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Redalyc.A REVISION of the MEXICAN and CENTRAL Lankesteriana International Journal on Orchidology ISSN: 1409-3871 [email protected] Universidad de Costa Rica Costa Rica Soto Arenas, Miguel A.; Dressler, Robert L. 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 Lankesteriana International Journal on Orchidology, vol. 9, núm. 3, enero, 2009, pp. 285- 354 Universidad de Costa Rica Cartago, Costa Rica Available in: http://www.redalyc.org/articulo.oa?id=44339819002 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative LANKESTERIANA 9(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†2,3 & ROBERT L. DRESSLER4,5 2 Instituto de Ecología, UNAM, Circuito Exterior s.n. Ciudad Universitaria, Coyoacán 04510 México D.F., MEXICO 3 Herbario AMO, Montañas Calizas 490, México, D.F. 11000, MEXICO 4 Lankester Botanical Garden, University of Costa Rica, P.O. Box 302-7050 Cartago, Costa Rica 5 Corresponding author: [email protected] ABSTRACT. We present a revision of the Mexican and Central American species of Vanilla. There are 15 different species in the area; Vanilla costaricensis, V. cribbiana, V. dressleri, V. martinezii and V. sarapiquensis are here proposed as new taxa, and V. pompona subsp. pittieri and V. pompona subsp. grandiflora are recognized at subspecific rank. Vanilla calyculata, V. hartii, V. helleri, V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia, V. pompona 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 presenta una revisión de las especies mexicanas y centroamericanas de Vanilla. Existen 15 especies reconocidas en el área. Vanilla costaricensis, V. cribbiana, V. dressleri, V. martinezii y V. sarapiquensis se proponen aquí como nuevas especies, V. pompona subsp. pittieri y V. pompona subsp. grandiflora también se reconocen con status subespecífico.Vanilla calyculata, V. hartii, V. helleri, V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia, V. pompona y V. trigonocarpa también se describen, ilustran y se discuten aspectos de su nomenclatura, tipificación, distribución, relaciones y otros puntos de interés. Adicionalmente, se incluye una clave de identificación de las especies. Varias especies mexicanas y centroamericanas de Vanilla están cercanamente relacionadas a V. planifolia y V. pompona, las dos especies más frecuentemente cultivadas, y por lo tanto representan un germoplasma importante para este cultivo. Adicionalmente se presenta un análisis 1 Editor’s note: Miguel Ángel 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 Hágsater, 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. 286 LANKESTERIANA cladístico de secuencias nucleotídicas de la región de los espaciadores internos transcritos (ITS) de los genes nucleares ribosomales, donde se muestra que las secuencias de este marcador molecular ampliamente utilizado son específicas, lo que permite, salvo algunas excepciones que se discuten, el reconocimiento de muestras estériles y además lo hacen un buen marcador molecular para estudios filogenéticos en este género. Se comenta la similitud y relación entre V. tahitensis* y V. odorata, y el origen híbrido de la primera. KEY WORDS: Orchidaceae, Vanilla, Vanilla calyculata, V. hartii, V. helleri, V. inodora, V. insignis, V. odorata, V. phaeantha, V. planifolia, V. pompona, V. trigonocarpa, Mexico, central Americas, new species The Pantropical genus Vanilla Plumier ex Miller is the hemiepiphytic growth habit (e.g. leafy and leafless a group of orchid vines with about 107 species (Soto shoots and differently shaped leaves on the same Arenas, 2003; Soto Arenas & Cribb, 2010). Vanilla individual plant as in many hemiepiphytic aroids; Putz is an ancient genus (Chase, 2001; Cameron, 2000, & Holbrook, 1986; Ray, 1990), the poor preservation 2003) and the largest in the subfamily Vanilloideae of the membranaceous, strongly three-dimensional (Soto Arenas, 2003). Some Vanilla species are flowers make them difficult to study, the notable grown as the source of an extract for flavoring and floral similarities between many species, in part due perfumery, especially Vanilla planifolia Jacks., with to the pollination by Euglossine bees (which permits V. x tahitensis J.W.Moore and V. pompona Schiede the maintenance of species with little morphological being much less cultivated (Correll, 1944; Purseglove, differentiation but with different floral fragrances (in 1975). Vanilla exports generate US $60-80 million in other genera of orchids); Williams, 1982), and the foreign exchange for producing countries (Smith et problems derived from the less than perfect quality al., 1992), perhaps more at present, and it is the most of type specimens. Additionally, some species are profitable crop of the warm tropics. very rare, with sparse populations, and the vines are The culture of V. planifolia and V. pompona long-lived perennials that only flower when they have originated in Mexico, probably in northern Veracruz attained considerable size. Small pieces of these plants (Bruman, 1948). Although frequently cited as native of may be transplanted to greenhouses or gardens, but Mexico and elsewhere, there were until recently very few of them ever attain the size and strength to flower. few records of V. planifolia from wild populations, There are abundant data that indicate that V. planifolia and its precise original distribution is uncertain (Soto is in danger of extinction in Mexico (Soto Arenas et Arenas, 1999). Many Vanilla species are similar al., 2004). Scarcity of material can be illustrated by the to V. planifolia in floral and vegetative traits, and flowering behaviour of the only Mexican population abundant material in herbaria, both flowering and of V. hartii, which was found in anthesis only after 7 sterile, is usually misidentified as V. planifolia. Even years of observations. Vanilla helleri and V. martinezii some Mexican plantations planted with wild Oaxacan are each known only from two flowering collections, Vanilla vines include a mixture of V. cribbiana, V. while V. costaricensis and V. sarapiquensis are each insignis, V. odorata, and V. pompona, besides the known from a single pressed specimen. Vanilla true V. planifolia. Similar “mixed” plantations occur phaeantha and V. helleri are here reported for the first in Ecuador (P. Lubinsky, pers. com.) and Guatemala time from Mexico, based on sterile material, whose (Beza, pers. com.). ITS sequences match with properly identified material It is well-known that Vanilla has peculiarities that of these taxa. Most cultivated specimens of Vanilla in make it a taxonomically difficult genus (Wood, 2003), botanic gardens and living collections never produce mostly due to the scarcity of flowering material (a flowers and therefore cannot be identified. result of the gregarious, ephemeral flowering, usually Vanilla plantations face several agricultural at the top of the forest canopy), the huge vegetative problems, the most important being the root rot variation and phenotypic plasticity characteristic of disease caused by Fusarium batatis f. vanillae (Childers et al., 1959). It has
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