Abstract
With more than 60 species, the Neotropics is the most species rich region among Vanilla. Currently, many Brazilian species have been described or rediscovered, and taxonomic realignments have been carried out based on integrative taxonomy. The accuracy in species delimitation is important for studies on phylogeny and ecology, and because some Brazilian vanilla seems to be inappropriate to human consumption. Vanilla is monophyletic, with three main lineages, two of which occur throughout the Neotropics. The pollination system of Neotropical Vanilla is not species-specific, and their flowers are commonly adapted to pollination by euglossine males. The pollination mechanism of euglossinophilous Vanilla is based on a combination of perfume collection and nectar seeking. At least one species, V. palmarum, is pollinated by hummingbirds representing an evolutionary disruption in the pollination system among Vanilla. With regards to seed dispersal, the mesocarp of species with dehiscent fruits are rich in raphid idioblasts that can be harmful to seed dispersers. For this reason, in Vanilla with dehiscent fruits, birds and mammals commonly access the sclerotic seeds through fruit valves. Seeds of Vanilla with indehiscent fruits are dispersed by terrestrial herbivores. Besides the nutritive rewards, fruits are rich in polyphenols that can be toxic or unpalatable to omnivores. The chemical scarification of seed coat by the digestive acids is crucial for synchronizing the biological processes involved in seed germination. The understanding of the processes involved in Vanilla reproduction is very important in the selection of favorable genotypes for commercial vanilla production.
References
Cameron, K. M. (2003). Vanilloideae. In: Pridgeon, A., Cribb, P., Chase, M.W., Rasmussen, F. (Eds.), Genera Orchidacearum. Oxford University Press, Oxford, 281–334. https://doi.org/10.1093/oso/9780198507116.001.0001
Pansarin, E. R., Menezes, E. L. F. (2023). A new remarkable Vanilla (Orchidaceae) endemic from Brazilian campos rupestres: their phylogenetic position and evolutionary relationships among Neotropical congeners. Phytokeys, 227, 151–165 https://doi.org/10.3897/phytokeys.227.101963
Pansarin, E. R. (2024a). Rediscovery and revalidation of the Brazilian endemic Vanilla schwackeana Hoehne (Orchidaceae): its distribution and phylogenetic position. Plant Ecology and Evolution, 157, 32–41. https://doi.org/10.5091/plecevo.110331
Pansarin, E. R. (2025a). Vanilla lindmaniana and V. palmarum (Orchidaceae) are distinct allopatric species. Plant Ecology and Evolution, 158, 53–62. https://doi.org/10.5091/plecevo.134103
Hoehne, F. C. (1945). Orchidáceas. In: Hoehne, F.C. (Ed.) Flora Brasílica, Fasc. 8 (Vol. XII, II; 13–43). Secretaria da Agricultura, Indústria e Comércio de São Paulo, São Paulo, 1–389 (+ 209 Tabs.).
Pansarin, E. R. (2010). Taxonomic notes on Vanilleae (Orchidaceae: Vanilloideae): Vanilla dietschiana, a rare south American taxon transferred from Dictyophyllaria. Selbyana, 30, 198–202.
Pansarin, E. R., Miranda, M. R. (2016). A new species of Vanilla (Orchidaceae: Vanilloideae) from Brazil. Phytotaxa, 267, 84–88. https://doi.org/10.11646/phytotaxa.267.1.9
Pansarin, E. R., Salatino, A., Pansarin, L.M., Sazima, M. (2012). Pollination systems in Pogonieae (Orchidaceae: Vanilloideae): A hypothesis of evolution among reward and rewardless flowers. Flora (Jena), 207, 849–861. https://doi.org/10.1016/j.flora.2012.09.011
Pansarin, E. R. (2022). Vanilla flowers: much more than food-deception. Botanical Journal of the Linnean Society, 198, 57–73. https://doi.org/10.1093/botlinnean/boab046
Pansarin, E. R. (2023). Non-species-specific pollen transfer and double-reward production in euglossine-pollinated Vanilla. Plant Biology, 25, 612–619. https://doi.org/10.1111/plb.13523
Pansarin, E. R. (2025b). Systematics of the Vanilla chamissonis complex (Orchidaceae): a study based on integrative taxonomy. Plant Ecology and Evolution, 185, 260-278. https://doi.org/10.5091/plecevo.154789
Soto Arenas, M. A., Cribb, P. (2010). A new infrageneric classification and synopsis of the genus Vanilla Plum. ex Mill. (Orchidaceae Vanillinae). Lankesteriana, 9, 355–398. https://doi.org/10.15517/LANK.V0I0.12071
Karremans, A. P., Chichilla, I. F., Rojas-Alvarado, G., Cedeño-Fonseca, M., Damian, A., Léotard, G. (2020). A reappraisal of neotropical Vanilla with a note on taxonomic inflation and the importance of alpha taxonomy in biological studies. Lankesteriana, 20, 395–497. https://doi.org/10.15517/lank.v20i3.45203
Batista, J. A. N., Castro, C., Sambin, A., Cruz-Lustre, G., Pansarin, E. R. (2023). Clarifying the identity of the Cleistes rosea complex (Orchidaceae) based on integrative taxonomy. Systematics and Biodiversity, 21, 2207575. https://doi.org/10.1080/14772000.2023.2207575
Fraga, C. N., Couto, D.R., Pansarin, E. R. (2017). Two new species of Vanilla (Orchidaceae) in the Brazilian Atlantic Forest. Phytotaxa, 296, 63–72. https://doi.org/10.11646/phytotaxa.296.1.4
Pansarin, E. R., Miranda, M. R. (2024). Clarifying the dubious identity of Vanilla dubia Hoehne (Orchidaceae) and its relatives. Plant Ecology and Evolution 157: 236–243. https://doi.org/10.5091/plecevo.121915
Soto Arenas, M. A., Dressler, R. L. (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. Lankesteriana, 9, 285–354. https://doi.org/10.15517/lank.v0i0.12065
Flanagan, N. S., Navia-Samboni, A., Vargas, W. G., Diaz-Rueda, D. M., Cruz, J. Y. S. R., Pansarin, E. R., Castano, A. (2025). Some taxonomic clarifications in Vanilla subgenus Xanata (Orchidaceae), and the resurrection of Vanilla calyculata. Kew Bulletin, 80, 1–14. https://doi.org/10.1007/s12225-024-10210-7
Hoehne, F. C. (1944). Orchidaceas novas para a flora do Brasil, dos hervarios do Instituto de Botânica, Jardim Botânico, Rio de Janeiro, e comissão de linhas telegráficas, estratégicas de Mato-Grosso ao Amazonas. Arquivos de Botânica do Estado de São Paulo, 1, 125–137.
Ferreira, A. W. C., Franken, E. P., Pansarin, E. R. (2020). Vanilla hartii Rolfe (Orchidaceae, Vanilloideae): first record for Brazil. Check List, Journal of Species List and Distribution, 16, 951–956. https://doi.org/10.15560/16.4.951
Krahl, D. R. P., Silva, J. B., Damian-Parizaca, A., Flanagan, N. S., Chiron, G., Krahl, A. H., Cantuaria, P. C., Pansarin, E. R. (2025). Unveiling the taxonomic identity of Vanilla sprucei (Orchidaceae, Vanilloideae): its phylogenetic position and new records for Brazil, Guyana and Venezuela. Acta Amazonica, 55, e55bc23227. https://doi.org/10.1590/1809-4392202302272
Pansarin, E. R. (2025c). Vanilla × robusta (Orchidaceae: Vanilloideae), the first natural Vanilla hybrid for South America. Nordic Journal of Botany, e04743. https://doi.org/10.1111/njb.04743
Carvalho, D. N., Meneguzzo, T. E. C., van den Berg, C. (2023). Vanilla calamitosa (Orchidaceae), a new aphyllous species from eastern Brazil. Kew Bulletin, 78, 309–317. https://doi.org/10.1007/s12225-023-10092-1
Pansarin, E. R., Aguiar, J. M. R. B. V., Ferreira, A. W. C. (2012). A new species of Vanilla (Orchidaceae: Vanilloideae) from São Paulo, Brazil. Brittonia (Bronx, N.Y.), 64, 157–161. https://doi.org/10.1007/s12228-011-9215-z
Zotz, G., Almeda, F., Arias. S., Hammel, B., Pansarin, E. R. (2021). Do secondary hemiepiphytes exist? Journal of Tropical Ecology, 37, 286–290. https://doi.org/10.1017/S0266467421000407
Pansarin, E. R., Ferreira, A. W. C. (2022). Evolutionary disruption in the pollination system of Vanilla (Orchidaceae). Plant Biology, 24, 157–167. https://doi.org/10.1111/plb.13356
Pansarin, E. R., Pansarin, L. M. (2014). Floral biology of two Vanilloideae (Orchidaceae) primarily adapted to pollination by euglossine bees. Plant Biology, 16, 1104–1113. https://doi.org/10.1111/plb.12160
Pansarin, E. R. (2024b). Specialized seed dispersal in Neotropical Vanilla reveals fruit unpalatability to omnivores. Plant Biology 26, 1185–1192. https://doi.org/10.1111/plb.13726
Pansarin, E. R. (2025d). Monkey as seed dispersers of Neotropical Vanilla involves social learning. Plant Biology 27, 584–591. https://doi.org/10.1111/plb.70018
Pansarin, E. R., Suetsugu, K. (2022). Mammal-mediated seed dispersal in Vanilla: Its rewards and clues to the evolution of fleshy fruits in orchids. Ecology, 103, ecy.3701. https://doi.org/10.1002/ecy.3701
Pansarin, E. R., Aguiar, J. M. R. B.V., Pansarin, L. M. (2014). Floral biology and histochemical analysis of Vanilla edwallii Hoehne (Orchidaceae: Vanilloideae): an orchid pollinated by Epicharis (Apidae: Centridini). Plant Species Biology 29, 242–252. https://doi.org/10.1111/1442-1984.12014
Rodolphe, G., Séverine, B., Michel, G., Pascale, B. (2011). Biodiversity and evolution in the Vanilla genus. In: Grillo, O., Venora G. (eds.) The dynamical processes of biodiversity - case studies of evolution and spatial distribution. InTech, Rijeka, pp. 1–27. https://doi.org/10.5772/24567
Anjos, A. M., Barbarena, F. F. V. A., Pigozzo, C. M. (2017), Biologia reprodutiva de Vanilla bahiana Hoehne (Orchidaceae). Orquidário 30, 67–79.
Chaipanich, V. V., Wanachantararak, P., Hasin, S. (2020). Floral morphology and potential pollinator of Vanilla siamensis Rolfe ex Downie (Orchidaceae: Vanilloideae) in Thailand. The Thailand Natural History Museum Journal 14, 1–14.
Pansarin, L. M., Pansarin, E. R., Sazima, M. (2008). Reproductive biology of Cyrtopodium polyphyllum (Orchidaceae): a Cyrtopodiinae pollinated by deceit. Plant Biology, 10, 650–659. https://doi.org/10.1111/j.1438-8677.2008.00060
Soto Arenas M. A. (1999). Filogeografía y recursos genéticos de las vanillas de México. Informe final SNIB-CONABIO proyecto No. J101. Instituto Chinoin AC, México DF:
Householder E. et al. (2010). Diversity, natural history and conservation of Vanilla (Orchidaceae) in Amazonian wetlands of Madre De Dios, Peru. Journal of the Botanical Research Institute of Texas 4, 227–243. https://doi.org/10.17348/jbrit.v4i1.8365
Faegri, K., van der Pijl, L. (1979). The principles of pollination Ecology. Pergamon Press, Oxford.
Stebbins, G. L. (1957). Self-fertilization and population variability in the higher plants. The American Naturalist 91, 337–354. https://doi.org/10.1086/281999
van der Pijl, L., Dodson, C. H. (1966). Orchid flowers: their pollination and evolution. University of Miami, Coral Gables.
Pansarin, E. R. (2021). Unravelling the enigma of seed dispersal in Vanilla. Plant Biology, 23, 974–980. https://doi.org/10.1111/plb.13331
Pansarin, E. R. (2025e). Epiphyte Vanilla relies on birds as long-distance seed dispersers. Plant Biology, 27,1497–1504. https://doi.org/10.1111/plb.70086
Schaefer, H. M., Ruxton, G. D. (2011). Plant-animal communication. Oxford Univ. Press, Oxford, UK. https://doi.org/10.1093/acprof:osobl/9780199563609.001.0001
Jordano P. (2000) Fruits and frugivory. In Fenner M. (ed.). Seeds: the ecology of regeneration in plant communities: 2nd ed. CAB International, Wallingford, UK, p.125-165. https://doi.org/10.1079/9780851994321.0125
Karremans, A. P., Bogarín, D., Otárola, M. F., Sharma, J., Watteyn, C., Warner, J., Herrera, B. R., Chinchilla, I. F., Carman, E., Valerio, E. R., et al. (2023a). First evidence for multimodal animal seed dispersal in orchids. Current Biology, 33, 364–371. https://doi.org/10.1016/j.cub.2022.11.041
Karremans, A. P., Watteyn, C., Scaccabarozzi, D., Pérez-Escobar, O. A., Bogarín, D. (2023b). Evolution of seed dispersal modes in the Orchidaceae: has the Vanilla mystery been solved? Horticulturae, 9, 1270. https://doi.org/10.3390/horticulturae9121270
Sierra-Vásquez, F., Castro-Luna, A. A., Espinosa-Francisco, E. K., Andrade-Torres, A. (2025). Molecular evidence of frugivorous bats (Chiroptera: Phyllostomidae) consuming Vanilla planifolia (Orchidaceae). Studies on Neotropical Fauna and Environment, 1–6. https://doi.org/10.1080/01650521.2024.2440986
Hiller, J. L., Benda, G. I., Rahatzad, M., Allen, J. R., Culver, D. H., Carlson, C. V., Reynolds, J. W. (1986). Benzyl alcohol toxicity: impact on mortality and intraventricular hemorrhage among very low birth weight infants. Pediatrics, 77, 500–506.
Granato, D., Mocan, A., Câmara, J. S. (2020). Is a higher ingestion of phenolic compounds the best dietary strategy? A scientific opinion on the deleterious effects of polyphenols in vivo. Trends in Food Science & Technology, 98, 162–166. https://doi.org/10.1016/j.tifs.2020.01.010

