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 Dedication
 Front Matter
 Index
 Title Page
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Title: Lankesteriana
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Permanent Link: http://ufdc.ufl.edu/UF00098723/00024
 Material Information
Title: Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Physical Description: v. : ill. (some col.) ; 25 cm.
Language: English
Creator: Jardi´n Bota´nico Lankester
Jardín Botánico Lankester
Publisher: Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Place of Publication: Cartago Costa Rica
Cartago Costa Rica
Publication Date: December 2009
Frequency: three times a year[2002-]
irregular[ former 2001]
three times a year
regular
 Subjects
Subject: Botany -- Periodicals -- Costa Rica   ( lcsh )
Epiphytes -- Periodicals -- Costa Rica   ( lcsh )
Orchids -- Periodicals -- Costa Rica   ( lcsh )
Plantkunde   ( gtt )
Botanische tuinen   ( gtt )
Genre: periodical   ( marcgt )
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
Bibliographic ID: UF00098723
Volume ID: VID00024
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 48491453
lccn - 2001240973
issn - 1409-3871

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Table of Contents
    Cover
        Cover
    Dedication
        Dedication
    Front Matter
        Front Matter
    Index
        Page i
        Page ii
        Page iii
    Title Page
        Title Page
    Main
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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.


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








LANKESTERIANA


(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


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








LANKESTERTANA


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.








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,
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.








LANKESTERIANA


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

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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


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"* 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

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









LANKESTERTANA


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









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








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

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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
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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

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








LANKESTERTANA


*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.


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








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.
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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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