and south towards San Ramn, its a reniform, velutinous, short-emarginate lip, 15-19 mm wide. Epidendrum bisulcatum Ames, has shorter and narrower leaves, a smaller, sub-orbicular lip and linear petals. Epidendrum incomptum Rchh.f. which also occurs in Costa Rica can be distinguished by its 3-lobed, ecallose lip, the 3 lobes more or less triangular. Outside Costa Rica, only Epidendrum foldatsii the same architecture, and a bilobed lip, but the x 7.5 mm. We take pleasure in dedicating this species to our late fellow orchid researcher from Mexico, Miguel ngel Soto Arenas, (1963-2009) who shared his knowledge and experience with us. Apart from his many contributions to science, reading through the narrative of the different ecosystems in The Orchids of Mexico , gives us a glimpse of how well he came to understand the relationship between orchids and their various habitats, as well as the threats to biodiversity. ACKNOWLEDGEMENTS: We wish to thank Diego Bogarn (JBL) for sharing his material and knowledge of the species and preparing sketches of the new species, and Elizabeth Santiago (AMO) for once more going through all records, and ample discussion of the manuscript, and revision of the descriptions. We are also thankful to G. Wece (WUR) for the tactical support. LITERATURE CITED Barringer, K., 1986, Schlechter’s Costa Rican Types, Field. Bot. 17: 6-7. Hgsater, E., M. A. Soto Arenas, G. Salazar, R. Jimnez The Orchids of Mexico. Instituto Chinon, Mexico City. 304 pp. Epidendrum brenesii (Mexico) 8: 814. Schlechter, R. 1923. Epidendrum Brenesii Schltr. Repert. Spec. Nov. Regni Veg. Beih. 19: 209. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. KARREMANS & HGSATER. — Confusion in Epidendrum brenesii409

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ONCIDIUM ORNITHORHYNCHUM, UNA ESPECIE MAL INTERPRETADA Y UN NOMBRE PARA UNA VIEJA ESPECIE: ONCIDIUM SOTOANUM (ORCHIDACEAE) ROLANDO JIMNEZ MACHORRO & ERIC HGSATER 411. 2010. LANKESTERIANA 9(3): Una de las especies ms populares de Oncidium se conoce como “Oncidium ornithorhynchum”. Su arqueadas en todas direcciones, lo que forma un bouquet perfecto como centro de mesa. Ha sido utilizada tambin en la creacin de numerosos hbridos y clones seleccionados que han recibido reconocimientos en todo el mundo. Theodor Hartweg envi plantas colectadas en Oaxaca, Mxico, a Loddiges quien la introdujo al cultivo por primera vez en Europa en 1836. Ese de Guatemala, misma que ilustr en su famosa obra The Orchidaceae of Mexico and Guatemala con el nombre de Oncidium ornithorhyncum e incluyendo un comentario que posteriormente repiti Lindley (1840), haciendo referencia a la obra de Kunth, el cual menciona que hay copias coloreadas a mano donde el dibujo se hizo a partir de un espcimen seco. Segn en U, BM-Bloomsbury y la Biblioteca Nacional de Paris. En la primera entrega de The Orchidaceae of Mexico and Guatemala (1837) apareci la especie de Mxico y Centroamrica ilustrada por primera vez como “O. ornithorhynchum”. Es probable que la planta dibujada por Bateman haya sido determinada por Lindley, quien la public por segunda vez con una nueva ilustracin (Lindley, 1840). Es probable que el autor se haya dejado llevar por la ilustracin y la localidad de origen mencionada en la obra de Kunth. Adems en aqul el nmero de estas por seudobulbo. Todos los autores posteriores a Bateman aceptaron su ilustracin como la representacin de O. ornithorhynchum. En 1990 Miguel ngel Soto, tuvo la oportunidad de ver en el herbario de Pars el ejemplar tipo de Oncidium ornithorhynchum y se dio cuenta que la especie que tenemos en Mxico sorprendentemente no corresponda con dicho ejemplar. Lamentablemente del material. Sin embargo, el hecho qued grabado en RESUMEN . Se aclara la identidad de Oncidium ornithorhynchum Kunth, una especie mal interpretada, debido a que el tipo depositado en el herbario de Pars (P) no corresponde con la especie que ha recibido ese nombre en Mxico y Centroamrica, sino con un taxn de Ecuador, Colombia y Per conocido como O. pyramidale Lindl. Se propone como nueva a la especie de Mxico y Centroamrica: O. sotoanum y se describe una nueva subespecie: O. sotoanum ssp. papalosmum R.Jimnez, conocida nicamente de los estados de Oaxaca y Chiapas, Mxico. ABSTRACT . The true identity of Oncidium ornithorhynchum Paris herbarium (P). It does not correspond to the well known Mexican and Central American species, but to O. pyramidale Lindl. from Ecuador, Colombia and Peru. The Mexican and Central American species is proposed as new: O. sotoanum O. sotoanum ssp. papalosmum R.Jimnez, known only from the states of Oaxaca and Chiapas, Mexico. PALABRAS CLAVE / KEY WORDS : Oncidium, ornithorhynchum, sotoanum, papalosmum, pyramidale

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.412 LANKESTERIANA FIGURA 1. Foto del Tipo de Oncidium ornithorhynchum Kunth en el Herbario de Pars. Foto cortesa de A. Espejo Serna.

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su memoria y lo comentamos en varias ocasiones. Fue hasta 1997 que Adolfo Espejo, botnico mexicano, viaj a Pars y obtuvo una fotografa del ejemplar tipo de O. ornithorhynchum como parte de su investigacin de los tipos de las monocotiledneas mexicanas (Espejo et al. 1998), del cual amablemente nos proporcion una copia (Fig. 1). Al examinar los ejemplares tipo de los “oncidiums” respaldada por el tipo de Oncidium ornithorhynchum Kunth en el herbario de Paris (P), no concordaba con la especie que se distribuye en Mxico y Centroamrica y que se conoce universalmente con este nombre. Ya con la imagen del tipo en nuestras manos, se procedi a hacer un anlisis del ejemplar, as como para comparar este material con la planta mexicana que se conoce como O. ornithorhynchum. El material original muestra que las vainas foliosas de la base del seudobulbo tienen el margen membranceo muy erecta, con las ramas de la base alargadas, acortndose gradualmente hacia el pice (arquitectura piramidal), el pednculo y raquis son casi rectos, ligeramente sinuosos; el callo del labelo se compone de dos quillas cortas hacia los lados y otras tres hacia el pice; las alas de la columna son erectas y cuneadas. el callo amarillo; el margen de las vainas foliosas es membranceo pero este es inconspicuo, midiendo tan solo 1 mm de ancho. Cada seudobulbo produce las ramas de la base del raquis cortas, las de la mitad largas y las del pice cortas. El raquis es en zigzag. El callo es distinto y las alas son oblicuamente dolabriformes y paralelas al cuerpo de la columna. Llama la atencin que la localidad tipo de O. ornithorhynchum dada por Kunth, “El Puerto Andaracuas, entre Guanajuato y Valladolid, Michoacn”, es evidentemente errnea (Jimnez, 1990). Esa localidad corresponde a la provincia meridional y El Bajo, una zona muy seca, donde abundan los encinares secos, muy pobres en orqudeas Laelia speciosa (Hgsater et al. 2005). El ejemplar tipo (Fig. 1), tiene una etiqueta con la leyenda “America Ecuatorialis” por lo que no es difcil deducir que el ejemplar probablemente fue colectado en Sudamrica. De acuerdo con las caractersticas de alas de la columna erectas y la estructura del callo, las nicas especies que presentan stas son Oncidium pyramidale Lindl. que se distribuye en Per, Ecuador y Colombia, y Oncidium elephantotis Rchb.f. de Ecuador. Oncidium elefantotis (alrededor de 1.5 cm de largo en comparacin con las de Oncidium ornithorhynchum que miden alrededor de 2 cm de largo). Por su parte, Oncidium pyramidale del labelo y las alas de la columna coinciden con las del dibujo de la obra de Kunth. Por lo tanto, hemos llegado a la conclusin de que el verdadero Oncidium LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. JIMNEZ MACHORRO & HGSATER. — Oncidium sotoanum413 FIGURA original de Oncidium ornithorhynchum en la obra Nova Genera et Species Plantarum, la cual se bas en el ejemplar colectado por Humboldt y Bonpland depositado en el herbario de Pars.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.414 LANKESTERIANA FIGURA 3. Dibujo de Oncidium ornithorhynchum Kunth, basado en , Colombia, Cundinamarca (AMES). Dibujo de R. Jimnez M.

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ornithorhynchum es lo que conocemos como O. pyramidale Lindl. de Ecuador, Colombia y Per. En consecuencia se reduce O. pyramidale a sinnimo del primero. Oncidium ornithorhynchum Pl. 1: 345, t. 80. 1815. TIPO: Amrica Ecuatorialis, Humboldt s.n. (Holotipo: P);Crescit in temperatis Regni Mexicani prope El Puerto Andaracuas, inter urbes Guanaxuato et Valladolid de Mechoacan, alt. 1010 hex. Floret Septembri”. SIN”NIMO : Oncidium pyramidale Nat. Hist. 15: 384. 1845. TIPO: COLOMBIA . Pasto: Hartweg s.n. (Holotipo: K-L! Probable isotipo: Hartweg s.n. (ex herbario Lindleyano, W-R 47588, a los 3340 m de altitud. Florecen en enero. Oncidium ornithorhynchum erectas, angostamente cuneadas, con el pice bilobado (Fig. 3). Es probable que Humboldt y Bonpland hayan colectado el Oncidium ornithorhynchum en su viaje a Sudamrica y posteriormente con el transporte de las plantas hubo una confusin en la localidad. No hemos encontrado ningn registro de una colecta de que tenemos en Mxico y Centroamrica. ESPECMEN EXAMINADO . COLOMBIA . Cundinamarca: Pramo de Guasca, 2850-3340 m, Jaramillo 3211 (AMES!, testigo de ilustracin). Se ha mencionado en la literatura a O. chrysopyramis Rchb.f. como sinnimo de O. pyramidale, pero esta es una especie diferente distinguindose por la Mxico y Centroamrica como nueva: Oncidium sotoanum sp. nov. TIPO: MXICO . Veracruz: Orizaba, Cerro San Cristbal, al sur de Orizaba, a 500 m de la carretera que va a Lemboglossum [Rhynchostele] bictoniense y Encyclia [Prosthechea] vitellina amarillo-naranja, aroma intenso agradable, dulce, col. 16 abr 1987, prep. mat. cult. 23 oct 1990, R. Jimnez 626 (Holotipo: AMO, isotipos: K-L, MO, clonotipos: prep. mat. cult. 31 octubre 1987, F; prep. mat. cult. 31 oct. 1989, AMES). Species Oncidio ornithorhyncho Kunth similis, architecturae non pyramidalis, alis columnae parallelibus corpo columnae, callo labelli distincto recedit. SIN”NIMOS : Oncidium ornithorhynchum Auct., non Kunth. Oncidium ornithorhynchum var. Rchb.f., enviada a H. G. Reichenbach por Day, (W-R, no visto). Hier ba Races delgadas, 0.5-1.2 mm de grosor. Rizoma corto. Seudobulbos agregados, ovoideelipsoides, comprimidos, generalmente bifoliados, ocasionalmente trifoliados, lisos, sin costillas, con arrugas longitudinales con la edad, frecuentemente con un canal o surco lateral a ambos lados; verdeglaucos, speros, 4-9 x 2.0-3.5 cm; cubiertos en la base por 4-5 vainas subcoriceas, verde-claras, con margen eroso escarioso inconspicuo (ocupando lo escarioso 1 mm de ancho), 3.5-9.0 cm de largo, las dos superiores foliosas, con lmina foliar articulada, semejante a las hojas 28-45 x 1.5-3.2 cm. Hojas 2-3 en el pice del seudobulbo, erecto-arqueadas, linear-elpticas a lanceoladas u oblanceoladas, largamente acuminadas, subcoriceas, verde-claras, 16-45 x 1.5-3.3 cm. originada en la base del seudobulbo, aparece cuando el brote nuevo est en desarrollo, madura, 2 a 3 por seudobulbo, raramente 4, paniculada, de largo, verde a ligeramente rojizo; raquis en zigzag; provista de brcteas ovado triangulares, herbceas, membranceas, amplexicaules, las del pednculo tubulares en la base, de 10-17 mm de largo, las de LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. JIMNEZ MACHORRO & HGSATER. — Oncidium sotoanum415

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.416 LANKESTERIANA FIGURA 4. Oncidium sotoanum R. Jimnez 626, Orizaba, Veracruz, Mxico, AMO (ilustracin tomada de Icon. Orchid. (Mexico). 1: pl. 77).

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. JIMNEZ MACHORRO & HGSATER. — Oncidium sotoanum417la base de las ramas no tubulares; agudas, de 3-6 x 3-5 mm. cuculadas, triangulares a ovadas, herbceo-membranceas, amplexicaulas, agudas, 1.5-4.0 x 2.5-4.0 mm. Ovario pedicelado, tenuemente sulcado, 7-9 mm de largo, 0.8-1.0 mm de grosor. Flores vistosas, 16-18 mm de dimetro, spalos y ptalos rosados, labelo rosado ms intenso en la parte basal con los lbulos laterales blancos, callo amarilloanaranjado, columna blanca con el pice rosado, tbula infraestigmtica blanca, alas rosadas, antera guinda o intensamente rosada; fragancia intensa, dulce, diurna. Spalos unguiculados a atenuados, obtusos a redondeados, carinados dorsalmente, el dorsal largo; oblanceolado a obovado o elptico, mrgenes x 2.0-4.5 mm; los laterales ligeramente incurvados a extendidos, falcados, brevemente connados en la trinervados a 5-nervados, 8.0-11.5 x 2.0-3.5 mm. Ptalos incurvados a extendidos, oblicuos, cortamente a obovado elpticos o nicamente elpticos, obtusos, con mrgenes ondulados, carinados dorsalmente, 5-nervados, 6-10 x 3-5 mm. Labelo trilobado, plano, con la base cordiforme, formando un ngulo de 225 con respecto a la columna, 12-18 x 6-9 mm de ancho entre los lbulos laterales; stos envuelve la base de los spalos laterales para formar un pliegue que secreta un lquido oleaginoso, 3 x 4-7 mm; 2-3 mm de ancho; lbulo medio obovado a obcordado, pice hendido, bilobulado, mrgenes enteros, 5-7 x 4-7 mm de ancho; callo prominente, ocupando un cuarto de la longitud total del labelo, formado por 5 quillas lateralmente aplanadas, con el borde eroso, dos cortas hacia los extremos, dos laterales hacia el interior que rematan en el pice en un cuerno alargado agudo y una central que termina antes de los cuernos alargados agudos, 3.0-3.7 x 4.5-5.2 mm. Columna delgada, ms ancha en la base que en el pice, 4-7 mm de largo, alada, con tabula infraestigmtica ensanchada en la parte basal, sulcada; alas delgadas, subcuadradas u oblicuamente dolabriformes, margen sinuoso a eroso, paralelas a la columna, 1-2 x 2.02.5 mm; rostelo alargado, angostamente triangular; cavidad estigmtica subtriangular a subcuadrada, truncada, cncava, brillante, blanca; clinandrio ovado, guinda o rosa intenso, con una raya blanca que va desde el pice hasta la parte media. Antera claviforme, bilocular, pice largamente acuminado, 3.5-4.0 x 1.5-2.0 mm. Polinario de 3.5 mm de largo, formado por dos polinios sulcados, elptico obovoides, 0.7 x 0.5 mm; con estpite tubular, 2.3 mm de largo, muy alargado, que termina en una lmina triangular por la cual se sostienen los polinios por medio de las pardo. Cpsula elipsoide a obovoide, redonda en corte transversal, lisa, 1.3 x 1.1 cm, pedicelo 1 cm de largo (Fig. 4 y 5). ETIMOLOGA : Dedicamos esta especie a nuestro colaborador y maestro, recientemente fallecido, Miguel ngel Soto Arenas, por su dedicacin al conocimiento de la biodiversidad, particularmente de las orqudeas, por haber compartido sus conocimientos en salidas al campo y discusiones. FIGURA 5. Foto de Oncidium sotoanum Hgsater; M A. Soto 4900, Chiapas, Mxico. Foto de R. Jimnez M.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.418 LANKESTERIANA Se distribuye en Mxico, en la parte oriental del Eje Volcnico Transversal, Sierra Madre Oriental, Macizo Central de Chiapas y Sierra Madre de Chiapas en los estados de Veracruz, Tabasco, Oaxaca y Chiapas. Se ha registrado para Centroamrica en Guatemala, El Salvador, Nicaragua y Costa Rica. perennifolia, bosque caducifolio, bosque hmedo de encino y ocasionalmente en selva mediana subperennifolia (Montane rain-forest, Oak-pineLiquidambar forest y Evergreen seasonal forest [Breedlove, 1981]), son bosques clido templados crece a 890-1700 m de elevacin. Florece de octubre a Se reconoce por sus seudobulbos casi glaucos, aroma dulce, rosadas, siendo una de las pocas especies de ste gnero que presentan el color rosado, con el callo amarillo y por el rostelo alargado parecido al pico de un ave. Las plantas no se confunden con ninguna otra pueden confundir con Oncidium iricolor Rchb.f (sin.: O. pollardii rosadas con manchas y puntos morados o magenta, el labelo rosado, la base amarilla, con dos manchas anaranjado-rosadas en el disco a ambos lados del callo y Si bien se trata de una especie que por ahora no est amenazada, algunas poblaciones han disminuido o desaparecido debido a la destruccin intensiva de su hbitat por actividades humanas. Est sujeta a una fuerte presin de colecta, especialmente en el Estado de Veracruz. Es muy apreciada en horticultura por intensamente aromticas. Existen individuos albinos, muy raros, dentro de las poblaciones de Oncidium sotoanum, en uno de estos se bas H. G. Reichenbach para describir en el siglo antepasado. Los colores plidos son raros pero stos no son los ms atractivos; los ms vistosos son los colores ms oscuros que dan un buen contraste entre el callo amarillo y el labelo. ES PE C ME N E S : MXICO . Sin localidad: 1905, C.A. Purpus 44 , (W-R, diapositiva AMO!); “Presented by Messrs. F. Nov 1886, H. Veitch s.n. (K-L, diapositiva AMO!). Veracruz : Orizaba, Escamela, 30 sept 1865-1866, E. Bourgeau 3234 (NY, Px2, BRx2, K-L, diapositivas J. Linden 195 (K-L, diapositiva AMO!); sin localidad precisa, S. Rosillo de V. 99 (AMO!). Tabasco : Teapa, en el Ro Puyacatengo del cruce-UACH, 6 may 1988, A. Guadarrama et al. 1018 (MEXU!). Oaxaca : Sin localidad precisa, 1840, H. Galeotti 5019 (P, K-L, BRx3, diapositivas AMO!); km 70 de la carretera Oaxaca-Tuxtepec, despus de La Esperanza rumbo a Tuxtepec, 1260 m, 13 nov 1987, R. Jimnez 792 et al. (AMO!); ibid. , prep. mat. cult. 10 nov 1992, R. Jimnez M. 795 et al. (AMO!); ibid. , prep. mat. cult. 10 ago 1990, (AMOx2!); Sierra San Pedro Nolasco, Talea, 1843-4, C. Jrgensen 644 (BM, diapositiva AMO!); Ixtln entre Puerto Eligio a Comaltepec, km 148 entre Tuxtepec y Comaltepec, 26 oct 1965, G. Martnez-Caldern 436 (MEXUx2!); ibid. , km 149, 16 nov 1965, G. Martnez-Caldern 503 de cerro de la Divisin, cerca de 5 km al E de Benito Jurez, cerca de la frontera con Chiapas, cerca de 41 km en lnea recta al NNE de San Pedro Tapanatepec, 10 nov 1984, S. Maya 881 (MEXU!); ibid., arroyo El Caracol, parte inferior, al O de Benito Jurez, cerca de 39-40 km en lnea recta al N de Tapanatepec, 15 nov 1984, S. Maya 912 (MEXU!); San Miguel Chimalapa, Filo de la Culebra, entre Cerro Guayabito y Cerro Salomn, ca. 7 km en lnea recta al NW de benito Jurez, ca. 42 km en lnea recta al N de San Pedro Tapanatepec, 25 nov 1985, S. Maya 2571 (CHAPA!); San Jos Tenango, Teotitln a 3 km de San Martn, 1500 m, 26 oct 1992, S. Salas 335 (AMO!); Santa Mara, al pie de la vertiente N de la Sierra de Tres Picos, 1 abr 1996, G.A. Salazar 5526 (AMO!); camino Oaxaca-Tuxtepec km 114, 4500’, oct 1977, Thurston 1739 sub E. Hgsater 5436 (AMO!). Chiapas : Camino Palenque Ocosingo, km 15, prep. mat. cult. 24 dic 1976, E. Hgsater 2359 (AMOx3!); Las Margaritas, a 12 km al E de Lago Tziscao camino a Ixcan, 1250 m, 16 nov 1984, E. Martnez 8631 (AMO!); monte Ovando, 14 nov 1939, E. Matuda 4019 (MEXUx2!); Lagos de Montebello, M. Rodrguez s.n. , (AMO!); ca. km 3 del camino de Tziscao-Mrquez de

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. JIMNEZ MACHORRO & HGSATER. — Oncidium sotoanum419Comillas, cerca de los lmites del Parque Lagunas de Montebello, 13 oct 1985, M.A. Soto 1644 et al. (AMO!); Ocosingo, alrededores de la Laguna Ocotalito entre Mezabok y Monte Lbano, 950 m, 16 jun 1986, M.A. 2691 (AMOx3!); 500 m debajo de Rayn, 1340 m, 10 abr 1989, 4900 (AMO!); Las Margaritas, ca. de Nuevo Momn, 41.28 km al E de Las Margaritas, por el camino a Cruz del Rosario, 1500-1600 m, 9 ago 1992, Solano 7163 (AMO!); La Independencia, km 41.7 del camino Montebello-Mrquez de Comillas, 17 jul 1999, 9040 (AMOx3!); Union Jurez, 16 nov 1985, 2733 (ENCB!); between southern slopes Sierra de Soconusco to crest, 6 Nov 1945, (MEXUx3!). EL SALVADOR : sin localidad, prep. mat. cult. 4 dic 1998, E. Salaverria sub E. Hgsater 7845 (AMOx2!); Ibid., prep. mat. cult. 24 nov 2000, (AMO!). OTROS REGISTROS : MXICO . Veracruz: Orizaba, nov O. Nagel sub Oestlund 1761 (dibujos y datos [notas E. Oestlund] AMO!). En los estados de Oaxaca y Chiapas, existe una poblacin de Oncidium sotoanum callo, la cual se propone a continuacin como nueva para el cultivo. Oncidium sotoanum papalosmum R.Jimnez, ssp. nov. TIPO: MXICO . Oaxaca: km 178 del camino OaxacaPto. Escondido, “Ro salado”, 1350 m. Pinar de Pinus oocarpa Restrepia xanthophthalma [muscifera], Mexicoa [Oncidium] ghiesbregtiana, Trichocentrum hoegii, Trichosalpinx blaisdelli y Oncidium karwinskii, eptita. Tpalos lila oscuro, labelo lila plido, callo amarillo intenso, aroma dulce, intenso (tarde). Escaso en la localidad, noviembre 1987, prep. mat. cult. 10 ago 1990, M.A. . (Holotipo: AMO, Isotipo: AMES; testigo de ilustracin). Formae typicae similis, plantis minoribus, Mexicanorum Oaxacensis et Chiapasensis distributa. Hier ba Races delgadas, 0.5-1.0 mm de grosor. Rizoma corto. Seudobulbos agregados, ovoide-elipsoides, comprimidos, bifoliados, lisos, sin costillas, con arrugas longitudinales con la edad, frecuentemente con un canal o surco lateral a ambos lados; verde-glaucos, speros, 3.0-6.5 x 1-3 cm; cubiertos en la base por 3-5 vainas subcoriceas, verdeclaras, con margen eroso-escarioso inconspicuo, 1.54.5 cm de largo, las dos o tres superiores foliosas, con lmina foliar articulada, semejante a las hojas, 4-22 x 0.8-3.0 cm. Hojas 2 en el pice del seudobulbo, erecto arqueadas, linear elpticas a lanceoladas, acuminadas, subcoriceas, verde-claras, 9-26 x 1.0-2.8 cm. originada en la base del seudobulbo, aparece cuando el brote nuevo est en desarrollo, madura, 1 a 2 por seudobulbo, paniculada, erectosimultneas, con pednculo de 4-6 cm de largo, provista de brcteas ovado triangulares, herbceas, membranceas, amplexicaules, las del pednculo tubulares en la base, 0.6-1.3 mm de largo, las de la base de las ramas no tubulares, agudas, 6 x 3 mm. cuculadas, triangulares a ovadas, herbceo-membranosas, amplexicaulas, agudas, 2 x 4 mm. Ovario pedicelado, tenuemente sulcado, 7-17 mm de largo, 0.5-1.0 mm de grosor. Flores vistosas, 16-27 mm de dimetro, spalos y ptalos rosados, labelo rosado, magenta-oscuro intenso en la base, con lbulos laterales blancos, callo amarillo, manchado de anaranjado, columna blanca con el pice rosado, tabula infraestigmtica blanca, alas rosadas, antera guinda o intensamente rosada; fragancia intensa, dulce, a Ppalo (hierba del gnero Phorophyllum ), diurna. Spalos unguiculados, obtusos a redondeados, carinados dorsalmente, 5-nervados, el dorsal mm de largo; oblanceolado a obovado, mrgenes

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.420 LANKESTERIANA FIGURA 6. Oncidium sotoanum papalosmum R.Jimnez, basado en M.A. Soto 2803, Chiapas, Mxico. Dibujo de R. Jimnez M.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. JIMNEZ MACHORRO & HGSATER. — Oncidium sotoanum421ligeramente ondulados, 5-nervados, 7.5-10.0 x 2.5-5 mm; los laterales extendidos, falcados, brevemente oblanceolado-elpticos, 5-nervados, 8-12 x 2.5-4 mm. Ptalos de incurvados a extendidos, oblicuos, ovado-elpticos, redondeados, mrgenes ondulados, carinados dorsalmente, 5-nervados, 7-10 x 4-5 mm. Labelo la base cordiforme, 13-16 x 5.5-11 mm de ancho entre los lbulos laterales; stos semiorbiculares de los spalos laterales, y en donde se localiza un pliegue que secreta un lquido aceitoso, 3 x 4 mm; 3 mm de ancho; lbulo medio obovado a obcordado, pice hendido, bilobulado, mrgenes enteros, 5-6.5 x 5-6.5 mm; callo prominente, ocupando un tercio de la longitud total del labelo, formado por 5 quillas lateralmente aplanadas, dentadas, dos cortas en los extremos, dos laterales hacia el interior que terminan en el pice en un cuerno agudo, recurvado y una central baja que termina antes de los cuernos agudos, 3-4 x 3-5 mm. Columna delgada, ms ancha en la base q ue en el pice, delgada, 4-7 mm de largo, alada, con tabula infraestigmtica ensanchada en la parte basal, sulcada; alas delgadas, semiovadas o dolabriformes, margen sinuoso a eroso, paralelas a la columna, 1-2 x 2-2.5 mm; rostelo alargado, angostamente triangular; cavidad estigmtica subtriangular truncada, cncava, brillante, blanca; clinandrio ovado, color guinda o rosa intenso, con una raya blanca que va desde el pice hasta la parte media. Antera claviforme, bilocular, pice largamente acuminado, papilosa, 3.5-4 x 1.5-2 mm. Polinario de 3-3.5 mm de largo, formado por dos polinios sulcados, elptico obovoides, 0.7 x 0.5 mm; con estpite tubular, 2.3-2.8 mm de largo, muy alargado, que termina en una lmina triangular, por la cual se sostienen los polinios, por medio de las pardo. Cpsula no vista (Fig. 6 y 7). ETI M OLO G A . El nombre “ pa palosmum ” proviene del Nahuatl papaloquilitl (plantas del gnero Phorophyllum cocina tradicional y conocida coloquial mente como “Ppalo”. Se distribuye en las laderas de la Sierra Madre del Sur y la Sierra Madre de Chiapas en su vertiente Florece de agosto a diciembre. 30 cm de largo, con ramas fuertemente en zig-zag; intenso en la base; por el aroma herbceo, parecido a una planta de la familia de las Compuestas del gnero Phorophyllum , la cual es comestible y en Mxico recibe el nombre comn de “Papaloquelite” o “Ppalo”, muy distinto al aroma dulce intenso de la especie; por las alas de la columna semiovadas o dolabriformes y por su distribucin en las laderas do s de Oaxaca y Chiapas. OTROS E S P E C ME N E S . MXICO. Oaxaca: Entre Finca Copalita y Pluma Hidalgo, 950-1100 m, 11 may 1994, M.A. Soto 7617 et al. (AMO!). Chiapas: Jiquipilas, Cerro La Palmita, 1340 m, 7 nov 1994, J. J. Castillo et al. 403 (AMO !); sin localidad precisa, prep. mat. cult. FIGURA 7. Oncidium sotoanum ssp. papalosmum, M.A. Soto 2803, Chiapas, Mxico. Foto de R. Jimnez M.

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LANKESTERIANA21 oct 1981, E. Hgsater 536 (AMO!); Mapastepec, Reserva El Triunfo, 1200 m, 25 XI 1989, M. Heath (AMO!); Parte media del Cerro Tres Picos, ladera norte, al SE de Villa Flores, 1300 m, 10 VI 1986, M.A. Soto 2803 et al. (AMO!, testigo de ilustracin); Cerro Tres Picos, 47-48 km de la desviacin Cintalapa-Arriaga, ca. 1200 m, 13 VII 1989, prep. mat. cult. 23 dic 1987, M.A. Soto 3193 et al. (AMO!). AGRADECIMIENTOS . Deseamos agradecer al Dr. Gustavo Romero del herbario AMES, por el prstamo de material de Oncidium piramidale y la informacin amablemente proporcionada sobre la obra de Kunth. Al Dr. Adolfo Espejo imagen del tipo de O. ornithorhynchum del herbario de Paris (P). A Astrid H. de Restrepo de la Sociedad Colombiana de Orquideologa por la imagen de O. piramidale. Al Dr. Rodolfo Solano del CIIDIR-Oaxaca, la revisin crtica del manuscrito y sus valiosas sugerencias. A Franco Pupulin por las diagnosis latinas. LITERATURA CITADA Bateman, J. 1837-1842. Oncidium ornithorhynchum. Pl. 4 in: The Orchidaceae of Mexico and Guatemala Johnson Reprint Corporation, New York. Francisco. , 1: Lista de referencia, Parte VII. Orchidaceae (2). Consejo de la Flora de Mxico/Universidad Autnoma Metropolitana-Iztapalapa/ CONABIO, Mxico D.F. Orqudeas de Mxico. Instituto Chinon, Mxico, 304 p. Kunth, K.S.1815. Nova Genera et Species Plantarum. Paris. Jimnez M., R., 1990. Ondidium ornithorhynchum. Pl. 77 in Icon. Orchid. (Mexico) 1. Jimnez M, R. 2008. Una revisin del gnero Oncidium Sw. (Orchidaceae) en Mxico, Tesis de Licenciatura, Facultad de Lindley, J. 1840. Oncidium ornithorhynchum. Edward’s Bot. Reg. 26: t.10. dates, commentaries and types. Vol. II: H-Le. 2nd publishers, The Hague. Pp 369-370.422LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.

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TWO NEW SPECIES OF LEPANTHES FROM COSTA RICA CLOSE TO L. SCHIZOCARDIA (ORCHIDACEAE: PLEUROTHALLIDINAE) FRANCO PUPULIN 1 , DIEGO BOGARN 1,3 & CHRISTINA M. SMITH 11Lankester Botanical Garden, University of Costa Rica2ngel Andreetta Research Center on Andean Orchids, University Alfredo Prez Guerrero, Ecuador3Harvard University Herbaria, Cambridge, MA, U.S.A.4Corresponding author: franco.pupulin@ucr.ac.cr 423. 2010. LANKESTERIANA 9(3): The genus Lepanthes Sw. is probably the most the largest plant genera in the world, with more than 700 species distributed in the West Indies and from Mexico to Bolivia and Brazil. According to the geographic records, most of the species are very restricted in distribution, often with narrow regional of comparing an unusually large number of taxa, species of Lepanthes are mostly easy to identify on generalized morphological scheme of Lepanthes a lip provided with thickened, erect lateral lobes and a minute apical appendix) has impeded until recently to the grouping proposed by Luer, who divided the genus into subgenera, sections, subsections and series (Luer 1986, 1987a, 1987b, 1993, 1996), most of the taxa are assigned to subgen. Lepanthes sect. Lepanthes subsect. Lepanthes ser. Lepanthes. This large group, mostly characterized by a congested raceme with morphology, but several informal groups of species can be recognized by some peculiar combinations of characters. ABSTRACT . We revise the group of species close to Lepanthes schizocardia, and we describe and illustrate two new species from Costa Rica. Lepanthes montis-narae is compared with the Panamanian L. schizocardia, from which it mainly differs by the pendent plants, the dark gray green leaves, the ovate to orbicular, subacuminate lower lobe smaller than the upper lobe, narrowly ovate, the scarlet lip and the conic, apically ciliate appendix. Lepanthes sotoana is similar to L. maxillaris from Ecuador. It can be distinguished by the pendent plants, the L. schizocardia to facilitate species comparison. RESUMEN . Revisamos el grupo de especies cercanas a Lepanthes schizocardia y describimos e ilustramos dos nuevas especies de Costa Rica. Lepanthes montis-narae L. schizocardia, de la cual se diferencia principalmente por las plantas pendulosas, las hojas verde oscuro grisceas, las hojas ligeramente ovado, el labelo escarlata y el apndice cnico, apicalmente ciliado. Lepanthes sotoana es similar a L. maxillaris de Ecuador. Puede ser distinguida por las plantas pendulosas, las hojas verde oscuro grisceas, laterales subfalcados que se tocan el uno al otro y esconden la base del apndice. Tambin ilustramos por primera vez a L. schizocardia para facilitar la comparacin de las especies. KEY WORDS : Orchidaceae, Pleurothallidinae, Lepanthes montis-narae, L. sotoana, L. schizocardia, L. maxillaris, new species, Costa Rica

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FIGURE 1. Lepanthes schizocardia Luer. A — Habit. B — Flower. C — Dissected perianth. D — Ovary, column and lip. lateral view. E — Lip, adaxial view. F — Pollinarium and anther cap. Drawing by F. Pupulin and C. M. Smith based on D. Bogarn 5940 (JBL).LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.424 LANKESTERIANA

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN et al .— The Lepanthes schizocardia group425 We refer here to one of these morphological alliances with the name of Lepanthes schizocardia group, characterized by a cleft lip with the blades scarcely distinct from the connectives, sometimes mandible-like in shape, and particularly long the group, L. schizocardia, from Panama (Luer 1984), characteristic, cleft, heart-shaped lip. The species was not illustrated with the protologue (Luer 1984), and we take this opportunity to present here a composite plate based on Panamanian material originally collected at the type locality (Fig. 1, 4). Two closely allied species (L. maxillaris L. didyma Hirtz) were subsequently described from Ecuador (Luer 1996). The group ranges to the North to Costa Rica, where a few collections were previously Costa Rica and in the living collection of Lankester Botanical Garden (LBG) as L. schizocardia. In 2007, researchers at LBG began working to a complete survey of the species of Pleurothallidinae in Costa Rica, eventually intended as a contribution to the Flora Costaricensis. In the framework of this project, we are critically revising the identity of the species previous recorded from Costa Rica, comparing specimens from local populations with the original materials and, whenever possible, with living plants of the concerned taxa, preferably from type localities. combined with extensive examination of natural orchid populations are many times either found only in a narrow geographical range or in reduced numbers it is essential that studies be carried out over a broad geographical areas in a thorough manner. In revising Costa Rican materials previously assigned to L. schizocardia, we discovered two species that we described here as new to science. The main differences between the three taxa are summarized in Table 1. Lepanthes montis-narae Smith, sp. nov. TYPE: COSTA RICA . San Jos: Dota, crest of Cerro Nara, 910-1000 m, epiphytic in premontane rain forest, disturbed primary forest, 20 Feb. 2000, F. McFarland (holotype: JBL; isotype: USJ). FIG. 2, 4. A Lepanthes schizocardia Luer similis, sed planta pendula, foliis fuscatis viridi-griseus ovatisportatis abaxialibus, lobulis petalorum inaequalibus, lobo superno ovato acuto, lobo infero minore, anguste ovato, labello scarlatino appendice conica ciliata recedit. Epiphytic, caespitose, pendent herb, up to 14.5 cm tall. Roots Ramicauls pendent, up to 10.5 cm long, enclosed by 9 – 11 glabrous, lepanthiform sheaths, the ostia glabrous. Leaves coriaceous, dark green, ovate to orbicular, conduplicate, subacuminate, emarginate, with a short apiculus, 3.0.0 x 1.4.0 cm, the TABLE 1: Comparison of characteristics of the different taxa. Character L. montis-narae L. schizocardia L. sotoana Habit pendent erect pendent Leaves not reticulated reticulated not reticulated ovate to orbicular, subacuminate elliptic, acute ovate to orbicular, subacuminate Apical petal lobe ovate oblong oblong Lower petal lobe smaller than upper lobe equal to upper lobe equal to upper lobe Lip lobes hiding the column hiding the column not hiding the column Lip color scarlet orange scarlet Lip appendix cylindric, ciliate oblong, bristly oblong, ciliate

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FIGURE 2. Lepanthes montis-narae side view. D — Dissected perianth. E — Lip, frontal view. F — Lip, lateral views. G — Petals, lateral views. H — Pollinarium and anther cap. Drawing by F. Pupulin and C. M. Smith from the holotype.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.426 LANKESTERIANA

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN et al .— The Lepanthes schizocardia group427rounded base narrowing into a petiole less than 1 mm long. racemose, distichous, glabrous, than leaves, up to 2.8 cm long, peduncle 1.1.9 cm long, rachis 0.3.2 cm long. Floral bracts 1 mm long, glabrous. Pedicels 2.5.0 mm long, persistent. Ovary to 2 mm long. Flowers orange with red-scarlet, the sepals orange tinged with red, the petals red-scarlet, the lip scarlet. Dorsal sepal ovate, acute, connate to the lateral sepals for about 0.4 mm, 3.0 x 1.5 mm. Lateral sepals ovate to elliptic, acute, connate for 0.9 mm, 2.6 x 1.2 mm. Petals transversely bilobed, entire, 1.1 x 3.1mm, the upper lobe ovate, obtuse, the lower lobe smaller than the upper lobe, narrowly oblong, the apex rounded. Lip bilobate, adnate to the column, the blades oblong with rounded ends and the apex entire, falcate, 1.3 x 0.9 mm, the connectives terete, to 1 mm long, lifting the blades above the column, the body oblong, connate to the base of the column, the appendix small, oblong, pubescent, slightly curved upward. Column cylindric, 0.7mm long, the anther apical, the stigma ventral. Pollinia two, ovoid. Anther cap cucullate. PARATYPE : COSTA RICA . San Jos: Dota, crest of Cerro Nara, 910-1000 m, epiphytic in premontane rain forest, disturbed primary forest, 20 Feb. 2000, McFarland (JBL-spirit, USJ!). DISTRIBUTION: only known from Costa Rica. HABITAT AND ECOLOGY. The only known specimens were found epiphytic in disturbed primary vegetation in premontane rain forest, between 900 and 1000 m of elevation. Apparently, it is restricted to the crest of ETYMOLOGY: Named after Cerro Nara, the mountain where the type specimen was collected. Lepanthes montis-narae is most similar to the Panamanian endemic Lepanthes schizocardia Luer. It can be distinguished by the pendent plants (vs. erect), the dark gray green leaves (vs. reticulated in purple beneath), ovate to orbicular, subacuminate leaves (vs. (vs. beneath the leaf), the petals with unequal lobes (vs. subequal), the apical lobe ovate (vs. oblong), the lower lobe smaller than the upper lobe, narrowly ovate (vs. as long as the upper lobe, ovate), the scarlet lip (vs. orange) and the conic, apically ciliate appendix of the lip (vs. ovate, apically bristly). Lepanthes sotoana sp. nov. TYPE : COSTA RICA secondary vegetation and remnants of primary, lower montane rain forest, 2 December 2001, F. Pupulin (holotype: JBL; isotypes: CR, JBL). FIG. 3, 4. A Lepanthes maxillaris Luer similis, sed planta folii portatis abaxialibus, sepalis lateralibus acutis, lobis lateralibus labelli subfalcatis apicalibuis contiguis, base appendicis occulta. Epiphytic, caespitose, pendent herb, up to 9.3 cm tall. Roots Ramicauls slender, pendent, up to 5 cm long, enclosed by 4 glabrous, lepanthiform sheaths, the ostia glabrous and non dilated. Leaves coriaceous, dark grayish green, lightly suffused with purple beneath, ovate to orbicular, conduplicate, acute, emarginated with a short apiculus, 2.2.0 x 1.2 – 2.6 cm, the rounded base narrowing into a petiole less than 1 mm long. racemose, distichous, glabrous, 1.5 cm, peduncle 0.8.2 cm long, rachis 0.3.5 cm long. Floral bracts 1 mm long, glabrous. Pedicels 3 mm long, persistent. Ovary to 2 mm long. Flowers orange with red-scarlet, the sepals orange tinged with red, the petals red-scarlet. Dorsal sepal ovate, acute, connate to the lateral sepals for about 0.6 mm, 3.4 x 1.7 mm. Lateral sepals ovate, acute, connate for 0,7 mm, 2.8 x 1.4 mm. Petals transversely bilobed, entire, 0.8 x 2.7 mm, the upper lobe oblong, obtuse, the lower lobe, oblong to ovate, obtuse. Lip bilobate, adnate to the column, the blades oblong with rounded ends and the apex ciliate, subfalcate, 1.5 x 1.3 mm, the connectives terete, to 1 mm long, perpendicular to the column and leaving it exposed, the body oblong, connate to the base of the column, the appendix small, oblong, convex, ciliate, with

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.428 LANKESTERIANA FIGURE 3. Lepanthes sotoana D — Dissected perianth. E — Lip, frontal view. F — Lip, lateral views. G — Petals, lateral views. H — Pollinarium and anther cap. Drawing by F. Pupulin and C. M. Smith from the holotype.

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a short apicule. Column cylindric, 1 mm long, the anther apical, the stigma ventral. Pollinia two, ovoid. Anther cap cucullate. PARATYPES . COSTA RICA Berln, november 1998, collected by G. Chacn Valverde s.n. number JBL-00269, 19 september 2002, JBL-00269 Cruz, 1350 m, epiphytic on secondary vegetation and remnants of primary, lower montane rain forest, 2 December 2001, F. Pupulin 3483, D. Castelfranco in cultivation at San Isidro del General, 2 May 2000, F. Pupulin 2319 (USJ). DISTRIBUTION: only known from Costa Rica. HABITAT AND ECOLOGY: Plants of this species are found in tropical wet forest, tropical moist forest, premontane belt transition and premontane rain forest, between 800 and 1400 m of elevation. The species is apparently restricted to the El General slope of Talamanca range in Costa Rica. EPONYMY: of the late Miguel ngel Soto Arenas, a Mexican orchidologist who made great contributions to the knowledge of the family in the Neotropics. Lepanthes sotoana is closely related to Lepanthes maxillaris Luer from Ecuador. It can be distinguished by the pendent plants (vs. erect), the dark gray green leaves (vs. reticulated in purple beneath), the leaf), the acute lateral sepals (vs. acuminate), the lip with subfalcate lateral lobes touching each other, hiding the base of the appendix (vs. falcate, separated, leaving exposed the appendix). Lepanthes montis-narae and L. sotoana had been collected and cultivated at Lankester Botanical those of L. schizocardia. After studying the type of L. schizocardia, we found both species consistently different. The plants are pendent, lacking the purple borne above the leaf. Lepanthes sotoana can be distinguished from L. montis-narae by the oblong petals (vs. ovate), the lateral lobes of the lip touching apically each other and do not hide the column, which is placed just above the lip (lateral lobes of the lip hide the column and do not touch each other apically) and the appendix of the lip is conic (vs. oblong) and conspicuous, up to 1 mm long (vs. inconspicuous, less than 1 mm long). We have not seen specimens of Lepanthes schizocardia in Costa Rica. ACKNOWLEDGEMENTS . services of Ministerio del Ambiente, Energa y Telecomunicaciones de Costa Rica (MINAET) and Sistema Nacional de reas de Conservacin (SINAC) for issuing in this study were collected. The present paper is part of by the Vice-Presidency of Research, University of Costa Rica. LITERATURE CITED Luer, C. A. 1984. Miscellaneous new species in the Pleurothallidinae. Phytologia 55(3): 175. Luer, C. A. 1986. Icones Pleurothallidinarum I. Systematics of the Pleurothallidinae. Monogr. Syst. Bot. Missouri Bot. Gard. 15: 29. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN et al .— The Lepanthes schizocardia group429 FIGURE 4. Lepanthes sotoana A — Lepanthes schizocardia (Bogarn 5940). B — L. montis-narae (Pupulin et al. 2103). C — L. sotoana (Pupulin et al. 3484 ). Scale bar = 5 mm.

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Luer, C. A. 1987a. New Lepanthes species from Ecuador — 4. Lindleyana 2(2): 9511. Luer, C. A. 1987b. New Lepanthes species from Ecuador — 5. Lindleyana 2(3): 125. Luer, C. A. 1993. New series and species of Lepanthes from Ecuador. Novon 3: 442. Luer, C. A. 1996. Icones Pleurothallidinarum, XIV. The genus Lepanthes, subgenus Lepanthes in Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 61: 1.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.430 LANKESTERIANA

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STANHOPEINAE MESOAMERICANAE, V. EL AROMA FLORAL DE LAS STANHOPEAS DE MEXICO GNTER GERLACH Botanischer Garten Mnchen-Nymphenburg, Menzinger Str. 65, 80638 Mnchen, Deutschland gerlach@extern.lrz-muenchen.de431. 2010. LANKESTERIANA 9(3): Este artculo est dedicado a Miguel ngel Soto Arenas quien falleci victima de un asesinato. Miguel era un gran amante de las Stanhopea y por esta razn contribuy bastante al conocimiento de este gnero. l no solamente es autor de dos especies de Stanhopea de Mexico (Stanhopea dodsoniana Arenas, Stanhopea whittenii G.Gerlach), sino tambin, ha visto y colectado la gran mayora de ellas en su habitat natural. Sin l mi parte para la Flora Mesoamericana hubiera sido muy pobre, porque l puso a mi disposicin los datos basicos de las especies de Stanhopea de Mxico. Pens en publicar Stanhopea de Mxico al terminar de investigar todas las especies, pero la muerte de Miguel me impidi de hacerlo; ahora sin incluir la S. maculosa, la cual todavia no ha publicar el trabajo prematuramente. Indroduccin El gnero Stanhopea est presente en Mxico con 13 especies aqui reconocidas (Tab. 1). Todos pertenecen al subgnero Stanhopea y aqu a las secciones Saccata y Wardii. Mxico representa un segundo centro de diversidad del gnero; el otro es en Colombia con 19 especies de los 3 subgneros y en las 4 secciones. Con Stanhopea maculosa, la subtribu Stanhopeinae encuentra su lmite norte de distribucin en el Edo. de Sonora, en los bosques de encinos y pinos en un habitat con un perodo de sequa muy pronunciado. Todo el resto de las especies de Stanhopea de Mxico se encuentra meridional del Trpico de Cncer, en climas calientes o templados, en bosques y selvas hmedas. Las especies de Stanhopea de Mxico segn Gerlach (1999) se agrupan en: Seccin Saccata1S. hernandezii [= S. cavendishii Lindl. ex W.Baxter, S. devoniensis Lindl., S. expansa P.N.Don, S. lyncea (Lindl.) P.N.Don] S. intermedia Klinge (= S. novogaliciana S.Rosillo) S. maculosa = S. fregeana Rchb.f., S. marshii Rchb.f., S. schilleriana Rchb.f.) S. martiana Lindl. (= S. implicata Westc. ex Lindl., S. velata C.Morren) S. pseudoradiosa Jenny ABSTRACT Stanhopea from Mxico are presented here. Some of the species, such as S. hernandezii, S. martiana, S. tigrina, S. intermedia and S. graveolens are S. ruckeri and S. intermedia have fragrances, just as their taxonomy and morphology also are not easily described and clear. . RESUMEN Stanhopea de Mxico. Algunas, como S. hernandezii, S. martiana, S. tigrina, S. intermedia y S. graveolens, se pueden caracterizarar muy fcilmente por su olor, mientras que otras dos especies, S. ruckeri y S. intermedia, presentan dos quimotipos. Adems, existen dos grupos con tres especies cada uno que no se delimitan por sus aromas; igual, su taxonoma y morfologa no es fcil y clara. PALABRAS CLAVE / KEY WORDS : Orchids, Orqudeas, Stanhopea 1 La lista de sinnimos no est completa. Se mencionan solamente los ms conocidos y usados.

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S. radiosa Lem. S. saccata Batem. S. tigrina Batem. ex Lindl. [= S. nigroviolacea (C.Morren) Beer] Seccin Wardii S. dodsoniana S. graveolens Lindl. (= S. aurata Beer, S. venusta Lindl.) S. oculata (= S. minor Schltr., S. purpusii Schltr.) S. ruckeri Lindl. (= S. inodora Lodd. ex Lindl.) S. whittenii Muestras de las plantas investigadas estn depositadas en el Herbario del Botanische conservadas en lquido en una solucin de agua / alcohol etlico / formol / cido acetico / glicerina (40/45/5/5/5). Para la co una hoja de aluminio. El aire cargado del perfume por 3 a 4 horas (headspace technique). Despus las hexano de alto grado de pureza e injectadas en un fue hecha con un espectrmetro de masas y con su tiempo de retencin en una columna polar (WAX fase). Todos los anlisis eran realizados por R. Kaiser en el laboratorio de Givaudan en Dben dorf, Suiza (Fig. 1—2).LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.432 LANKESTERIANA TABLA 1. Lista de plantas investigadas.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. GERLACH — Stanhopeinae Mesoamericanae V433 FIGURA S. whittenii FIGURA (generosidad de R. Kaiser).

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TABLA LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.434 LANKESTERIANA Styrene Terpinolene Terpinen-4-ol (E)-Nerolidol S. tigrina 10/16/96 S. tigrina 09/07/90 S. martiana 99/3317 S. martiana 99/3315 S. hernandezii 92/3098 S. saccata 99/3323 S. pseudoradiosa 08/0915 S. pseudoradiosa 08/0915 S. pseudoradiosa 99/3316 S. radiosa 99/3319 S. radiosa 99/3339 S. radiosa 99/3302

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. GERLACH — Stanhopeinae Mesoamericanae V435 Butyl Acetate (Z)-Ocimene epoxide Acetic acid (E)-Ocimene epoxide 1,4-Dimethoxybenzene Methyl 3-phenylpropionate Methyl (Z)-cinnamate Methyl (E)-cinnamate S. graveolens 02/2290 S. ruckeri Soto 5115 S. ruckeri 99/3300 S. whitteni Soto 8068 S. whitteni i 99/3306 S. oculata 73/577 S. dodsoniana 99/3327 S. dodsoniana 02/2291 S. intermedia Soto 3131 S. intermedia 99/3313 S. novogaliciana 00/3843 S. tigrina 99/2068 TABLA

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.436 LANKESTERIANA No todas las sustancias presentadas en la Tabla 2 tienen importancia en la atraccin de los polinizadores, pero en la mayora de los casos no sabemos si son percibibles por estos animales o no. Hay muchas sustancias que son comunitarias como mencionan sabineno, mirceno, limoneno, eucaliptol (1,8-cineol) y los ocimenos). Sabemos que entre ellos hay unos de los mas fuertes atrayentes para los euglosidos, pero como estan presentes en casi todas las especies no pueden servir como rasgos diferenciales. Como consecuencia estas sustancias no estn consideradas en la discusin. SECCI”N SACCATA a) Grupo de S. saccata (S. pseudoradiosa, S. radiosa y S. saccata) Estas tres especies forman un grupo de gran similaridad morfolgica. Tienen el labelo profundamente saccato y una coloracin muy parecida de amarillo hasta crema con una puntuacin del hipoquilo. Jenny (2004) muestra muy bien un gradiente de aperturas en 5 especimenes de S. radiosa y S. pseudoradiosa2. Poniendo estos labelos con este de S. saccata (Fig. 5F) en lnea uno puede observar un gradiente de anchura de las aperturas de S. pseudoradiosa (Fig. 3D) a S. radiosa (Fig. 5D) hacia S. saccata. Las tres Occidental pasando por la Sierra Madre del Sur a S. radiosa S. saccata. Entonces el gradiente Jenny las especies no crecen mixtas, pero todava hay muy pocas observaciones para comprobar ste factor con seguridad. Las 8 muestras de aroma publicadas por Whitten presentadas aqu tienen (E)-ocimeno como 87,3%. El cinamato de metilo junto con sus derivados representa otra caracterstica en algunos clones. En la Tabla 2 se presentan dos analisis de la misma planta de S. pseudoradiosa diferentes; es interesante ver la variabilidad del aroma Williams que en este grupo empieza una especiacin alopatrica. b) Stanhopea intermedia y S. novogaliciana se pueden distinguir. Conservadas en lquido y pueden diferenciar. Unicamente el color es el caracter diagnostico para las dos, S. intermedia es casi unicolor de un amarillo plido (Fig. 4A, 5A), mientras S. novogaliciana tiene el color amarilla fuerte con manchas grandes marrones en las bases de los ptalos y del labelo (Fig. 5C). Rosillo (1984) publica otro argumento para la diferencia de las dos. l dijo que las dos crecian juntas en la misma barranca, pero solamente la S. novogaliciana era visitada por euglosidos de color azul metlico, mientras que la S. intermedia frequente si uno ve varias Stanhopeinae de la misma fuertemente mientras que otros carecen de visitantes. Eso depende del comportamiento de los euglosidos, los aromas y despus de cierto tiempo se van. Por esta razn supongo que era un efecto casual. Observaciones al respecto seran muy agradecidas! diferencia bien. Las sustancias mayoritarias comunes de las dos son: mirceno, limoneno, eucaliptol y S. novogaliciana investigada contiene de alta concentracin. Puede ser que se trata de un artefacto en la recoleccin del aroma, porque el indol no es muy volatil y por esta razn se colecta con ms otro lado tambin se puede aclarar con la presencia de dos quimotipos que tenemos tambien en otras especies, por ejemplo en S. ruckeri, una especie con un concepto morfologico bien claro.2 La foto esta invertida, o hay que cambiar izquierdo por derecho.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. GERLACH — Stanhopeinae Mesoamericanae V437 La declaracin de Rosillo (1984) del euglosido azul comparndole con el polinizador de S. radiosa (por el Euglossa; con alta probabilidad se trata de Euglossa mixta una especie con amplia distribucin. Por la morfologa tan similar de ambos y el crecimiento en la misma regin se hace bien visible que se trata de una sola especie que aparece en dos coloraciones diferentes y dos quimotipos. Por las reglas del cdigo botnico hay que usar el nombre mas antiguo, as S. novogaliciana cae en la sinonima de S. intermedia. c) Resto de la Seccin (S. hernandezii, S. maculosa, S. martiana y S. tigrina) En la taxonoma del resto de la seccin no hay problemas en la delimitacin de las especies. Todas tienen el epiquilo con los margenes erectos, formando un V en corte transversal, asi con el pice tridentado, con la punta central en la mayora de las especies mas cortas que los laterales. Todos tienen manchas, a veces muy grandes y en algunos clones de S. tigrina ocupando su color basico (S. tigrina f. nigroviolacea). Estas especies son endmicas de Mxico, vienen de bosques mesophyticos desde los Edos. de Sonora y Tamaulipas en el norte hasta Oaxaca y Veracruz en el sur. La composicin qumica de la fragrancia de tres especies de este grupo fue analizado, dejando solamente S. maculosa sin investigacin de aroma S. martiana (Fig. 4B) y S. tigrina (Fig. 6A) est dominado por el acetato de feniletilo, una sustancia de aroma muy fuerte y pesado. Las dos se diferencian en su aroma por tener acetato de benzilo y salicilato de metilo en S. tigrina y linalool en S. martiana en porcentajes mayores. S. hernandezii (Fig. 3B) tiene un aroma bien diferente de las dos, reminiscente a canela. Grandes cantidades de acetato de cinamilo y acetato de benzilo son responsable de este olor. Estas tres especies se SECCI”N WARDII a) Grupo de S. oculata (S. dodsoniana, S. oculata y S. whittenii) Este grupo es el ms difcil en su taxonoma entre todas las especies de Stanhopea . Las plantas vienen desde la Sierra Madre Oriental hasta Nicaragua. S. oculata (Fig. 3C) es conocida de los Edos. de San Luis Potosi, Puebla, Veracruz, Oaxaca y Chiapas de Mxico llegando hasta Nicaragua en el sureste, S. dodsoniana (Fig. 5B) de Veracruz, Oaxaca y Chiapas y llega tambin a Nicaragua, mientras que S. whittenii (Fig. 6B) tiene su distribucin desde Chiapas en Mxico hasta Guatemala y Belice. Entonces en el Edo. de Chiapas todas las tres estn presentes, S. dodsoniana y S. oculata tienen la distribucin en los Edos. de Veracruz, Oaxaca unas 30 muestras de aroma investigadas, todava no se ha detectado un carcter discriminador. Es probable que hay hbridos naturales en las muestras investigadas, porque son morfologicamente bien parecidos y hay algunos clones que son intermedias en su forma entre dos especies. Aunque estas tres especies son las mas comunes de las Stanhopeas de Mxico todava casi no por Eufriesea caerulescens en Chiapas de una planta trada de un jardn de Oaxaca. Como en esta poca solamente se conoca S. oculata , no podemos decir con seguridad si trataba verdaderamente de esta especie. Aqu unicamente la observacin de la polinizacin y el estudio de poblaciones puede aclarar la confusin al respecto de su taxonoma. b) S. ruckeri Los dos clones investigados de S. ruckeri presentan un aroma totalmente diferente. Una huele muy agradable a canela (derivados de alcohol cinmico), la otra (Fig. 3E) est dominada por el alcohol y el acetato de feniletilo, as tiene un aroma pesado. Sera sumamente interesante investigar ms clones de esta especie para ver si tiene dos quimotipos o si el aroma c) S. graveolens Esta especie (Fig. 3A) es la ms fcil de distinguir de todas las Stanhopea. Su aroma es algo desagradable causado por un alto porcentaje de indol (10%). Dos clones mas de sta especie de Guatemala fueron investigados y muestran el mismo patrn en

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.438 LANKESTERIANA FIGURE 3. A — Stanhopea graveolens 02/2290. B — S. hernandezii 92/3098. C — S. oculata 73/577. D — S. pseudoradiosa 99/3316. E — S. ruckeri 99/3300. F — S. saccata 99/3323. Photo by G. Gerlach.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. GERLACH — Stanhopeinae Mesoamericanae V439 FIGURE 4. A — Stanhopea intermedia 99/3313. B — S. martiana. Photo by G. Gerlach.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.440 LANKESTERIANA FIGURE 5. A — Stanhopea intermedia 99/3313. B — S. dodsoniana 99/3327. C — S. novogaliciana 00/3843. D — S. radiosa 99/3302. Photo by G. Gerlach.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. GERLACH — Stanhopeinae Mesoamericanae V441 FIGURE 6. A — Stanhopea tigrina 99/2068. B — S. whittenii 99/3306. Photo by G. Gerlach.

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442 LANKESTERIANALANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.similares de indol se encuentran adems en S. hernandezii, S. intermedia y S. martiana. El aroma de S. graveolens casi no se puede distinguir del de S. intermedia. Conclusiones Tomando en cuenta que los polinizadores de las especies de Stanhopea son solamente Euglsidos en la bsqueda de aromas preferidos hay que echar el ojo especialmente a este carcter. Los anlisis de las aromas florales muestran que algunas especies estn caracterizadas por el aroma como S. hernandezii, S. martiana, S. tigrina, S. intermedia y S. graveolens , mientras que otras presentan dos quimotipos o son bastante variable en su composicin. Lamentablemente las investigaciones de aromas florales estn restringidas a un laboratorio y la coleccin del aroma floral en el campo es bastante difcil. La taxonoma est basada en muestras de herbario y descripciones que no incluyen informaciones de la fragancia de la respectiva especie. La dificultad es que son dos disciplinas que no cuadran, en el campo y en el herbario no se puede analizar los aromas, solamente al revs funciona, a tener la planta en flor uno puede hacer el anlisis del aroma floral y cuadrarlo con el polinizador y la taxonoma. Este mtodo consume mucho tiempo, as hay pocos colegas que investigan en esta direccin. Observar plantas en el campo en muchas ocasiones es un caso de suerte, primero la floracin es muy corta (2 a 4 das), segundo la las plantas observadas deben de estar al alcance del observador y no en el dosel; adems, el tiempo debe ser suficiente para que lleguen los polinizadores. Como es impredecible saber si uno puede tener xito en estas observaciones, es ms probable que este trabajo pueda ser llevado a cabo por un aficionado o cientfico que no depende de la carrera. Es imposible proponer a estudiantes un tema como tal para una tesis, por sus resultados impredecibles. De todos modos quiero animar a todos los amantes de la naturaleza para que vayan al campo: necesitamos urgentemente informaciones de este tipo. AGRADECIMIENTOS . Quiero agradecer especialmente a Eric Hgsater (Mxico) por su inmensa ayuda en relacin a las plantas y por brindarme informacin sobre las localidades de Stanhopeinae mexicanas, a Gerardo Salazar, Mark Whitten (en orden alfabtico) por su discusin al respecto de estas plantas, al Dr. Roman Kaiser (Givaudan Research, mi esposa Corina Gerlach por la ayuda con la correcin del laboratorio y la diagramacin, y a Bert Klein y su equipo de jardineros por cultivar nuestras plantas preciosas. LITERATURA CITADA Gerlach, G. 1999. 854. Stanhopea Frost ex Hook. 1829(333). En Schlechter: Die Orchideen III / A, Berlin 2355-2372. Jenny, R. 2003. The Genus Stanhopea 1th Part S. anfracta to S. napoensis. Caesiana 22: 1-145. Jenny, R. 2004. The Genus Stanhopea 2nd Part. Caesiana 22: 146-291. Rosillo de Velasco, S. 1984. Stanhopea novogaliciana Rosillo, una nueva especie del occidente de Mexico. Orquidea (Mx.) 9(2): 251-260. Stanhopea (Orchidaceae) from Mexico. Lindleyana 16: 144-148. Soto Arenas, M.A. 2003. Stanhopea dodsoniana Soto Soto Arenas, M.A. 2003. Stanhopea hernandezii (Kunth) Schltr. Icon. Orchid. 5-6: t. 674. Soto Arenas, M.A. 2003. Stanhopea maculosa Westc. Icon. Orchid. 5-6: t. 675. Soto Arenas, M.A. 2003. Stanhopea martiana Bateman ex Lindl. Icon. Orchid. 5-6: t. 676. Soto Arenas, M.A. 2003. Stanhopea oculata (G. Lodd.) Lindl. Icon. Orchid. 5-6: t. 677. Soto Arenas, M.A. 2003. Stanhopea tigrina Bateman. Icon. Orchid. 5-6: t. 678. Stanhopea whittenii Orchid. 5-6: t. 679. Stanhopea (Orchidaceae). Lindleyana 7(3): 130-153.

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CROSSOGLOSSA SOTOANA (ORCHIDACEAE: MALAXIDEAE), A NEW SPECIES HONORING THE LATE MEXICAN BOTANIST , MIGUEL NGEL SOTO ARENAS FRANCO PUPULIN 1 & ADAM P. KARREMANS 1,21Lankester Botanical Garden, University of Costa Rica2ngel Andreetta Andean Orchid Research Center, University Alfredo Prez Guerrero, Ecuador3Harvard University Herbaria, Cambridge, MA, U.S.A.4Corresponding author: franco.pupulin@ucr.ac.cr443. 2010. LANKESTERIANA 9(3): Robert L. Dressler and Calaway H. Dodson created Crossoglossa in 1993 to accommodate several species previously placed in Microstylis (Nutt.) Eaton section Blephariglottis Schltr., together Liparis Rich. and Malaxis Crossoglossa can be distinguished from the latter genera by the lack of pseudobulbs, the more or less elongate stems with several distichously arranged leaves, the short, straight column clasped by basal lobes or auricles of the lip, the incumbent anther (shared with Liparis ), and the structure of the fruit, with the wide valves being thin and papery and the narrow ones being thick and woody and free of connecting tissue. The genus Crossoglossa blephariglottis (Schltr.) Dressler. In 1999, in their studies on Liparis , Garay and Romero reduced Crossoglossa (as Crassoglossum ) as a synonym under Liparis sect. Tipuloidea (Ridl.) pseudobulbous plants with ascending stems from a decumbent base, basally imbricate by the bases of distichous leaves not articulated with leaf-sheaths; and a short column reminiscent of that of Malaxis close relationship of Crossoglossa with Liparis , with the species of the former genus forming the sister group to a complex of Central American Liparis . Even though at this time there are no published phylogenetic analyses including species referable to Crossoglossa , Noguera-Savelli and her co-workers (2008) suggest that in future generic realignments within the Malaxideae, Crossoglossa could be in Liparis . However, updated phylogenetic results currently being prepared for publication demonstrate that Crossoglossa s closest relatives are the reptant Andean “ Liparis ” such as “ L. ” crispifolia and “ L. ” wageneri . Crossoglossa is only distantly related to the clade that includes the type species of Liparis [ L. loeselii (L.) Rich.] and, according to DNA sequences of ITS and matK, deserves to be maintained as distinct (G.A. Salazar, pers. comm., 2009). As actually circumscribed, Crossoglossa includes some 25 species, mainly distributed in southern Central America and Andean South America, from Colombia to Bolivia, with the highest diversity found in Colombia and Ecuador. No species have been recorded from Venezuela so far (Noguera-Savelli et al ., 2008). The number of taxa rapidly diminishes toward the north, with a single species recorded in Nicaragua (Hamer, 2001). The real diversity of the genus, however, is still unclear, considering that more than half of the species attributed to Crossoglossa ABSTRACT: Crossoglossa sotoana, a new species from Costa Rica, is described and illustrated. It is compared with Ecuadorian C. barfodii, from which it differs by the strongly undulate margins of the leaves, the resupinate the callus composed by two transversely rectangular lamellae. It can be distinguished from C. boylei , also KEY WORDS : Orchidaeae, Malaxideae, Crossoglossa, C. sotoana, Costa Rica, new species

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were published in the last 15 years, after the proposal 1995; Ortz, 1995; Dressler 1997; Vsquez, 1999; Noguera-Savelli et al ., 2008 ). Here we describe a new species from Costa Rica: Crossoglossa sotoana sp. nov. TYPE: COSTA RICA . Alajuela: San Ramn, ngeles, Alberto M. Brenes BiologicalReserve, 10” N 841” W, 800-900 m, shores of Ro San Lorencito, cultivation at Lankester Botanical Garden, University of Costa Rica, 5 Apr. 2005, D. Bogarn 817 (holotype, USJ; isotype, JBL). Similis Crossoglossae barfodii Dodson, a qua recedit resupinatis, linearibus incurvis, labello obovato-sub-pandurato callo duobus transverse rectangularibus lamellis composito. Terrestrial, erect herb , to 15 cm tall excluding the Roots ca. 2 mm in diameter. Stem terete, 3-5 mm in diameter, 4-5 cm long, laxly concealed by the leaf bases. Leaves elliptic-spathulate, obtuse, from a narrow, conduplicate base clasping the stem, the margins strongly undulate, 50-80 15-20 mm above the middle. an erect, successively manydistinct longitudinal keels provided with glandular trichomes, to 15 cm long. Floral bracts triangular, acuminate, the margins with glandular hairs, 6.07.0 2.5 mm. Ovary terete-subconical, glabrous, proximally white, distally green, to 6 mm long including the pedicel. Flowers resupinate, spreading, slightly facing downwards, sepals and petals with green toward the base, the central vein and the basal calli yellow; column green, anther yellow. Dorsal sepal ovate, rounded, 3-veined, concave toward the apex, 2.6 1.6 mm. Lateral sepals narrowly ovate, rounded, 3-veined, 2.5 1.5 mm, sometimes shortly connate at base. Petals ligulate, obtuse, 1-veined, slightly porrect-incurved in natural position, 2.5 1.0 mm. Lip obovate-subpandurate, retuse, 5-veined, basally concave, then convex, 3.0 ciliate, the base subcordate with the auricles partially clasping the column, provided with a basal, ring-like callus around the column, the callus bearing one transversely rectangular, thickened, slightly concave projections on each side of the midvein. Column subterete, clavate, 1 mm long, the distal portion expanded around the subapical stigma. Anther cucullate, transversely elliptic-reniform, 2-celled. Pollinia 2, obovate-subspherical. DISTRIBUTION : Known only from Costa Rica. ECOLOGY : Plants of C. sotoana grow as terrestrials in loose, moist soils and leaf litter in the understory of primary vegetation. The only known population of this species was found in the premontane wet forest of the Tilarn mountain range, on the Caribbean watershed of the continental divide in central Costa Rica, at about 800-900 m elevation. EPONYMY: Named in honor of the recently deceased Mexican botanist, Miguel ngel Soto Arenas, to commemorate his fundamental contributions to orchidology. Crossoglossa sotoana closely resembles the Ecuadorian C. barfodii Dodson, but that species has obcordiform-pandurate (vs. obovate-pandurate) lip, provided at the base with two erect, triangular lamellae (vs. low, transversely rectangular). Crossoglossa boylei Dodson, also from Ecuador, can be distinguished broadly ovate dorsal sepal and ovate petals. In Costa Rica, C. sotoana can be distinguished from C. fratrum (Schltr.) Dressler and C. tipuloides (Lindl.) Ktze. by the short, nearly acaulescent plants and its distinctly lobed, obovate-subpandurate, retuse lip. Crossoglossa aurantilineata Pupulin has longer and narrower leaves, an orange stripe along the midrib of the lip, while C. blephariglottis (Schltr.) Dressler presents an acute lip with longer linear petals. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.444 LANKESTERIANA

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN & KARREMANS — Crossoglossa sotoana445Crossoglossa sotoana perianth. E — Column and lip, oblique view. F — Column, adaxial view. G — Anther and pollinia. Drawn by

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ACKNOWLEDGEMENTS . services of Ministerio del Ambiente, Energa y Telecomunicaciones de Costa Rica (MINAET) and Sistema Nacional de reas de Conservacin (SINAC) for issuing in this study were collected. The present paper is part of by the Vice-Presidency of Research, University of Costa drawing that illustrates the new species. LITERATURE CITED Atwood, J.T. 1989a. Malaxis blephariglottis. Icon. Pl. Trop. 14: sub pl. 1331. Atwood, J.T. 1989b. Malaxis tripuloides. Icon. Pl. Trop. 14: sub pl. 1332. Cribb, P.J. 2005. Crossoglossa. Pp. 460 in: A. M. (eds.). Genera Orchidacearum, Vol. 4. Epidendroideae (part one). Oxford University Press, Oxford. Dodson, C.H. 1995. Confusion concerning the genus Crossoglossa (Malaxideae: Orchidaceae). Lindleyana 10(1): 3. Dressler, R.L. 1997. Novedades en orqudeas mesoamericanas – I. Orquideologia 20(3): 253. Crossoglossa. Pp. 148 in: C. H. Dodson, Native Ecuadorian orchids. Vol. 1: Aa—Dracula. Dodson Trust, Sarasota, USA. Orchidum II. Harvard Pap. Bot. 4(2): 475-488. Hamer, F. 2001. Orchidaceae. In: W.D. Stevens, C. Ulloa Nicaragua. Monogr. Syst. Bot. Missouri Bot. Gard. 85(2). Gonzlez. 2008. Description of a new species and notes on Crossoglossa (Orchidaceae: Epidendroideae: Malaxideae) from the eastern Andes in Colombia and Venezuela. Brittonia 60(3): 240. Ortz, P. 1995. Orqudeas de Colombia. 2nd ed. Corporacin Capitalina de Orquideologa, Santafe de Bogot. Pupulin, F. 2000. New species of Costa Rican Orchidaceae. Lindleyana 15(1): 21. Crossoglossa neirynkiana (Orchidaceae, Malaxidinae), a new species from Ecuador. Ann. Bot. Fennica 38: 91. Vsquez, R. 1999. Crossoglossa: un registro nuevo para C.dodsonii una especie nueva del departamento de Cochabamba. Rev. Soc. Boliv. Bot. 2(2): 153.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.446 LANKESTERIANA

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DOS ESPECIES NUEVAS DE PLEUROTHALLIDINAE (ORCHIDACEAE) DE MXICO RODOLFO SOLANO G”MEZ Centro Interdisciplinario de Investigacin para el Desarrollo Integral Regional, Unidad Oaxaca, Instituto Politcnico Nacional. Hornos 1003, Santa Cruz Xoxocotlan, 71230, Oaxaca, Mxico. solanogo@yahoo.com.mx.447. 2010. LANKESTERIANA 9(3): la subtribu Pleurothallidinae para Mxico ha hecho que ste sea uno de los grupos mejor estudiados de la Arenas, 2003; Garca-Cruz et al. , 2003; LpezVelzquez et al 2007; Solano Gmez et al Arenas, 2008). La diversidad de este grupo en Mxico est representada por 20 gneros, 203 especies y 3 et al ., 2007). Si bien para la mayora de las especies su delimitacin taxonmica ha quedado bien establecida, permanecen varias que no han sido publicadas, por lo que en este trabajo dos de ellas son publicados como nuevas a partir de material vivo y herborizado; cada especie proporciona un mapa para indicar su distribucin conocida en Mxico. Acianthera sotoana R.Solano, sp. nov. TIPO: MXICO . Oaxaca: Municipio Totontepec Villa de 1800 m, colectado septiembre 1977, prensado 23 julio 1983, O. Surez 566 (holotipo, AMO; isotipo, AMO). FIG . 1, 3. extus et intus, petalis non falcatibus, labello acuto ad apicem, et habitu differt. Hierba rizomatosa, hasta de 36.5 cm de altura. Races Rizoma muy corto, engrosado, de 8-10 mm de longitud entre tallos adyacentes, 4.5-6.0 mm de grosor. Tallos erectos, cilndricos, formados por 4-5 entrenudos, 4.017.5 cm de largo, 1.6-3.5 mm de dimetro; cubiertos totalmente por vainas lateralmente comprimidas, a lo largo de las nervaduras, caedizas, densamente manchadas de prpura. Hoja ssiles, una por tallo, apical, ssil, elptica, obtusa, coricea, cortamente bilobada y mucronada en el pice, ligeramente recurvada en los mrgenes, 10-19 x 4-6 cm. RESUMEN: Se describen e ilustran dos especies nuevas de la subtribu Pleurothallidinae como Acianthera sotoana, de Oaxaca y Veracruz, y Stelis sotoarenasii, de Oaxaca. Estas orqudeas haban sido confundidas previamente con especies similares, la primera con Acianthera johnsonii, la segunda con Stelis retusa. Para cada especie se proporciona informacin sobre distribucin, hbitat, fenologa, estado de conservacin y un mapa de sus localidades conocidas. ABSTRACT: Two new species of subtribe Pleurothallidinae are described and illustrated as Acianthera sotoana, from Oaxaca and Veracruz, and Stelis sotoarenasii, from Oaxaca. These orchids have been previously confused with similar species, the former with Acianthera johnsonii, the latter with Stelis retusa. For each species information about distribution, habitat, phenology, conservation status, and a map with their known localities are provided. PALABRAS CLAVE / KEY WORDS: Acianthera sotoana, endemismos, Oaxaca, orqudeas, Stelis sotoarenasii, Veracruz

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racemosa, generalmente de la base del tallo y colgante, a veces tambin del pice del tallo, ms larga que la apical; pednculo cilndrico, 1-3 cm rodeado por una brctea espatcea, conduplicada, triangular, obtusa, escariosa, caediza; brcteas del imbricadas, obtusas, membranceas, densamente cubiertas de manchas prpura, 6.0-7.7 cm de largo. similares a las del pednculo pero ms grandes, 7.9-8.7 mm de largo. Ovario subpiramidal, subtriangular en seccin transversal, engrosado hacia el pice, verde con manchas prpura, glandularpubescente; articulado en un ngulo casi recto a un pedicelo cilndrico. Flores casi perpendiculares al racimo, poco abiertas, 17-21 mm de largo; spalos caf-anaranjados o amarillo-anaranjados, densamente caf-anaranjados con manchas prpura; labelo anaranjado con manchas prpura; columna y antera prpura. Spalos casi paralelos a la columna, carnosos, mucronados en el pice, glandular-pubescentes en en los mrgenes; spalo dorsal lanceolado a oblongolanceolado, subagudo, 7-nervado, carinado sobre la nervadura media, 17.0-18.5 x 5 mm; spalos laterales unidos entre s por dos tercios de su longitud para formar un sinspalo cncavo, carnoso, oblongolanceolado, bipartido, 18.4-21.5 x 10.0-13.4 mm, cada spalo 7-nervado, carinado sobre la nervadura media. Ptalos paralelos a la columna, cartilaginosos, lanceolado-subrmbicos, oblicuos, acuminados, angulados en la mitad, ciliados en los dos tercios superiores, 3-nervados, 11.2-12.3 x 2.0 mm. Labelo columna, en posicin natural paralelo con respecto a la columna, arqueado, carnoso, conduplicado, papiloso x 2.1 mm; cuando extendido cordado-sagitado, subagudo, 6.7 x 3.7 mm; con dos callos engrosados, subparalelos y ligeramente erectos sobre la lamina, ms un callo prominente, rojizo e incurvado hacia la base. Columna delgada, ligeramente arqueada, el cuerpo de 5 mm de largo y 1 mm de ancho a nivel del estigma; con un pie de columna bien desarrollado, incurvado, casi la mitad de largo que el cuerpo de la columna; clinandrio cubriendo totalmente la antera, 3-dentado. Estigma la columna; rostelo ventral, laminar, convexo. Antera ventral, esfrico-ovoide, diminutamente glandulosa ancho. Polinario formado por dos polinios ovoides, lateralmente comprimidos, amarillos, 0.6 mm de largo, unidos por caudculas granulosas. OTROS ESPECMENES : MXICO . Oaxaca: Municipio el banco del ro, 1800 m, septiembre 1977, prensado 20 junio 1981, O. Surez 566 (AMO). Municipio el banco del ro, 1800 m, septiembre 1977, Greenwood s.n. (AMO). Municipio Totontepec Villa de Morelos, septiembre 1977, O. Surez 2500 (OAX). Municipio Totontepec Villa de Morelos, Santa Mara Huitepec, ro Llano, bosque de galera con Platanus rodeado de selva mediana perennifolia, 1497 m, 17.27’’N, 96.35’’O, rupcola, abundante a la orilla del ro, 13 marzo 2009, (OAX). Veracruz: Municipio Pajapan, cima del volcn San con puntos morados, 2 septiembre 2007, T. Krmer (MEXU). Municipio de Tatahuicapan de Jurez, faldas del volcn Santa Martha, bosque 28 agosto 2008, T. Krmer 3696 (MEXU). Municipio regular, con cpsulas, 19 octubre 2008, Castaeda 199 (MEXU). DISTRIBUCI”N : Esta especie es endmica de Mxico la vertiente del Golfo de Mxico (Fig. 3), en el Eje Volcnico Transversal y la Sierra Madre Oriental. HBITAT : : Las poblaciones de esta especie se localizan entre 1125 y 1800 m de altitud, en zonas con selva mediana perennifolia, con presencia de arroyos permanentes y neblinas frecuentes durante todo el LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.448 LANKESTERIANA

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Figura 1. Acianthera sotoana R.Solano, basado en O. Surez 566, (AMO, holo.), dibujo de R. Solano. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOLANO — Dos nuevas Pleurothallidinae para Mxico449

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encima de una capa de musgo o materia orgnica, pero Las localidades de esta especie se conocen en zonas de clima semiclido hmedo, con un rgimen de lluvias en verano, temperatura media anual de 18C y FENOLOGA junio y septiembre, ha sido colectada con frutos en desarrollo en octubre y se han visto frutos secos poco despus de haber liberado las semillas en marzo, poca en la que empieza el crecimiento vegetativo. ESTADO DE CONSERVACI”N . No evaluado. Las poblaciones conocidas de esta especie se presentan dentro o en las inmediaciones de reas naturales protegidas: el Parque Nacional Pico de Orizaba-Cofre de Perote (Veracruz), Reserva de la Biosfera Los Tuxtlas (Veracruz) y Reserva Comunal de Santa Mara Huitepec (Oaxaca), donde an existe hbitat favorable para la orqudea. ETIMOLOGA : Esta especie est dedicada a Miguel ngel Soto Arenas, un maestro, amigo y colega para la mayora de orquideologos mexicanos en activo, por su invaluable aportacin al estudio de las orqudeas mexicanas. Esta especie, junto con A. herrerae (Luer) R.Solano A. johnsonii forman un grupo que se del tallo y de su pice, el cual recientemente fue incluido, al igual que las especies de Echinosepala (Luer, 2004). Aunque los primeros ejemplares de Acianthera sotoana fueron colectados desde 1977, stos fueron confundidos con A. johnsonii, una especie centroamericana que en Mxico se conoce de la regin de los Altos de Chiapas; sin embargo, la segunda es arqueados y el labelo con el pice redondeado y el callo 2003). Por su parte, A. herrerae tiene spalos con el mucrn ms largo, ptalos abruptamente acuminados y atenuados, labelo con una forma muy diferente y provisto de lbulos laterales uncinados (Luer, 1991, 2004). Previamente se haba usado el nombre indito “Acianthera mixe” para la especie aqu descrita y as haba sido registrada por algunos autores (Soto Arenas et al., 2007; Solano Gmez et al., 2008). Stelis sotoarenasii R.Solano, sp. nov. TIPO: MXICO . Oaxaca: Municipio Villa Sola de Vega, La Cumbre, carretera Oaxaca-Puerto Escondido, cerca de las torres de microondas, 2179 m, 16.32’’N, abundante, 26 abril 2009, R. Solano 2550 (holotipo, OAX; isotipos, AMO, MEXU). FIG . 2, 3. similaris sed folio latior ad apicem rotundato, racemis Hierba Races 0.7-1.2 mm grosor. Tallos formados por 2 entrenudos, 2-6 cm largo, 1.0-1.5 mm dimetro, cubiertos totalmente por vainas tubulares, obtusas, carinadas y mucronadas, escariosas, caedizas; con un anillo engrosado cerca del pice. Hoja una por tallo, apical, carnosa, erecta, oblanceolada, redondeada, ligeramente recurvada en el pice, 3.5-9.5 x 1.31.8 cm, la base subpeciolada, atenuada, acanalada.. surgiendo del anillo del tallo, racemosa, abren sucesivamente; pednculo, cilndrico, 1.32.3 cm de largo, rodeado en la base por una brctea espatcea, conduplicada, triangular, obtusa, escariosa; brcteas del pednculo 2, la basal cubierta por la brctea espatcea, tubulares, oblicuas y obtusas en el pice, membranceas. tubularinfundibuliformes, obtusas, cortamente apiculadas, membranceas, 3.5-5.0 mm de largo. Ovario arqueado, ensanchado hacia el pice, subtriangular en seccin transversal, 4.3-4.9 mm largo; articulado a un pedicelo de 3.6-7.0 mm de largo. Flores bilabiadas, ligeramente colgantes, 9.0-9.5 mm de largo (de la base al pice de los spalos), 7.3-9.2 mm de apertura (entre el pice del spalo dorsal y los pices de los laterales), 3.0-3.7 mm de ancho (entre los ptalos); los segmentos amarillos, densamente rayados y manchados de prpura oscuro, columna blanca con manchas prpura, antera amarilla con un mancha apical prpura. Spalos carnosos, 3-nervados; spalo dorsal oblongo-lanceolado, obtuso, LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.450 LANKESTERIANA

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Figura 2. Stelis sotoarenasii R.Solano, basado en O. Suarez 2014, (AMO), dibujo de R. Solano. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOLANO — Dos nuevas Pleurothallidinae para Mxico451

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a lo largo de la nervadura media, 9.0-9.4 x 3.0-3.4 mm; spalos laterales connados por casi dos tercios de su longitud en un sinspalo arqueado-recurvado, conduplicado, cortamente bilobado en el pice, con la porciones libres adherentes, cuando extendido formando una lamina oval, obtusa, 5.5-6.0 x 3.3-3.5 mm; el pice de cada spalo cortamente apiculado. Ptalos erectos, cartilaginosos, falcado-espatulados, subredondeados, abruptamente atenuados en la mitad a lo largo de las nervaduras, 4.0-4.3 x 1.5-2.0 mm. Labelo columna, carnoso, arqueado-recurvado, conduplicado, 3-nervado, 1.8-2.0 x 1.3-1.5 mm en posicin natural; la x 0.6-0.8 mm; la lamina extendida ligeramente 3-lobada, obovadasubcordiforme, redondeada, 2.0-2.2 x 1.7 mm; con dos quillas erectas, prominentes que siguen el contorno de los mrgenes y se extienden en los dos tercios basales de la lmina, mas otros dos callos internos el tercio basal de la lmina, aproximados en el pice y divergentes hacia la base. Columna con un cuerpo delgado, fuertemente arqueado, alado, 3.5-3.6 mm de largo, 1 mm de ancho; provista de un pie prominente, descendente, ligeramente ms corto que el cuerpo; clinandrio cubriendo totalmente la antera, con el margen denticulado; alas prominentes, proyectadas hacia el labelo, con el margen denticulado. Estigma ventral, cncavo, subcuadrado, cubierto por una sustancia viscosa y transparente; rostelo ventral, laminar, convexo. Antera ventral, ovoide, 0.6 x 0.4 mm. Polinario formado por 2 polinios piriformes, lateralmente comprimidos, 0.35 mm de largo, provistos de caudculas granulosas. OTROS ESPECMENES : MXICO . Oaxaca: Municipio Villa Sola de Vega, La Cumbre, carretera Oaxaca-Puerto Escondido, cerca de las torres de microondas, 2179 m, 16.32’’N, 97.2’’O, bosque de pino-encino, G. Cruz s.n. (OAX). Abundante. Municipio San Juan Tepeuxila, camino CuicatlnTlacolula, casi 4 km despus de Tepeuxila, 2100 m, 10 julio 2004, R. Jimnez 2451, G. Salazar y J. Reyes (AMO). Municipio Oaxaca de Jurez, Sierra de San Felipe, 17’N, 96’O, bosque de pino-encino, O. Surez s.n. (OAX). Municipio Villa Sola de Vega, km 90 de la carretera Oaxaca-Puerto Escondido, O. Surez 2014, AMO. Municipio Villa Sola de Vega, km 114.7 de la carretera Oaxaca-Puerto Escondido, 2 noviembre 1974, E. Greenwood s.n. [AMO (diapositiva)]. DISTRIBUCI”N . Esta especie es endmica de Mxico y hasta ahora solo se conoce de las vertientes interiores de las Sierra Madre Oriental y la Sierra Madre del Sur en Oaxaca ( FIG . 3). HBITAT : Esta especie se encuentra entre 1500 y 2200 m de elevacin, en bosques de encino y pinoencino con una estacin de sequa bien marcada. gruesas de encinos, directamente sobre la corteza o sobre una capa de musgo. Prospera en zonas de clima templado subhmedo, donde las corrientes de viento son constantes y eso hace que no sean tan hmedos al interior. FENOLOGA . En campo se han visto ejemplares de esta se extiende hasta octubre. No se han observado ejemplares con frutos desarrollados. ESTATUS DE CONSERVACI”N . No evaluado. Aunque solo una de sus localidades se presenta en un rea protegida, la Reserva de la Biosfera Tehuacn-Cuicatln, en las otras el hbitat es an favorable y sus poblaciones pueden llegar a ser relativamente abundantes, formando en un solo rbol hospedero colonias integradas por hasta algunas decenas de individuos. ETIMOLOGA . El nombre de esta especie honra a Miguel ngel Soto Arenas por su excelente contribucin al estudio de la taxonoma y conservacin de las orqudeas mexicanas. En el pasado esta especie ha sido confundida con Stelis retusa, considerndola como una forma centro y occidente de Mxico, en el Eje Volcnico Transversal, mientras que S. sotoarenasiii presenta una distribucin restringida a Oaxaca. Otra especie muy similar es S. aristocratica, pero en esta las plantas son LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.452 LANKESTERIANA

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dorsal no est recurvado; su distribucin se restringe a la Sierra Madre del Sur en Guerrero. Para la especie aqu propuesta como Stelis sotoarenasii se haba usado el nombre indito “Stelis zapotecana”, con el cual fue citada por algunos autores recientemente (Soto-Arenas et al Solano Gmez et al., 2008). AGRADECIMIENTOS. A Eric Hgsater y dos revisores annimos por sus cometarios para mejorar el manuscrito, a Salvador Acosta Castellanos por traducir las diagnosis e latn y a la Secretara de Investigacin y Posgrado del proyecto “Orqudeas en riesgo del estado de Oaxaca” (clave 20090393). LITERATURA CITADA Epidendreae. Pp. 1-173 in de Rzedowski (eds.), Flora del Bajo y de Regiones Adyacentes, Fasciculo 119. Instituto de Ecologa, A.C.CONACYT-CONABIO, Ptzcuaro, Michoacn. Orchid. (Mexico) 1: Orchids of Mexico 1. Asociacin Mexicana de Orquideologa A.C., Mxico, D.F. (Mexico) 5-6: Orchids of Mexico 2-3. Herbario AMO. Mxico, D.F. (Mexico) 10: Orchids of Mexico 4. Herbario AMO. Mxico, D.F. Farrera. 2007. Primer registro de Phloeophila Acta Bot. Mex. 78: 77-83. Luer, C.A. 1991. New species of Pleurothallis . Lindleyana 6: 100-103. Luer, C.A. 2004. New genera and new combinations in the Pleurothallidinae. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 253-265. Lepanthes Sw. en Mxico. Orqudea (Mex.) 14: 1-231. Cuicatln, Oaxaca, Mxico. Fundacin para la Reserva de la Biosfera Cuicatln A.C., Instituto de Biologa Figura 3. Mapa mostrando la localizacin de las poblaciones conocidas de Acianthera sotoana R.Solano (triangulos) y Stelis sotoarenasii R.Solano (crculos). LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. SOLANO — Dos nuevas Pleurothallidinae para Mxico453

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UNAM, Comisin Federal de Electricidad y Sociedad Mexicana de Cactologa A.C. Mxico, D.F. 172 p. Solano-Gmez, R. 1993. El gnero Stelis Sw. (Orchidaceae: Pleurothallidinae) en Mxico. Orqudea (Mex.) 13: 1-112. Solano-Gmez, R. 1999. Orchidaceae III. Stelis . Pp. 1-26 in Fasciculo 113. Instituto de Ecologa A.C. y Universidad de California, Mxico. Solano-Gmez, R. 2000. Additions to Stelis Sw. (Pleurothallidinae) from Mexico. Lindleyana 15: 46-52. species of Stelis (Orchidaceae, Pleurothallidinae) from Guerrero, Mexico. Rev. Mex. Biodiv. 78: 253-256. Garca. 2008. Diversidad, distribucin y estrategias para la conservacin de las Pleurothallidinae (Orchidaceae) en Oaxaca. Bol. Soc. Bot. Mxico 82: 41-52. Acianthera johnsonii in: (Mexico) 5-6: Orchids of Mexico 2-3. Herbario AMO. Mxico, D.F. Stelis retusa in (Mexico) 10: Orchids of Mexico 4. Herbario AMO. Mxico, D.F. Soto-Arenas, M. 1987. Una revisin de las especies mexicanas de Trichosalpinx subgnero Trichosalpinx . Orqudea (Mex.) 10: 247-277. Solano-Gmez. 2007. Orqudeas de Mxico. Herbario AMO-Instituto Chinon. A.C. y CIIDIR Unidad Oaxaca, CONABIO proyecto No. P107, Mxico, D.F. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.454 LANKESTERIANA

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MASDEVALLIA SOTOANA (ORCHIDACEAE: PLEUROTHALLIDINAE), A NEW SPECIES FROM ECUADOR HUGO MEDINA 1,4 & FRANCO PUPULIN 1,2,31 Centro de Investigacion en Orquideas de los Andes “ngel Andreetta”, Universidad Alfredo Prez Guerrero, Ecuador2 Jardin Botanico Lankester, Universidad de Costa Rica, P.O.Box 302-7050 Cartago, Costa Rica3 Harvard University Herbaria, Cambridge, MA, U.S.A. 4 Corresponding author: hugoamt5@gmail.com455. 2010. LANKESTERIANA 9(3): The genus Masdevallia includes today some 400 recognized species (Pridgeon, 2005) and more than 1100 published names (Tropicos, 2009). Although the genus has a long taxonomic history, with many species already known and cultivated during the 19th Century, the diversity of Masdevallia was substantially revealed in the last decades of the last Century, with a record of new descriptions between 1978 and 1988, when 265 Masdevallia names were proposed (references in Luer 1983, 1996— 2002, 1986a, 1986b, 2000a, 2000b, 2000c, 2001a, 2002, 2003). In the course of the actual decade, new species of Masdevallia appeared at a constant rate Sijm 2003; Luer 2001b, 2001c, 2001d, 2002, 2004, taxa, however, have been published in the last few 2007, Valenzuela 2008), with an average of only six new species per year since 2005 (compared with more than 14 species/year between 2000 and 2005). This obtaining plant material from tropical countries, due to the enforcement of controls on wild plants trade, which particularly affect the interchange of samples the big effort made by Luer and his co-workers in the last 30 years to illustrate the diversity of Masdevallia is During the process of reviewing the taxonomic identity of the vast Masdevallia collection at Ecuagenera, to be documented and incorporated as Andean Orchid Research Center (CIOA, for its acronym in Spanish), University Alfredo Prez Guerrero, and in the related databases, we found a number of taxa species. The new species proposed here has apparently no close relatives, with the exception of M. strobelii Masdevallia sotoanasp. nov. the collection of Ecuagenera at Gualaceo, accesin No. 002293, 14 Oct. 2009, H. Medina 72 (holotype, QCNE). Fig. 1. Masdevalliae strobelii Sweet et Garay similis, foliis ABSTRACT: A new species of Masdevallia, M. sotoana, is described and illustrated from Ecuador. The new species is similar to M. strobelii , from which is mainly distinguished by the smaller size of the plant and the much shorter column foot . RESUMEN : Se describe e ilustra una nueva especie de Masdevallia de Ecuador, M. sotoana. La nueva especie es similar a M. strobelii el pice, as como el pie de la columna mucho mas corto. KEY WORDS : Orchidaeae, Malaxideae, Crossoglossa, C. sotoana, Costa Rica, new species

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.456 LANKESTERIANAMasdevallia sotoana ovary, column and lip, lateral view. E — Left petal. F — Lip, spread. G — Column, ventral and three quarters views. H — Pollinarium and anther cap. Drawn by F. Pupulin from the holotype.

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trichomatibus parvibus ornatis, petalis lineari-oblongis dente basali perparvo instructis, labello anguste columnae multo breviore recedit. Epiphytic, caespitose, erect herb, to 7 cm tall. Roots Stems (ramicauls) shorter than the leaf, erect, ca. 5 mm long, concealed by 2 tubular, loose, scarious sheaths, 3 mm long. Leaves petiolate, the blade elliptic, acute, minutely tridenticulate, green, the base of the petiole darker, 2.8.5 x 1.0.2 cm; petiole conduplicate, to 2.4 cm long. singlefrom the apex of the stem, to 3.5 cm long. Floral bract tubular-lanceolate, amplectent, glumaceous when young, becoming dry-papyraceous with age, 6 mm long. Pedicel terete, to 6 mm long. Ovary articulated with the pedicel, terete, glabrous, 4 mm long. Flowers forming a sepaline cup and apically spreading into base, the apical tails yellow, the petals, the lip and the column white. Sepals united to form a tube, ventrally slightly gibbous, minutely papillose-hirsute toward the apex of the blade, 3-nerved, abaxially carinate along the midnerve, 1.4 cm long, each 6 mm wide, basally connate for ca 1.1 cm, the free portion semi-rounded to trasnversely elliptic, extending at apex into a narrow, Petals linear-oblong, slightly sygmoid, minutely tri-dentate, 1-nerved, the lower marging produced at the base into a short, triangular teeth, thickened along the nerve, 5.0 x 1.7 mm. Lip narrowly lanceolate, acuminate, subtruncate-bilobed at base, the margins of the basal lobes suberect, apically Column straight, 5 mm long, with low marginal wings increasing toward the apex to form a hood over the clinandrum; anther ventral, stigma elliptic. Pollinia 2, narrowly oblong, on a rounded viscidium. PARATYPE: Ecuador. Same locality as the holotype, collection of Ecuagenera at Gualaceo, accession No. 001094, 19 Aug. 2008, H. Medina 69 (QCA). EPONYMY : Dedicated to Miguel Angel Soto Arenas, for his outstanding contributions to the study and knowledge of Neotropical orchids. DISTRIBUTION AND ECOLOGY : Know only from southeastern Ecuador. A single population of this species has been found in the submontane wet forests about 1800 meters of elevation. According to Luer’s (2002) scheme of subgeneric M. sotoana belongs to subgen. Masdevallia sect. Masdevallia subsect. Saltatrices (Rchb.f.) Luer. The group was not affected by generic recircumscriptions and nomenclatural changes recently proposed by Luer (2006) in his reconsideration of Masdevallia. Masdevallia subsect. Saltatrices, which is distributed in the Andes from Venezuela into Peru, is mostly characterized by the deeply connate sepals into a sepaline tube that is often, but not always ventricose (Luer 1996, 2002). The subsection includes 26 species, some of them not closely related. As Luer (2002) himself recognizes, the distinction between subsections Saltatrices and Masdevallia is not sharp. Among the taxa of subsect. Saltatrices, M. sotoana has apparently only a single close relative, M. strobelii. H. Dodson and independently collected in the same region, M. sotoana and M. strobelii are sympatric. Masdevallia sotoana may be distinguished from M. strobelii by the smaller size of the plant (leaves 2.8— 3.5 vs. 4-6 cm in M. strobelii <3 cm vs. >5 cm), the petals provided with a small basal teeth (0.5 mm vs. 2 mm), the narrowly lanceolate and the much shorter column foot (2 mm vs. 5 mm). ACKNOWLEDGEMENTS . We would like to express our most sincere tanks to Jos “Pepe” Portilla and his family for allowing us free access to the plant collection of Ecuagenera and giving any kind of facilities. To the Ministry of Environment of Ecuador for extending the management permits of the living collections where the species treated here have been documented. To the Foundation for Orchid Research and Conservation, Vancouver Orchid Society, Canada, for its kind sponsoring of the AORC. The present paper is part of the Project 814-A7-015, “Inventario y — Orchidaceae”, sponsored by the Vice-Presidency of Research, University of Costa Rica. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. MEDINA & PUPULIN — A new Masdevallia from Ecuador457

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LITERATURE CITED Campacci. M. A. 2005. Uma nova especie de Orchidaceae brasileira. Boletim CAOB 59: 92. Kniger, W. 2000. Neue Arten der Gattungen Masdevallia , Oncidium, Sigmatostalix and Trigonochilum. Arcula 10: 266. Kniger, W. 2001. Neue Arten der Gattungen Masdevallia , Oncidium, Scelochilus and Sigmatostalix. Edition:1 Arcula, Botanische Abhandlungen.11: 282. Kniger, W. 2003. Neue Arten der Gattungen Cyrtochilum , Masdevallia und Incidium. Arcula 14: 349. Masdevallia , Oncidium und Sigmatostalix . Arcula, 12: 298. Luer, C. A. 1983. Thesaurum Masdevalliarum: a monograph of the genus Masdevallia . Verlag Helga Kniger, Munich, Germany. Luer, C. A. 1996. Ein Schatz von Masdevallia : eine Monographie der Gattung Masdevallia . A treasure of Masdevallia : A Monograph of the genus Masdevallia . Missouri Botanical Garden, St Louis, Missouri. Luer, C. A. 1986a. Icones Pleurothallidinarum. I. Systematics of the Pleurothallidinae (Orchidaceceae). Monogr. Syst. Bot. Missouri Bot. Gard. 15. Luer, C. A. 1986b. Icones Pleurothallidinarum. II. Systematics of Masdevallia (Orchidaceceae). Monogr. Syst. Bot. Missouri Bot. Gard. 16. Luer, C. A. 2000a. Icones Pleurothallidinarum. XX. Sistematics of the Jostia, Andinia, Barbosella, Barbrodia, Pleurothallis subgen. Antilla, subgen. Effusia, subgen. Restrepioidia. Addenda to Lepanthes, Masdevallia, Platystele, Pleurothallis, Restrepiopsis, Scaphosepalum, and Teagueia. Monogr. Syst. Bot. Missouri Bot. Gard. 79. Luer, C. A. 2000b. Icones Pleurothallidinarum. IX. Systematics of Masdevallia, part one. M. subgen. Polyantha , section Alaticaules. section Polyanthae. Monogr. Syst. Bot. Missouri Bot. Gard. 77. Luer, C. A. 2000c. Icones Pleurothallidinarum. XXI. Systematics of Masdevallia, part two. M. subgen. Masdevallia , section Coriaceae. Section Dentatae. Section Durae, Section Reichenbachianae, M. subgen. Pygmaeia . Monogr. Syst. Bot. Missouri Bot. Gard. 82. Luer, C. A. 2001a. Icones Pleurothallidinarum. XXII. Systematics of Masdevallia, part three. M. subgen. Masdevallia , section Minutae. Monogr. Syst. Bot. Missouri Bot. Gard. 86. Luer. C. A. 2001b. Miscellaneous new species in the Pleurothallidinae. Selbyana. 22(2): 103. Luer. C. A. 2001c. Miscellaneous new species in the Pleurothallidinae. Revista de la Sociedad Boliviana de Botanica. 30: 37. Luer. C. A. 2001d. Icones Pleurothallidinarum, XXII. Systematics of Masdevallia , part three. Monogr. Syst. Bot. Missouri Bot. Gard. 86: 519. Luer, C. A. 2002. Icones Pleurothallidinarum. XXIII. Systematics of Masdevallia, part four. M. subgen. Masdevallia , section Masdevallia. subsection Caudatae. subsection Oscilantes, subsection Saltatrices. Monogr. Syst. Bot. Missouri Bot. Gard. 87: 781. Luer, C. A. 2003. Icones Pleurothallidinarum. XXV. Systematics of Masdevallia, part . Monogr. Syst. Bot. Missouri Bot. Gard. 91. Luer, C. A. 2004. Miscellaneous new species in Brachionidium , Dracula, Lepanthes, Octomeria, Platystele , Pleurothallis , Porroglossum, and Masdevallia. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 233. Luer, C. A. 2005. Icones Pleurothallidinarum XXVII. Additions to the genus Masdevallia . Monogr. Syst. Bot. Missouri Bot. Gard. 103: 282. Luer, C. A. 2006. Icones Pleurothallidinarum. XXVIII. A reconsideration of Masdevallia. Systematics of Specklinia and vegetatively similar taxa. Monogr. Syst. Bot. Missouri Bot. Gard. 105. Masdevallia species from Peru. Selbyana 27(1): 15. in the Pleurothallidinae. Rev. Soc. Boliv. Bot. 3(1-2): 37. Ortiz, V. P. 2004. A new Masdevallia from Colombia. Orquideologia 23(2): 73-78. Ortiz, P. 2005. Nuevas especies de orqudeas de Colombia. New orchid species from Colombia. Orquideologa 24(1): 1-20. Pridgeon, A. M. 2005. Masdevallia . Pp. 367—370 in : A. M. (eds.), Genera Orchidacearum. Volume 4. Epidendroideae (part one). Oxford University Press, Oxford. Masdevallia from Ecuador. Amer. Orch. Soc. Bull. 35: 377. Tropicos, 2009. Tropicos.org. Missouri Botanical Garden, consulted on15 Oct. 2009. http://www.tropicos.org Valenzuela, G. A. L. 2008. A new species of Masdevallia : (Pleurothallidinae) from Peru. Lankesteriana. 8: 17— 19. Verlag KG, Stuttgart.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.458 LANKESTERIANA

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NEW SPECIES OF PORROGLOSSUM (ORCHIDACEAE: PLEUROTHALLIDINAE) FROM ECUADOR GILBERTO MERINO 1 , ALFONSO DOUCETTE 1,2 & FRANCO PUPULIN 1,3-61Centro de Investigacin en Orqudeas de los Andes “ngel Andreetta”, Universidad Alfredo Prez Guerrero, Ecuador 2Plant Sciences Department, Cornell University, Ithaca, New York, U.S.A.3Jardn Botnico Lankester, University of Costa Rica. 4Harvard University Herbaria , Cambridge, Massachusetts, U.S.A 5Marie Selby Botanical Gardens, Sarasota, Florida, U.S.A6Corresponding author: fpupulin@cariari.ucr.ac.cr459. 2010. LANKESTERIANA 9(3):Introduction. The genus Porroglossum Schltr. (Orchidaceae) includes 38 species restricted to Andean South America, where they range from Venezuela to Peru (Luer 1987). Ecuador is the genus’s center of diversity with 23 species recorded (Dodson 2004). With 8 species, Colombia has the second highest Porroglossum echidna (Rchb.f.) Garay, was discovered in Colombia and was attributed to the genus Masdevallia habit, short ramicauls, and sepals with long tails (Luer 1987). Rudolf Schlechter proposed the genus in 1920, based on P. colombianum [= P. mordax (Rchb.f.) Sweet] and P. muscosum (Rchb.f.) Schltr., mainly distinguishing Porroglossum from Masdevallia by the apical position of the anther and the presence of a long column foot with a free apex (Schlechter 1920). The unusually long column foot in Porroglossum functions to accommodate the active mobility of the labellum. The region of the labellum that rests against the column foot is sensitive to tactile stimuli and responds to touch by relaxing the labellum’s long strap-like (hypothetically) push the pollinator against the column (Bean 1887, Leavitt 1906, Luer 1987). Although the pollination system has been described in detail by Bean (1887), Oliver (1888) and Luer (1987), the not yet been recorded in nature. Sensitive labella have evolved independently in other taxa of the subtribe Pleurothallidinae Lindl. (Luer 1987, 2000). Examples include Acostaea Schltr. and Condylago Luer species, now treated as members of the genus Specklinia Lindl., and Masdevallia teaguei Luer, which was once recognized as the monotypic genus Jostia Luer. The presence of an irritable lip was once thought to be a key character for generic circumscription. Recent phylogenetic analyses based on combined molecular evidence (Pridgeon et al . 2001) show that in some cases it is unsuitable as a character for generic ABSTRACT . Three new species of Porroglossum from Ecuador are described and illustrated. Porroglossum miguelangelii is compared to the Peruvian P. lycinum, from which it differs by the longer, narrower petals with margins that are not obtusely angled below the middle and the presence of stigmatic tooth-like processes. Porroglossum merinoi is distinctive in that it is the only species with densely pubescent sepals. Porroglossum porphyreum is similar to P. schramii, but differs in the sepals with a thickened tail abruptly inserted at their apex, the petals with teeth on both the upper and lower margin, and the labellum provided with a longer and broader basal callus. KEW WORDS : Orchidaceae, Pleurothallidinae, Porroglossum, P. miguelangelii, P. merinoi, P. porphyreum, Ecuador, new species

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.460 LANKESTERIANAcircumscription within the Pleurothallidinae because it creates paraphyletic groups. In contrast, lip motility observed in the well supported monophyletic clade of Porroglossum can be considered a synapomorphy of the genus. Porroglossum is the only currently accepted and recognized genus in the Pleurothallidinae that can be characterized as a whole by the presence of actively motile labella. The systematics of Porroglossum was treated by Sweet (1970, 1972) and by Luer (1987). Luer recognizes two subgenera, subgen. Eduardia Luer and subgen. Porroglossum. Subgenus Eduardia is characterized by a repent growth habit and a Porroglossum is composed of three sections: section Echidnae (Rchb.f.) Luer which contains species characterized by peduncle indumentums, section Tortae Luer which contains species characterized by Porroglossum which contains the rest of the species (Luer 1987). The species described here all belong to the section Porroglossum. Luer (1987) attributed twenty-seven species names to the genus Porroglossum, considering P. colombianum and P. xipheres to be synonyms of P. mordax and P. muscosum, respectively. Since then, Luer has authored and co-authored thirteen new species, bringing the number of species names attributed to Porroglossum to forty and the number of species to thirty-eight (Luer 1988, 1989, 1991, 1994, 1995, 1998, 2004, 2006a, 2006b). In Ecuador, 23 species of Porroglossum have been previously recorded, representing over two thirds of the known species. In preparing a systematic revision of the Ecuadorian species of Porroglossum, we found three species that present enough morphological differences proposed as new to science. Porroglossum miguelangelii Pupulin, sp. nov. locality, 1800 m, collected by Hugo Medina, November 1993, cultivated by Ecuagenera Orchid 2008, G. Merino 1 (holotype: QCNE; isotype: Andean Orchids Research Center (CIOA, by the initials in Spanish), University Alfredo Prez Guerrero, Spirit Collection No. 000845). FIG . 1, 4A. A Porroglosso procul Luer foliis ellipticis obtusis, petalis lineari-ligulatis subfalcatis, in parte media leviter angustatis, lamina labelli late obtriagulariobcuneata marginibus lateralibus incurvatis recedit. Herb epiphytic, caespitose, 4 to 8 cm tall. Roots . Ramicauls terete, erect, abbreviated, blackish at the base 1 mm long, enclosed by 2 to 3 tubular sheaths. Leaf coriaceous, spathulate, elliptical-obovate 2-4 x 0.5-1.0 cm, narrowed at the base into a conduplicate petiole 1.5-2.0 cm long. raceme 8-10 cm long; the peduncle glabrous, 5.0-9.5 cm long. Floral bracts tubular, 4 mm long. Pedicel 5 mm long. Ovary terete, minutely verrucose, 6-sulcate, 5 mm long. Flowers resupinate, held erect, without a detectable fragrance, the dorsal sepal yellow-green suffused with brown, with three brown veins, the lateral sepals yellow-green, the petals translucent, amber with a brown mid-vein, the lip white, spotted with dark brown, the column greenish-white with a cream anther cap. Dorsal sepal broadly ovate, concave, tricarinate, 9.5-10.0 x 5.0 mm, contracted into a thick, erect tail, connate to lateral sepals for about 3 mm. Lateral sepals subrectangular, tricarinate, 10-11 x 6 mm when spread, connate for about 4 mm to form a laterally compressed cup, contracted at apex into a thickened, horizontally outstretched tails. Petals linear-ligulate, the apex rounded and broader than middle portion, 6 x 1 mm. Lip unlobed, obcuneate-obtriangular, 4 x 5 mm at its widest portion, hinged to the column foot by a short rectangular strap, motile, with a sulcus running from the apex to the center of the blade, with a basal low, broad callus. Column short, stout, semiterete, near the stigma. Anther cap transversely ovate, obtuseapiculate. Pollinia droplet shapped, 2 in number, each with a glandular caudicle. EPONYMY : Named to the memory of Miguel ngel Soto Arenas, in recognition of his outsantding contributions to the knowledge of orchids. DISTRIBUTION : Known only from southeastern Ecuador. HABITAT AND ECOLOGY : Epiphytic in shade in premontane and lower montane cloud forests, usually

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FIGURE 1. Porroglossum miguelangelii D — Column and lip, lateral view. E — Lip, spread. F — Anther cap and pollinarium. Drawn by A. Doucette from the holotype.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. MERINO et al. — New species of Porroglossum461

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in windy areas, at elevations of 1800 meters, where the plants mostly establish on small trees covered by mosses and lichens. Porroglossum miguelangelii is most similar to P. lycinum Luer from northern Peru, from which it differs by smaller plants, the longer, narrower petals with margins that are not obtusely angled below the middle, the presence of tooth-like processes before the stigma (absent in P. lycinum), the spreading, pale green sepals, suffused with yellow at their apices (vs. projected forwards, speckled and suffused with brown in P. lycinum). The two species have similarly shaped with dark purple towards their apex. Porroglossum merinoi sp. nov. TYPE: ECUADOR . Morona-Santiago: Tumbez, 1200 m, in cultivation, 12 June 2007, G. Merino 2 (holotype: QCNE; isotype: CIOA, Spirit Collection No. 000155). FIG . 2, 4B. Species Porroglosso lycino Luer brevioribus ligulato-rotundatis, petalis ligulatis dente triangulari margine inferno instructis, labello obtrullato obtuso glabro differt. Herb epiphytic, caespitose, to 6 cm tall. Roots Ramicauls erect, slender, 10--12 mm long, enclosed by 2 to 3 tubular, membranaeous sheaths that become papery with age. Leaf erect, coriaceous, long-petiolate, the blade elliptic-oblanceolate, sub-verrucose, 2.5-3.0 x 0.7-1.0 cm, narrowed at the base into a conduplicate petiole 2.6-3.0 cm long. a suberect, congested, peduncle terete, slender, glabrous, provided with 4-5 widely spaced, tubular bracts to 7 mm long. Floral bract tubular, 5 mm long. Pedicel terete, 7-8 mm long. Ovary terete, subverrucose, 5 mm long. Flowers resupinate, partially spreading, lightly fragrant of with solid yellow at the glabrous apexes, the nerves at the base; the petals translucent yellowish white, with a central stripe and the apex purple-red; the lip with a yellowish white anther cap. Dorsal sepal lanceolate, narrowly obtuse, tricarinate, concave at the base, parallel to the column below the middle and x 3 mm, connate to the lateral sepals for about 1.5 mm to form a slightly, laterally-compressed cup. Lateral sepals transversely obtrapezoidal, 3-veined, narrowed at the middle into thick, ligulate, narrowly obtuse-rounded, forwardly directed tails, 7.5 mm long x 8 mm at its widest portion when spread, basally connate for about 1.5 mm to the dorsal sepal, connate to each other and to the column foot about 5 mm to form a deep, subacute, rounded chin. Petals subfalcate, 1-nerved, the lower margin with a triangular tooth below the middle, rounded at apex 4.0 x 1.3 mm, Lip long-unguiculate, the blade obcuneate-spathulate, broadly obtuse, with a short, sulcate, rounded apiculum, 5.5 mm long x 3 mm at its widest portion, thickened at the base into the straplike claw, bent with tension toward the free apex of the column-foot. Column semiterete, truncate, 2.5 mm long, with narrowly triangular-subfalcate, descending substigmatic arms, produced at the base into a slender, curved foot about 6 mm long; the anther terminal, incumbent. Anther cap transversely ovate, obtuseapiculate. Pollinia droplet shapped, 2 in number, each with a glandular caudicle. EPONYMY : Named after Gilberto Merino, Ecuadorean orchid researcher with a strong interest in DISTRIBUTION : Collected once, known only from southeastern Ecuador. HABITAT AND ECOLOGY : The type specimen of P. merinoi was found growing as a shade epiphyte on a small tree covered by mosses, in cloud forest at about 1800 m of Porroglossum merinoi is one of the most distinctive species in the genus. It is the only species with truly pubescent sepals. P. hirtzii bears minute hairs on its sepals but they are sparsely distributed. P. merinoi is allied to P. andreettae, from which it differs by the petals with entire margins (vs. contracted into acute apices near the middle), a slightly broader labellum, and puberulent sepals provided with a thickened, glabrous, yellow apex. Both species are lightly fragrant. LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.462 LANKESTERIANA

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FIGURE 2. Porroglossum merinoi and lip, lateral view. E — Lip, spread. F — Pollinarium and anther cap. Drawn by A. Doucette from the holotype.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. MERINO et al. — New species of Porroglossum463

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FIGURE 3. Porroglossum porphyreum D — Column and lip, lateral view. E — Lip, spread. F — Pollinarium and anther cap. Drawn by A. Doucette from the holotype.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.464 LANKESTERIANA

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FIGURE 4. Porroglossum species. A — Porroglossum miguelangelii.. B — P. merinoi. C — P. porphyreum. Scale bars = 5 mm. Photographs by F. Pupulin.Porroglossum porphyreum Pupulin, sp. nov. TYPE: ECUADOR . Morona-Santiago: El Aguacate, 1600 m, collected by Jinsop Medina, 1992, cultivated by in cultivation, 20 July 2008, G. Merino 3 (holotype: QCNE; isotype: CIOA, Spirit Collection No. 000743). FIG . 3, 4C. A Porroglosso schramii Luer caudis sepalorum brevioribus incrassatis, sepalis lateralibus transverse ellipticis cauda apicali abrupte instructis, petalis dente acuto margo inferno munitis, labello toto dense pubescenti callo basali magno latiore recedit. Herb epiphytic, caespitose. 6 to 10 cm tall. Roots Ramicauls terete, erect, abbreviated, blackish at the base, 3--5 mm long, enclosed by 2 to 3 tubular, membraenous sheaths that become papery with age. Leaf coriaceous, ellipticobovate, obtuse to acute, 3-5 x 1.5-2.0 cm, narrowed at the base into a conduplicate petiole 1--3 cm long. raceme; the peduncle terete, slender, glabrous, 10-12 cm long. Floral bract tubular, 4mm long. Pedicel 5 mm long. Ovary terete, minutely verrucose, 6-sulcate, 6 mm long. Flowers resupinate, held erect to horizontally, without apparent fragrance, the sepals yellowish white heavily suffused with purple, the petals amber with a brown mid-vein, the lip white suffused with purple, the column and the anther cap white. Dorsal sepal ovate, acute, concave, tricarinate, 6 x 3 mm, contracted at apex into an erect to forward projecting, thickened, linear tail, connate to lateral sepals for about 4 mm, the tails 1 mm in diameter. Lateral sepals ovate, slightly compressed, tricarinate, 10 x 5 mm wide when spread, connate at the base for about 7 mm, abruptly contracted at apex into thickened tails that may project forwards or Petals subfalcate, truncate, 4 x 2 mm, the margins present two triangular teeth below the middle. Lip entire, obcuneate, 4 x 5 mm at its widest portion, hinged by a short rectangular strap to the column foot, motile, microscopically pubescent, with a sulcus running from the apex to the center of the lip, the base with a low, broad callus. Column tooth-like processes close to the stigma. Anther cap transversely ovate, obtuse-apiculate. Pollinia droplet shapped, 2 in number, each with a glandular caudicle. PARATYPES in cultivation at Ecuagenera Orchid Nursery in Gualaceo, 10 December 2008, accession No. CIOA001373 (Centro de Investigacin en Orqudeas de los Andes-Spirit); accession No. CIOA-001374 (Centro de Investigacin en Orqudeas de los Andes-Spirit). ETYMOLOGY : From the Greek porphyreos, purple, in DISTRIBUTION : Known only from southeastern Ecuador.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. MERINO et al. — New species of Porroglossum465

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HABITAT AND ECOLOGY : Plants of P. porphyreum grow as shade epiphytes on the moss covered branches of shrubs, in the premontane forests of southern Andean Ecuador. In its natural habitat, P. porphyreum has Porroglossum porphyreum is similar to P. schramii Luer. P. porphyreum can be distinguished fromP. Schramii by having sepals that abruptly (rather than gradually) contract into thickened (rather than thin) sepaline tails. The petals of P. porphyreum have teeth below the middle on both margins while the petals of P. schramii only bear a tooth margin on the upper margin. The labellum of P. porhpyreum has a basal callus that is longer and broader than the basal callus of P. schramii. ACKNOWLEDGEMENTS . We acknowledges Jos Portilla Andrade and his family for granting access and use of the vast orchid collections at Ecuagenera, as well as for the many facilities they made available at Gualaceo for orchid research. This paper is part of the project No. 814regin Mesoamericana”, supported by the Vice-Rectory of Research, University of Costa Rica. LITERATURE CITED Bean, W. 1887. A sensitive Masdevallia . Gard. Chron. 1: 836. Dodson, C.H. 2004. Native Ecuadorian Orchids. 4:822— 831. Leavitt, R. G. 1906. The blooming of an unusual orchid. The Plant World. 9: 75-80. Luer, C. A. 1980. Miscellaneous New Species in the Pleurothallidinae (Orchidaceae). Phytologia. 46(6): 374. Luer, C. A. 1985. Porroglossum hoeijeri Luer. Orchideen 6(1): 7. Luer, C. A. 1987. Systematics of the genus Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 24: 25 . Luer, C. A. 1988. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 26: 108. Luer, C. A. 1989. Addendum to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 31: 124. Luer, C.A. 1991. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 39: 152. Luer, C. A. 1994. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 52: 134. Luer, C. A. 1995. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 57: 144. Luer, C. A. 1998. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 72: 11415. Luer, C. A. 2000. Jostia: a new genus in the Pleurothallidinae. Monogr. Syst. Bot. Missouri Bot. Gard. 79: 1-4. Luer, C. A. 2006a. Addendum to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 239. Luer, C. A. 2006b. Addenda to Porroglossum. Monogr. Syst. Bot. Missouri Bot. Gard. 105: 253. Schlechter, R. 1920. Beschreibungen neuer Arten: Porroglossum colombianum. Repert. Spec. Nov. Regni Veg. Beih. 7: 83. Sweet, H. R. 1970. Orqudeas Andinas Poco Conocidas. Orquideologia 5: 166. Sweet, H. R. 1972. The Genus Porroglossum. Amer. Orchid Soc. Bull. 41: 519. Oliver, F. W., 1888. On the sensitive labellum of Masdevallia muscosa Rchb. f. Ann. Bot. (London) 1: 237-253. Pridgeon, Alec M., Rodolfo Solano and Mark W. Chase. 2001. Phylogenetic relationships in Pleurothallidinae (Orchidaceae): combined evidence from nuclear and plastid DNA sequences. American Journal of Botany. 88:2286-2308.LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.466 LANKESTERIANA

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ON THE IDENTITY OF MYOXANTHUS SCANDENS (ORCHIDACEAE: PLEUROTHALLIDINAE), WITH A NEW SPECIES FROM COSTA RICA FRANCO PUPULIN 1 , DIEGO BOGARN 1,2 & MELANIA FERNNDEZ 11Lankester Botanical Garden, University of Costa Rica2ngel Andreetta Research Center on Andean Orchids, University Alfredo Prez Guerrero, Ecuador3Harvard University Herbaria, Cambridge, MA, U.S.A.4Corresponding author: franco.pupulin@ucr.ac.cr467. 2010. LANKESTERIANA 9(3): Members of the subtribe Pleurothallidinae (Orchidaceae) are often taxonomically problematic. Both in herbaria and living collections, names of Pleurothallidinae are frequently applied without direct knowledge of the nomenclatural types. Moreover, the and their usually reduced dimensions make more taxa. The misunderstanding of the critical characters of natural variation, often lead to incorrect species circumscriptions. Under these circumstances, it is possible that undescribed species remain hidden under Myoxanthus scandens problem [Atwood, 1987, 1989 (both with illustration); Pupulin, 2002; Luer, 1976, 1982, 1992, 2003]. This species is member of a small group of Myoxanthus stems, and mostly narrowly elliptic leaves, including M. chlo M. epibator Luer M. frutex (Schltr.) Luer, M. parvilabius (C. Schweinf.) Luer, and Pleurothallis pennellia Luer. With the exception of M. scandens, which ranges northward to Costa Rica, this group is eminently South American in distribution, with the highest diversity in the Andes from Colombia to Bolivia. Only one species, M. parvilabius, is known from the Guyana shield (Luer, 1992). A. R. Endrs collected material later referred to as Myoxanthus scandens near the village of San Ramn, Costa Rica, around 1870. According to the collector’s vouchers deposited in W, the same species was also found in the vicinity of the old capital city of Costa Rica, Charles H. Lankester at La Estrella del Guarco, close to Cartago, and this specimen was eventually used by professor Ames to formally describe Myoxanthus scandens (as Pleurothallis scandens; Ames 1923). According to a manuscript note of the collector on the ABSTRACT: We present and illustrate a new species of Myoxanthus from Costa Rica, closely related to M. scandens. We present descriptions and illustrations of both species. The new species, Myoxanthus sotoanum differs from M. scandens, oblong, obtuse, subfalcate lateral lobes of the lip, the two keels extending nearly to the middle of the lip, and Myoxanthus sotoanum generally also occurs at lower elevations than M. scandens. RESUMEN : Presentamos e ilustramos una nueva especie de Myoxanthus para Costa Rica, junto a la descripcin e ilustracin de M. scandens basadas en material costarricense. La nueva especie , Myoxanthus sotoanum, es similar a M. scandens ptalos y el labelo amarillos, los lbulos laterales del labelo oblongos, obtusos y subfalcados, las dos quillas estigma . Myoxanthus sotoanum generalmente se encuentra a elevaciones inferiores que M. scandens. KEY WORDS : Myoxanthus scandens, Costa Rica, Myoxanthus sotoanum, new species, Orchidaceae

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.468 LANKESTERIANA raisin purple”. After studying specimens of this group in the collections of CR, INB, JBL, USJ and W, we noted that the name was apparently applied to two different additional distinguishing characters between these One of these species was recognized as Myoxanthus scandens, while the other is still in need of taxonomical recognition. With the aim of better comparing both entities, we describe and illustrate M. scandens as well as the species new to science. Myoxanthus scandens (Ames) Luer, Selbyana 7(1): 49. 1982. Pleurothallis scandens Ames, Sched. Orch. 5: 18-21. 1923. FIG . 1. TYPE. COSTA RICA . Cartago: La Estrella, C. H. A delightful scandent species occurring in forest-ridges. Flowers dull raisin purple (holotype, AMES; isotype, AMES). Plant epiphytic, caespitose, scandent, up to 60 cm long. Roots Ramicauls slender, cylindric, 0.1-0.15 cm in diameter, 0.6-30.0 cm long, enclosed by 3-13 tubular, pubescent sheaths 0.5-4.5 cm long. Leaves straight with the ramicaul, coriaceous, elliptic, conduplicate, acute, 2.5-11.0 x 0.5-1.0 cm, narrowly cuneate basally, subpetiolate. a fascicle of successive, singly, 2-4 mm long including the peduncle 2 mm long. Floral bracts tubular, ovate, acute, conduplicate, pubescent, 1 mm long. Pedicels 1 mm long, persistent. Ovary cylindric, 1.5 mm long. Flowers with brownpurple sepals and translucent green petals with longitudinalpurple stripes, the lip purple. Dorsal sepal ovate, acute, concave, 4.5 x 2.0 mm, 5-veined. Lateral sepals ovate, acute to obtuse, 4.0 x 1.5 mm, 3 to 5-veined, connate for approximately half their length, pubescent at the center of their common surface, the apex erose. Petals narrowly ovate, wider at the basal half, contracted into a thickened, apical, serrulate, acute tail, 4.0 mm long, 2.0 mm wide at the basal half. Lip oblong, scarcely trilobed, 3.0 mm long, hinged to the column foot, the lateral lobes near the middle, perpendicular, erect, the disc with a longitudinal pair of parallel lamellae from the lower third to near the apex, the base subtruncate with a pair of small lobules, the apex obtuse, minutely denticulate. Column short, stout, semiterete, 2 mm long, provided with an acicular, acuminate tooth on each side of stigma, the clinandrium denticulate, the anther incumbent, the stigma ventral. Pollinia two, pyriform, on an elliptic, recurved viscidium. MAT E RIAL STUDI E D : Costa Rica . Alajuela : [San Ramn], La Palma de San Ramn, [1300-1500 m], 29 Agosto 1924, A. M. Brenes 2291 (CR-34052). [Alajuela: San Ramn], La Palma de San Ramn, [1300-1500 m], 24 octubre 1924, A. M. Brenes 1536 (CR-26204). Cartago : 5.6 km San Jos side of road leading to Palmital, steep new dirt road up hill from Pan Am highway, 1700–1800 m, epiphytic on exposed tree near edge of extremely dark maroon almost black, 21 Setiembre 1979, K. S. Walter 79486 (CR). Oreamuno, Cipreses, 952’57”N, 8351’18” W, 1850 m, remnants of primary forest, 23 June 1998, F. Pupulin et al. 838 (JBL-Spirit). Oreamuno, Cipreses, 952’57”N, 8351’18” W, 1850 m, 23 June 1998, F. Pupulin et al. 837 (USJ). Turrialba, en cultivo en el Jardn Botnico Lankester, 25 enero 2001, M. Blanco 1820 (USJ). Heredia : Heredia, Vara Blanca, en los potreros y partes de bosque secundario pertenecientes al Pos Volcano Lodge cerca del centro de Vara Blanca, 1010’00” N, 8410’06” W, 1700 1900 m, 03 04 agosto 2006, Karremans (JBL-Spirit) (Fig. 2). Panama . Panama : road between El Llano and Cart, 300 m, 4 Mar. 1976, C. Luer et al. 1018 (MO, SEL, illustration); Llano-Cart road, 1 mile past saw mill on dirt road, 300-500 m, 11 Nov. 1979, C. Todzia et al. 1023 (CR). ETYMOLOGY : From the Latin scandens, “scandent” in allusion to the habit of the plant. DISTRIBUTION: Costa Rica and Panama. HABITAT AND ECOLOGY : Epiphytic in lower montane wet and very wet forests, and premontane rain forest at elevations between 1300 and 1900 m in Costa Rica, and epiphytic in premontane wet forests at elevations between 300 and 500 m in Panama. According to

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010. PUPULIN et al . — On the identity of Myoxanthus scandens469 FIGURE 1. Myoxanthus scandens (Ames) Luer. A — Habit. B — Detail of the habit. C — Flower. D — Dissected perianth. E — Ovary, column and lip, lateral view. F — Lip, spread. G — Pollinarium and anther cap. Drawn by D. Bogarn and A. Karremans 1322.

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LANKESTERIANA 9(3), January 2010. Universidad de Costa Rica, 2010.470 LANKESTERIANA February. Myoxanthus scandens is distinguished by the petals, lateral lobes of lip triangular, acute, erect with two keels extending to the apex of the lip, and the long, We are not including here Pleurothallis pennellia in the synonymy of M. scandens. The species was described in 1976 from Ecuador, on the basis of a Botanical Gardens (Luer, 1976). Luer (1976) noted the vegetative similarity to M. scandens, but described the be an artifact due to the juvenile, immature condition of the holotype specimen, and Luer later included it in the synonymy of M. scandens (Luer, 1992). The petals of P. pennellia are narrowly ovate and shortly obtuse at the apex, while in both M. scandens and M. sotoanum they are apically contracted into a thickened, acute tail. The lip of P. pennellia has two low keels above the middle, while these structures extend from the lower third to the middle of the lip in M. sotoanum, and from the base to near the apex in M. scandens. Furthermore, the clinandrium of P. pennellia is short, leaving the anther partially exposed, while M. scandens and M. sotoanum have a long, hooded clinandrium, completely covering the anther cap. In our opinion, P. pennellia is a true member of Myoxanthus, likely distinct from other Peruvian (or perhaps an Ecuadorian), unvouchered However, we refrain to make the new combination in Myoxanthus until we can study more material from the type locality. Myoxanthus sotoanum Fernndez sp. nov. TYPE: Costa Rica. Cartago: Jimnez, Pejibaye, La Marta, shore of Ro Pejibaye, 9”N 83”W, 690 m, epiphytic in secondary vegetation along the river, premontane wet forest, 10 Jan. 2004, F. Pupulin (holotype, CR; isotype, JBL-spirit). Fig. 2. Species Myoxantho scandenti (Ames) Luer aemulans, sed foliis linearibus, sepalis brunneis, subfalcatis, duobus carinis extensis fere dimidium labelli, dentibus lateralibus columnae acicularis acuminatis praecipue differt. Plant epiphytic, cespitose, scandent, up to 90 cm long. Roots Ramicauls slender, cylindric, 0.5 cm in diameter, 1-23 cm long, enclosed by 2-6 tubular, pubescent sheaths 0.8-4.0 cm long. Leaves straight with the ramicaul, coriaceous, elliptic, conduplicate, acute, 2.5-9.0 x 0.150.3 cm, narrowly cuneate, subpetiolate. apex of the ramicaul from a