Origins and Dispersal of Cultivated Vanilla (Vanilla ...

1 Origins and Dispersal of Cultivated Vanilla ( Vanilla planifoliaJacks. [Orchidaceae])1 PESACHLUBINSKY*,2,S VERINEBORY3,JUANHERN NDEZHERN NDEZ4,SEUNG-CHULKIM2,ANDARTUROG MEZ-POMPA22 Department of Botany and Plant Sciences, University of California, Riverside, CA, USA3 CIRAD, Station de Ligne Paradis, 97410, St-Pierre, La R union4 INIFAP, Mart nez de la Torre, 93600, Veracruz, M xico*Corresponding author; e-mail: and Dispersal of Cultivated Vanilla (Vanilla planifoliaJacks. [Orchidaceae]). Vanilla is a clonally propagated crop originating from Mesoamerica. Information regardingthe circumstances under which Vanilla cultivation began is incomplete.

2 Presumably, the Totonacpeople of Papantla (north-central Veracruz, Mexico) were the earliest to cultivate Vanilla ; however,the oldest reports of Vanilla use relate to the pre-Columbian Maya of southeastern Mexico/CentralAmerica, where Vanilla was a cacao-beverage spice. We utilized Amplified Fragment Length Poly-morphism (AFLP) marker diversity to infer the Origins and relationships among Cultivated andnon- Cultivated Vanilla in Mesoamerica and on islands in the Indian Ocean, which comprise today sprincipal production regions of Vanilla . Our results suggest that, genetically, Vanilla cultivatedoutside of Mesoamerica is most closely related to Cultivated stock from Papantla; whereas uniqueclones ofV.

3 Planifoliaare found in non- Cultivated and Cultivated individuals from elsewhere inMesoamerica. This is consistent with a single origin for Cultivated Vanilla outside of Mexico,along with multiple Origins for Cultivated material within Mexico. These data suggest that ves-tiges of pre-Columbian Maya Vanilla cultivars are not found in commercial production Words: Vanilla ; Orchidaceae; Mesoamerica; AFLP; crop is a value added spice crop produced intropical countries and consumed principally in theUnited States, France, and Germany (FAOSTAT).Most production occurs at the household level andinvolves tens of thousands of farmers is carried out in a supply chain thatstarts with local trade at the community andregional level, followed by re-sales to variouscurers/exporters, international brokers, and whole-salers/extract manufacturers (Ecott2004; Rain2004).

4 Two clonally propagated species are com-mercialized for their cured, fermented fruits: Vanilla planifoliaJacks. ( , Mexican or Bour-bon Vanilla ) andV. tahitensisJ. W. Moore( Tahitian Vanilla ) (Childers et ;Correll1953; ; Sauer1993). Of theestimated 5,000 to 10,000 metric tons (Mt) ofvanilla-bean production per annum, roughly 95%derives fromV. planifolia, with most coming fromMadagascar and Indonesia (FAOSTAT).The genusVanillaPlumier ex Miller (ca. 110species) belongs to one of the basal subfamilies ofthe Orchidaceae, the Vanilloideae (Cameron et ). The early origin ofVanillaamong theorchids is evidenced by its numerous primitivetraits such as fruits that arefleshy as well as hard,sclerotic seed coats (Cameron and Soto Arenas2003).

5 According to molecular clock estimates,Vanillais around 65 million years old (Ram rezet ). This age supports the hypothesis thatthe pan-tropical distribution of the genus, whichis rare among orchids, has resulted from theseparation of continental landmasses (Cameronand Soto Arenas2003). The onlyVanillareported to have aromatic fruits, about 30 species,Economic Botany, 62(2), 2008, pp. 127 138 2008, by The New York Botanical Garden Press, Bronx, NY 10458-5126 30 January 2008; accepted 11 April 2008;published online 5 June from the neotropics and the Caribbean, withthe exception ofV. tahitensis(Soto Arenas1999).

6 The principal commercial Vanilla species, (2n=32), is a rare, perennial climbingherb endemic to humid evergreen forests inMesoamerica, on karst substrates between 250and 750 meters above sea level (masl), in regionsthat receive more than 2,500 millimeters (mm) ofannual precipitation (H gsater et ;Port res1954; Soto Arenas1999; Tanaka andKamemoto1984). It is occasionally sympatricwith several other aromaticVanillaspecies in-cludingV. pomponaSchiede,V. odorataC. Presl,V. insignisAmes, andV. hartiiRolfe, as well aswith non-aromatic species likeV. inodoraSchiede(Soto Arenas1999).The Aztec elite and merchant class esteemedvanilla for itsflavor and rarity.

7 For his pleasuregarden in Huaxtepec, Moctezuma dispatchedmessengers to the tropical coast region with arequest to the Lord of Cuetlaxtla [San Juan deUlua, Veracruz] for plants with roots of thevanilla orchid (Nuttall1925; p. 456). Theseplants presumably came from the region ofPapantla (Tecolutla river basin), in the north-central section of what is today the state ofVeracruz. Papantla and its environs, whichinclude the Classic Period ( 250 900)archaeological site of El Taj n, are home to theTotonac people. In the eighteenth century,Papantla emerged as thefirst and only region inthe world to produce Vanilla for internationalexport (Kour 2004).

8 The sole destination forPapantla Vanilla was the European market, whichdemanded Vanilla as an ingredient for cacao-basedconfections (Coe and Coe1996). Cultivation ofvanilla by the Totonac was integrated into asystem of swidden agriculture whereinmilpafallows were managed as Vanilla agro-forests(Kelly and Palerm1952; Medell n-Morales1988; Toledo et ). Papantla lost itsvanilla monopoly in the mid-nineteenth century,when Vanilla production took off in Frenchcolonies in the Indian Ocean, most notablyMadagascar (Kour 2004). Mexico remains aminor supplier of Vanilla today, accounting forless than 1% of global production (FAOSTAT),with 20 30 Mt of cured Vanilla being producedannually in the Papantla Vanilla has been a Papantla cash cropfor roughly 250 years, there is deficient evidenceto suggest that it was a plant of any internalcultural value to the pre-Columbian Totonac.

9 Inthe original 1740s record of Vanilla use fromPapantla, Vanilla is gathered, not Cultivated , in thesurrounding forests and sold in town (Bruman1948). While observers claimed that this was elmodo que siempre se ha tenido [the way it hasalways been], a 1662 list of cash crops fromPapantla only mentions tobacco and chili pep-pers, without citing Vanilla (Bruman1948).The earliest suggestions of Vanilla gatheringand/or cultivation anywhere in pre-ColumbianMesoamerica contain two elements: They comefrom the Maya lowlands and they are ancillary toobservations of cacao (Theobroma cacaoL.)production (Bruman1948; Caso Barrera andFern ndez2006;Kour 2004).

10 Cacao-basedbeverages ( chocolate ) have been consumed inthe Maya lowland area, where cacao growsnaturally, for at least 2,500 years (Henderson ; Hurst et ; McNeil2006; Millon1955). During the Late Post-Classic ( 1350 1500), a trade in cacao, which had started in theMaya area and among the Maya elite, grew toencompass all of Mesoamerica, with the Aztecrulers of the Valley of Mexico (where cacao doesnot grow well) serving as the dominant, centrip-etal force (Bergmann1969). Plants that were usedto spice cacao and that grew sympatric with areasof cacao cultivation in the Maya lowlands becameincorporated into cultivation and trade.