identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
F87A879DFFEDFFAB0194FDCCFB5C930D.text	F87A879DFFEDFFAB0194FDCCFB5C930D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vanilla , Miller 1754	<div><p>Vanilla Miller (1754: without page number), nom. cons. prop.</p><p>Type species:— Vanilla planifolia Andrews (1808: 538), typ. cons. prop. (Karremans &amp; Pupulin 2023).</p><p>Etymology:—Latinized form of the Spanish vainilla, meaning small vaina, a tight-fitting holster for knives or swords, most likely in allusion to the shape of the fruit.</p><p>Replaced lectotype:— Vanilla mexicana Miller (1768: without page number).</p><p>= Vanillophorum Necker (1790: 134), nom. inval., opus utique oppr.</p><p>= Myrobroma Salisbury (1807: t. 82).</p><p>Type species:— Myrobroma fragrans Salisbury (1807: t. 82), nom. illeg. = Epidendrum rubrum Lamarck (1783: 178) = Vanilla rubra (Lam.) Urban (1920: 157) .</p><p>= Dictyophyllaria Garay (1986: 231) .</p><p>Type species:— Dictyophyllaria dietschiana (Edwall) Garay (1986: 231) = Vanilla dietschiana Edwall (1903: 192) .</p><p>= Miguelia Averyanov (2011: 45) .</p><p>Type species:— Miguelia somae (Hayata) Averyanov (2011: 49) = Vanilla somae Hayata, Icon. Pl. Formosan. 6: 88. 1916.</p><p>Description:—Nomadic, epiphytic, lithophytic or terrestrial, herbaceous vines, usually germinating in decomposing organic matter or humus rich soil. Plants typically notoriously dimorphic. Plantlets erect, producing running horizontal stems, often with scale-like leaves, that locate phorophytes or other tutors by means of negative phototropism. Hosts not specific, with apparent exceptions. Mature stems scandent, zigzagy, sparsely or profusely branched, terete or quadrangular, sometimes sulcate, slender to thick, brittle to succulent, smooth to warty, up to 20 or more meters in length, typically erect when mature, reaching the apex of the host, then horizontal and pendent, often transitioning between these; mature stems able to reach the ground again and transforming into horizontal runners with scale-like leaves until climbing on a new tutor. Leaves and stems of runners and new growths often covered by a removable, glaucous wax, that is sometimes retained in mature plants. Roots of three kinds; the first are terrestrial roots found only at the base of the stem, thick, terete, and villose when in soil; the second kind short, usually flattened, slender and smooth, produced at each node, growing perpendicular to the stem and used to clasp the tutor; the third kind are elongate, mostly aerial roots, produced by nomadic vines and originating along the stem nodes, growing downwards parallel to the stem until they reestablish the connection with the ground, especially when the stem is severed. Leaves non-articulate, distichous, mostly with well-developed blades, in several species retaining the scale-like leaves typical of juvenile growths or mature runners, which are often deciduous; sessile to petiolate, the blades narrowly lanceolate to oblong or elliptic, soft-membranaceous to thick coriaceous. Inflorescence mainly axillary, rarely apical, racemose, sometimes branched, congested or scant, short or elongate, acropetalous, with two to a few dozen flowers, usually a single flower open at a time; bearing short or large floral bracts, either similar or quite distinct to the vegetative axis. Floral bracts often small, scale-like, at times foliaceous. Ovary articulate to the perianth, sulcate, smooth, rarely granulose, unilocular, rarely provided with an inconspicuous calyculus, typically incurved, straightening a few days after fecundation and becoming recurved after that. Perianth deciduous after a few days except when the flowers are fertilized, then retained longer. Extrafloral nectar produced from the floral bracts and calyculate apex of the ovaries of certain species. Flowers usually showy, short-lived, lasting from a few hours to a few days; sepals and petals usually green, yellow, cream or white, the lip usually similar in color, often stained with pink, red or purple; produced in acropetalous succession, mostly gullet-shaped, resupinate, mostly with horizontal or sub-pendular orientation, with the perianth deciduous, sometimes strongly fragrant, mostly nectarless. Sepals free, spreading, flat to contorted and undulate, membranaceous to fleshy, usually smooth, apex sometimes thick, granulose or warty. Petals free, often dorsally keeled, spreading, reflexed or undulate-contorted, margins mostly straight, occasionally minutely serrulate apically. Lip diversely fused to the column margins, merely at the base to almost completely to the apex, forming an open, deeply saccate or funnel-shaped chamber, in most species with a narrow, papillose-hirsute, dry, basal nectary formed by the fusion of the column and the lip’s claw; the lip blade simple to variously lobed, usually long-clawed, disc with longitudinal keels, trichomes, warts or thickened-rugose veins, often with a penicillate callus formed by a cluster of fimbriate, retrorse scales, placed below the column apex. Column semiterete, straight, sigmoid or arcuate, glabrous to pubescent on the ventral surface, sometimes with basal keels, mostly footless, provided with apical, more or less developed, thin to thick wings. Stigma forming a transversely rectangular to transversely elliptic slit-like cavity with thickened, sticky margins, or more often enclosed and completely hidden between the apical, flap-like, rostellar lobe and a stigmatic basal lobe exerting into a recurved flap. Anther versatile, terminal, semi-erect, incumbent to ventral, non-articulate, often subtended by a distinct, transversely rectangular, flat filament remnant; connective conspicuous, sometimes with horn-like projections formed by the incumbent twist of the anther. Pollen in monads, forming two or four, easily disrupted masses, without accessory structures, but sticky and sometimes removed as a triangular unit. Fruit mostly a dehiscent capsule to slightly dehiscent berry, sometimes indehiscent, usually opening along two lines and forming unequal valves, mostly fragrant. Seeds small, obovoid, black, with a sclerotic, crustose seed coat, surface smooth to slightly warty.</p><p>The previous description is based on that of Soto Arenas (2003), significantly modified and expanded to include detailed personal observations by the authors. It is important to note that many characteristics of Vanilla plants and flowers are often neglected or misinterpreted in literature, and require further attention. It is rarely noted where plants germinate, whether or not their stems, leaves and roots are dimorphic and ornate in any way, if the inflorescence are axillary or terminal and arise from erect, horizontal or pendent stems, if extrafloral nectar and floral fragrances are produced, or if the fruits are naturally dehiscent or not, and if they are truly non-aromatic. Plants are often described as hemiepiphytes, which they are not. They are typically nomadic vines (but see exceptions cited below), usually germinating on the ground and then becoming climbers that don’t lose the connection to the soil or do so only temporarily. The inflorescence has been described as cymose or paniculate in some cases, but the illustrative and photographic evidence is inconclusive (see discussion below). The flowers of the relatives of V. planifolia have a well-developed nectary formed by the fusion of the claw of the lip and the column base. The nectary has plenty of papillae and trichomes, and may offer traces of exudates, but is often dry, and usually does not seem to elicit continued consumption and visitation by any animals. There seem to be few exceptions, the most notable is V. hartii Rolfe (1899: 133), which has measurable quantities of sweet nectar stored in an accordingly bloated nectary. Unlike most orchids, Vanilla flowers don’t have a true anther cap, but rather a versatile, flexible and quasi-ventral anther, nor do they have pollinia or a pollinarium, but rather loose masses of pollen monads. Fruits are often said to be non-aromatic when lacking the characteristic vanilla smell, but we have yet to find a Vanilla fruit that truly has no odor at all. Most, if not all, have a smell, albeit not always particularly strong or appealing. More careful characterization and documentation of the ecological, vegetative and flower features of Vanilla would be most useful.</p><p>Typification:—Upon publishing the name Vanilla, Miller (1754) did not select a type species. He included three species concepts detailed as “1. Vanilla flore viridi &amp; albo, fructu nigrescente ”, “2. Vanilla flore violaceo, fructu breviori rubro ” and “3. Vanilla flore albo, fructu breviori corallino ”. Of the first he noted that “The Fruit of these Plants is call’d by the Spaniards, in America, Vanilla, or Vinello; and is much used by them to scent their Chocolate. It is the first Species here mention’d, which is chiefly esteem’d. This grows plentifully in the Bay of Campechy, in the West-Indies; where they are usually sold for about Three-pence each Fruit, English Money”. He would later formally name this species of commerce Vanilla mexicana Miller (1768) . Mansfeld (1959) typified the generic name Vanilla with V. mexicana, which he considered, at least in part, to be a synonym of V. planifolia Andrews (1808) .</p><p>The typification of the name Vanilla mexicana itself has long been unclear. In the protologue of V. mexicana, Miller (1768) cited a single element (“ Volubilis siliquosa Mexicana, plantaginis folio. Cat. Car. 3. p. 7”) referring to an illustration published by Catesby (1747). Although not explicitly stated, Catesby’s illustration is without a doubt based on those of Plumier (1703), the originals of which are kept at the Muséum National d’Histoire Naturelle in Paris (reproduced in Karremans et al. 2020). Despite its specific epithet chosen by Miller, the plant depicted by Plumier and reproduced by Catesby was not of Mexican origin, but from Haiti. Careful inspection by Soto Arenas &amp; Dressler (2010) revealed that the illustration reproduced by Catesby is not the species of Vanilla that was cultivated in Mexico and much used by the Spaniards to scent their chocolate, as early authors such as Miller believed. In fact, not only is the Antillean taxon not known to occur in Mexico, it is not in cultivation on account of its non-aromatic fruits (Soto Arenas &amp; Cribb 2010; Karremans et al. 2020), but the two are now known to belong to separate subgenera and be quite distantly related (Bouétard et al. 2010). As clearly stated in the protologue, Miller had the aromatic, cultivated species in mind when describing Vanilla mexicana, mistakenly believing it was the same species illustrated by Plumier, and reproduced by Catesby. Given that Miller did not cite any other material under V. mexicana we must select the illustration by Catesby as the type for the name. However, Karremans &amp; Pupulin (2023) put forward a formal proposal to conserve the generic name Vanilla with a conserved type, replacing the non-aromatic V. mexicana for the well-known, widespread, commercially cultivated, aromatic V. planifolia Andrews as generitypus. This proposal, recommended by the Nomenclature Committee for Vascular Plants (Applequist 2024) and accepted by the General Committee at the XX International Botanical Congress (Wilson 2025), not only reflects the original intent of Miller, but also prevents any future attempt to segregate the aromatic species of Vanilla from the genus. However, it has the unintended consequence that the subgeneric names Vanilla subgen. Vanilla and Vanilla subgen. Xanata become synonyms as they are both typified by V. planifolia .</p><p>The authority of the generic name Vanilla is often incorrectly ascribed to Plumier in literature. The names ‘Vanilla’, ‘Vanillia’ or ‘Vaynilla’ had indeed been used prior to Miller’s formal publication in 1754. In 1703, Charles Plumier used the name as part of a polynomials to refer to species belonging to the genus. But these are all prior to the starting date for binomial botanical nomenclature in 1753. Genus Vanilla was validly published by Miller without any ascription to Plumier, or any other author, which means that their inclusion in the authority is taxonomically incorrect. To prevent arguments alleging that Art. 46.8 (Turland et al. 2018) provides that ascription of a name is to be accepted when indicated by typographical or stylistic distinctions in the text, such as the diagnosis of Vanilla published by Miller - which is written in cursive -, it must be noted that this specific typographic style is used throughout the entire text (Miller 1754) in the description of Miller’s own new genera. While it is true that starting with the seventh and all subsequent editions of his Dictionaire, Miller (1759) ascribed the name Vanilla to Plumier, Art. 46.8 (Turland et al. 2018) also states that in determining the correct author citation, “only internal evidence in the publication as a whole” must be used, while Art. 46.9 (Turland et al. 2018) is emphatic in that the use of external evidence to determine authorship of nomenclatural novelties included in a publication is advisable only “where there is no internal evidence of authorship”. Exactly the same occurs with the generic name Ceiba Miller (1754: without page number), which had also been used previously by Plumier, but cannot be formally ascribed to the latter.</p><p>Finally, another element of debate is the etymology of the specific epithet “ planifolia ” when used in Vanilla, provided that the flat leaves are a rather standard feature among members of the genus. Although not explicitly stated by Andrews, when historical elements are taken into account it becomes evident that the flat leaves to which the author alludes are the sepals and petals of the flower. In the protologue, Andrews mentions that his species has been confused with that of Plumier’s first species ( V. mexicana Miller), stating that “seldom seen two species of one genus so different in the blossoms”. Both Plumier (1703) and Miller (1754) describe this plant as having “an anomalous flower, consisting of six leaves”, and indeed V. mexicana is readily recognized by the notoriously twisted, undulate sepals and petals, which easily distinguishes it from V. planifolia, which has flat sepals and petals.</p></div>	https://treatment.plazi.org/id/F87A879DFFEDFFAB0194FDCCFB5C930D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Karremans, Adam P.;Pupulin, Franco;Damián-Parizaca, Alexander	Karremans, Adam P., Pupulin, Franco, Damián-Parizaca, Alexander (2025): A revised infrageneric classification of Vanilla (Orchidaceae) based on phylogenetic, morphological and biogeographical evidence. Phytotaxa 715 (3): 207-228, DOI: 10.11646/phytotaxa.715.3.2, URL: https://doi.org/10.11646/phytotaxa.715.3.2
F87A879DFFE1FFA80194FBB2FA2194E1.text	F87A879DFFE1FFA80194FBB2FA2194E1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vanilla (Vanilla)	<div><p>1. Vanilla subgen. Vanilla</p><p>= Vanilla subgen. Xanata Soto Arenas &amp; Cribb (2010: 359) .</p><p>Type species: — Vanilla planifolia Andrews (1808: t. 538).</p><p>= Vanilla subsect. Papillosae Portères (1951: 290), nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>= Vanilla subsect. Lamellosae Portères (1951: 291) . nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>= Vanilla subsect. Papillosae Portères (1954: 141), nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>= Vanilla subsect. Lamellosae Portères (1954: 148), nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>Description:—Members of Vanilla subgen. Vanilla are nomadic or epiphytic vines, germinating in decomposing organic matter on the ground or epiphytically. Leaves usually well-developed and thick coriaceous. Inflorescence racemose, mainly axillary, occasionally apical, producing two to many flowers, the floral bracts usually small, typically differing notoriously from the leaves. Some species offer floral fragrances and thus can be highly aromatic. The sepals and petals are coppery, greenish, yellowish or whitish, narrow and often similar in color to the lip. In fragrant species the sepals thickened, often warty near the apex. Petal with a prominent abaxial keel, apical margins often minutely serrulate. The lip forming a distinct floral tube, with the lip margins fused to the column for more than half of its length to form a narrow, papillose-hirsute, dry, basal nectary; the flowers appear, however, nectarless (except in V. hartii). The lip blade usually with three distinct sites of ornamentation: a papillae-rich basal claw, a distinct penicillate callus of retrorse scales (except in V. bicolor Lindley 1838: misc. 37 and V. palmarum Lindley 1840: pl. 436) placed above the middle of the blade, just below the rosteller flap of the column, and clumps of warts or papillae on a thickened plate or forming elevated keels near the apex of the lip. The column is straight or bent, often with a hirsute ventral surface, with a well-developed rostellum and another flap-like, retrorse proximal stigma lobe. Pollen in monads, forming two easily disrupted masses. The fruits calyculate, either dehiscent or indehiscent, sometimes differing within the same species, non-aromatic (V. sect. Palmarum) or aromatic (V. sect. Vanilla), the latter containing vanillic and anisic aromatic compounds.</p><p>Discussion:— Vanilla subgen. Vanilla includes 40 species and two nothospecies divided into two sections: Vanilla sect. Vanilla, including all commercially grown, aromatic vanilla, and Vanilla sect. Palmarum Karremans, Damián &amp; Pupulin. Our circumscription of Vanilla subgen. Vanilla aligns completely with Vanilla sect. Xanata sensu Soto Arenas &amp; Cribb (2010) and is strongly supported by multiple lines of evidence: biogeographically, through its exclusively Neotropical distribution; ecologically, by its predominantly allogamous breeding system and reliance on euglossine bees for pollination; and by DNA data across multiple scales. We tentatively include their so-called ‘ V. penicillata group’, made up of the cryptic V. calamitosa and V. penicillata Garay &amp; Dunst. (in Dunsterville &amp; Garay 1965: 324), in this subgenus pending further studies. These two species are odd among Neotropical Vanilla in that they are the only ‘leafless’ species occurring on the mainland, featuring the reduced or scale-like leaf habit. Soto Arenas &amp; Cribb (2010) listed the group among the relatives of V. planifolia on account of the presence of the tubular lip adnate to the column, the blade with a penicillate callus, and calyculate fruits, and we agree with this interpretation. However, the greenish-cream flowers with brown keels and slightly curved fruits are reminiscent of some members of V. subgen. Tethyos, which often also have a penicillate callus. DNA data would be critical to confirm the phylogenetic relationships of these two taxa, which might represent an interesting case of convergent evolution.</p></div>	https://treatment.plazi.org/id/F87A879DFFE1FFA80194FBB2FA2194E1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Karremans, Adam P.;Pupulin, Franco;Damián-Parizaca, Alexander	Karremans, Adam P., Pupulin, Franco, Damián-Parizaca, Alexander (2025): A revised infrageneric classification of Vanilla (Orchidaceae) based on phylogenetic, morphological and biogeographical evidence. Phytotaxa 715 (3): 207-228, DOI: 10.11646/phytotaxa.715.3.2, URL: https://doi.org/10.11646/phytotaxa.715.3.2
F87A879DFFE4FFAE0194F90BFB219792.text	F87A879DFFE4FFAE0194F90BFB219792.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vanilla (Gondwana) Karremans	<div><p>2. Vanilla subgen. Gondwana Karremans, Damián &amp; Pupulin, subgen. nov.</p><p>Type species:— Vanilla cameroniana Damián (in Damián-Parizaca et al. 2023: 223).</p><p>Etymology: —The name derives from the landmass that existed in the Southern Hemisphere uniting South America and Africa, alluding to the unresolved phylogenetic position and intermediate morphological features of its only known member.</p><p>Diagnosis:—Nomadic vines, with subcoriaceous, bullate, spathulate, reticulate, petiolate leaves. Stem pustulate. Inflorescence racemose, short, with small non-foliaceous floral bracts. Flowers with greenish, narrow sepals and petals and a white lip, forming a cone-like, floral tube made by the adnation of the column and lip margins. Lip with a distinct penicillate callus above the middle and an elevated, trullate, crenulate to dactilar, apical disc. Column straight with a well-developed rostellum. Pollen in monads, forming two easily disrupted masses.</p><p>Discussion:—This monotypic section is currently represented only by the odd V. cameroniana, a species exclusively known by a few collections from French Guiana and which does not seem to have any currently known allies. The thin leaves with reticulate margins and greenish flowers with a white lip are reminiscent of members of Vanilla subgen. Membranacea, but the penicillate callus and tubular lip with fused margins place it among the members Vanilla subgen. Vanilla . However, unlike any member of either Vanilla subgen. Vanilla and most members of Vanilla subgen. Tethyos, the leaves of this species are large, bullate, spathulate, broadly-spathulate to cuneate, with a well-developed petiole, and the tubular lip forms a narrow cone with an apical bulge. The stem is pustulate, a feature that also occurs in Vanilla subgen. Tethyos, the sepals and especially the petals are extremely narrow, almost linear, a rather rare character found only in a few members of Vanilla subgen. Vanilla . Members of both sections have diverse arrangements of verrucae and papillae, but none have an elevated, trullate, crenulate to dactilar, apical disc as the one found in the sole member of V. subgen. Gondwana . Despite the absence of DNA data for the current analysis, V. cameroniana ’s distinctive morphology strongly supports its recognition as a separate lineage.</p><p>Ecology:—The only species in this subgenus is poorly known. No information on the fragrances produced by their flowers or fruits, or anything regarding pollination and dispersal of this species is known.</p></div>	https://treatment.plazi.org/id/F87A879DFFE4FFAE0194F90BFB219792	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Karremans, Adam P.;Pupulin, Franco;Damián-Parizaca, Alexander	Karremans, Adam P., Pupulin, Franco, Damián-Parizaca, Alexander (2025): A revised infrageneric classification of Vanilla (Orchidaceae) based on phylogenetic, morphological and biogeographical evidence. Phytotaxa 715 (3): 207-228, DOI: 10.11646/phytotaxa.715.3.2, URL: https://doi.org/10.11646/phytotaxa.715.3.2
F87A879DFFE5FFAE0194FD05FE81939D.text	F87A879DFFE5FFAE0194FD05FE81939D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vanilla (Membranacea) Karremans	<div><p>3. Vanilla subgen. Membranacea Karremans, Pupulin &amp; Damián, subgen. nov.</p><p>Type species:— Vanilla mexicana Miller (1768: without page number).</p><p>= Vanilla subsect. Membranaceae Portères (1951: 291), nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>= Vanilla subsect. Membranaceae Portères (1954: 159), nom. inval. nud. (Art. 39.1; Turland et al. 2018).</p><p>Diagnosis:—Nomadic vines, with one terrestrial species retaining a juvenile growth habit ( V. dietschiana). Stem smooth, scandent, branched, terete, non-sulcate, succulent. Leaves sub-petiolate to sessile, membranaceous, sub-reticulate, developing conspicuous blades, rarely reduced. Inflorescences axillary, racemose, often with leaf-like bracts, developing either on the proximal or distal part of the stem. Ovary non-calyculate. Flowers resupinate, long-lived, produced in succession, slightly fragrant. Sepals and petals green, often dark or coppery green. Lip white, typically with salmon, pink, green or yellow markings especially on the keels, calli, and warts. Sepals free, spreading, mostly contorted and undulate, smooth. Petals free, dorsally sulcate, spreading, mostly undulate-contorted. Lip sessile or with a very short claw, fused to the column only at its base, simple to trilobed, midlobe often quite distinct, disc often with two large longitudinal lamellae, thickened cushion-like calli, or with narrow to thickened-rugose longitudinal keels, lacking a penicillate callus. Column short, about half the length of the lip, arched, in a few species with a short to long foot. Pollen in monads, forming two easily disrupted masses. Fruit cylindrical, mostly straight, bean-like, dehiscent, green, turning black when mature, never yellow or reddish, with a sweet-grassy or putrid, non-vanillic fragrance.</p><p>Discussion:—This subgenus of 17 species previously included what was the type of genus, V. mexicana . With the genus now being conserved with V. planifolia as type species instead, the membranaceous Vanilla are no longer the typical subgenus and require a subgeneric name. We propose Vanilla subgen. Membranacea, using the name originally used for this group by Portères. It is here divided into two sections, the broadly distributed Vanilla sect. Membranacea (11 spp.) featuring species with predominantly distal inflorescences and allogamous reproduction, and Vanilla sect. Dictyophyllaria (6 spp.) which is primarily distributed in the Atlantic Forest and bears mostly proximal inflorescences and autogamous reproduction. Both sections have consistently received strong support in phylogenetic studies, including our own analysis (Fig. 1) and the recent comprehensive phylogenomic reconstruction by Damian et al. (2025b).</p></div>	https://treatment.plazi.org/id/F87A879DFFE5FFAE0194FD05FE81939D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Karremans, Adam P.;Pupulin, Franco;Damián-Parizaca, Alexander	Karremans, Adam P., Pupulin, Franco, Damián-Parizaca, Alexander (2025): A revised infrageneric classification of Vanilla (Orchidaceae) based on phylogenetic, morphological and biogeographical evidence. Phytotaxa 715 (3): 207-228, DOI: 10.11646/phytotaxa.715.3.2, URL: https://doi.org/10.11646/phytotaxa.715.3.2
F87A879DFFE6FFAC0194FA32FC8A9199.text	F87A879DFFE6FFAC0194FA32FC8A9199.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vanilla (Tethyos) Karremans	<div><p>4. Vanilla subgen. Tethyos Karremans, Damián &amp; Pupulin, subgen. nov.</p><p>Type species:— Vanilla africana Lindley (1862: 137) .</p><p>Etymology:— Tethyos is the genitive of Tethys, a word of Greek origin which means ancient sea or sea goddess. Tethys was the name of the prehistoric ocean located between the ancient continents of Gondwana and Laurasia, and bordered by the continents of Africa, Eurasia, India, and Australasia.</p><p>Diagnosis:—Nomadic or lithophytic vines, the leaves non-articulate, distichous, mostly with well-developed blades, but sometimes retaining deciduous scale-like leaves in several unrelated groups of species. Differing from the typical subgenus by the having flowers white or yellow, the lip typically notoriously stained with red, pink or purple (vs. lip similar in color to sepals and petals, not red, pink or purple), conical to saccate, the basal claw diversely fused to the column, typically forming a narrow, dry, papillose-hirsute, basal nectary, but not always forming an extended floral tube; the apical portion of the lip free, often flabellate, broadly expanding, with margins recurved. Sepals and petals often elliptic, broad (vs. lanceolate, narrow). Column shorter than the lip, with or without hairy ventral surface (vs. ventral surface hirsute), stigma a transverse cavity, the lateral lobes non-emergent (vs. emerging, flap-like). Pollen in monads, forming four easily disrupted masses. Fruits curved (vs. straight), non-calyculate (vs. calyculate), dehiscent (vs. either dehiscent or non-dehiscent), non-aromatic (vs. aromatic), specifically lacking vanillic and anisic compounds.</p><p>Discussion:— Vanilla subgen. Tethyos is the most diverse and broadly distributed subgenus of Vanilla . It currently includes 70 species across tropical Africa, Asia and the Antilles. They produce fleshy fruits, that are apparently dehiscent, and so-called non-aromatic. The fruits are actually not odorless, but none are known to contain compounds in the vanillyl, p-hydroxybenzyl and anisyl families, which give the fruits their characteristic vanilla-like smell and commercial interest.</p><p>As noted by Soto Arenas &amp; Cribb (2010), Vanilla sect. Aphyllae as defined by Rolfe is polyphyletic. However, all species assigned by Rolfe to that section were placed by Soto Arenas &amp; Cribb in their newly proposed V. sect. Tethya. This means that the lectotypification of V. sect. Aphyllae with any of its 12 species renders V. sect. Tethya illegitimate. Therefore, we selected the same type species as the two are unavoidably nomenclatural synonyms. Most species of Rolfe’s Vanilla sect. Aphyllae, typified by V. phalaenopsis, are endemic to the Indian Ocean basin, forming a distinct clade (Fig. 1) featuring leafless plants, showy white or yellowish flowers with maroon blotches on the lip, broad petals, and a loss of the penicillate callus. They seem to be closely related to a group of leafless New World species, which are endemic to the Antilles and differ in having narrow sepals and petals and a penicillate callus. These taxa, together with a few species from mainland Africa form our expanded circumscription of V. sect. Aphyllae, a section here including 22 species and being the only one in the genus that shows pantropical distribution. They are sister to a clade that is here given the new name V. sect. Miguelia, and which includes 37 exclusively Asian species, endemic to India, the Indo-China region and South-East Asia. Finally, both clades are sister to yet another Old World group, V. sect. Tethyos which includes 11 species exclusively known to occur in the tropics of Central and Western Africa.</p><p>Ecology:—Most members of Vanilla subgen. Tethyos seem to have sweetly fragrant flowers that appear to be dependent on pollinating bees to ensure reproduction (Watteyn et al. 2023), but observations on the pollination of flowers of species belonging to this subgenus are scanty. While the New World species are apparently pollinated by bees in the genus Centris (Centridini), their Old World relatives seem to be pollinated by bees in the genera Allodape (Allodapini) and Trinchostoma (Halictini).</p><p>Although morphologically similar to species of Vanilla subgen. Vanilla, the members of Vanilla subgen. Tethyos are neither pollinated, nor dispersed by the exclusively Neotropical Euglossine bees. Florally, many species belonging to both subgenera are similar in being mainly food-deceptive, having a dry, narrow nectary formed by the fusion of the claw of the lip and the base of the column. The column is elongate, and a penicillate callus in the middle of the lip blade is key in placing and removing the pollen masses as it forces the insect against the anther and stigma while it attempts to reach the basal nectary. The long-tongued Euglossini are needed in order to reach the nectary in Vanilla subgen. Vanilla, and it is therefore not surprising that some members of Vanilla subgen. Tethyos are pollinated by long-tongued counterparts in the Allodapini (Gigant et al. 2014; 2016).</p></div>	https://treatment.plazi.org/id/F87A879DFFE6FFAC0194FA32FC8A9199	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Karremans, Adam P.;Pupulin, Franco;Damián-Parizaca, Alexander	Karremans, Adam P., Pupulin, Franco, Damián-Parizaca, Alexander (2025): A revised infrageneric classification of Vanilla (Orchidaceae) based on phylogenetic, morphological and biogeographical evidence. Phytotaxa 715 (3): 207-228, DOI: 10.11646/phytotaxa.715.3.2, URL: https://doi.org/10.11646/phytotaxa.715.3.2
