Vanilla chamissonis Klotzsch

Vanilla chamissonis Klotzsch View in CoL ( Klotzsch 1846: 564)

Figs 3 View Figure 3 , 4 View Figure 4 , 6 View Figure 6 , 5 View Figure 5 , 7 View Figure 7 , Table 1 View Table 1

Vanilla chamissonis var. brevifolia Cogn. ( Cogniaux 1893: 149). Based on Epidendrum vanilla Vell. ( Vellozo 1827: ic. 9, t. 1), nom. illeg. View in CoL

Vanilla gardneri Rolfe ( Rolfe 1895: 177), syn. nov. – Type: BRAZIL • Rio de Janeiro, Morro do Flamengo, near Rio de Janeiro; G. Gardner 245; lectotype (designated by Soto Arenas and Dressler 2010): K! [ K 000463752, K 000463753]. View in CoL

Vanilla vellozoi Rolfe ( Rolfe 1896: 467), as “ Vanilla vellozii ”, syn. nov. – Type: BRAZIL • Rio de Janeiro; 1882; A. Glaziou 14302; lectotype designated by Soto Arenas and Dressler 2010): K! [ K 000940259]. View in CoL

Vanilla carinata Rolfe ( Rolfe 1896: 468), syn. nov. – Type: BRAZIL • Rio de Janeiro, Organ Mountains; 1878; J. Miers s. n.; holotype: K! [ K 000463750]. View in CoL

Type

BRAZIL • Santa Catarina; A. von Chamisso s. n.; lectotype (designated by Christenson 1995: 33): LE! [ LE 00011144 ] .

Description

Nomadic vines, long scandent. Roots axillary, one per node; terrestrial roots up to 8 mm diam., fleshy, whitish, with hyaline absorbing hairs; aerial roots 2.2–3.2 mm diam., whitish to brownish. Stem climbing, cylindrical, fleshy, straight to sinuous, glabrous, glaucous to dark green, strongly furrowed under arid conditions; internodes of ascendant stems 8–14 × 0.7–1.3 cm. Leaves 5.2–18 × 2.5–5.5 cm, alternate, distichous, elliptic to oblong, asymmetric, fleshy, glabrous, pale green to dark green, pseudopetiolate, margin entire, base attenuate, apex acuminate; pseudopetiole 5–10 mm concave. Inflorescence axillary, racemose, with up to 22 flowers opening in succession; 1–2 flowers opening each morning; rachis 5–9 × 0.9–1.3 cm, terete, pale green to dark green; bracts 4–7.5 × 4.5–8 mm, progressively smaller toward the apex, triangular / deltoid, coriaceous, concave, green, patent, apex acute, not incurved. Flowers resupinate, pedicellate, abscission layer between perianth and ovary present; pedicel with ovary 46–50 × 4.2–6 mm, triangular in transverse section, straight to incurved, whitish at the base, green to the apex, with a calyculus (6–6.5 × 3–3.5 mm) at the apex. Sepals 4.9–6.1 × 0.9–1.3 cm, free, oblanceolate, fleshy, slightly concave, spreading, pale green to yellowish, margin entire, involute at the base, base attenuate, apex acute to obtuse; dorsal sepal symmetric; lateral sepals asymmetric. Petals 4.9–6.1 × 0.9–1.2 cm, free, obliquely oblong-elliptic, asymmetric, lower margin more arcuate, yellowish at the base, pale green to the apex, base attenuate, apex obtuse to rounded, adaxial surface with a central and longitudinally disposed keel. Labellum 1 - lobed to slightly 3 - lobed, 5.3–6.2 × 3.3–4.2 cm, tubular, deepening near the middle, yellowish to the base, white in distal portion, unguiculate, with a central crest from the unguiculus to the apex, and a penicillate callus just below the anther; unguiculus fused along the margins of the basal half (ca 29–32 mm) of column length forming a nectar chamber, nectar chamber 1.4–1.6 cm long, tubular; central crest yellowish from the nectar chamber to the penicillate callus, dark yellow to the apex; distal portion of the central crest swollen, low cushion, rugose-papillose at the apex, with a group of transversal yellow-orange scales near the penicillate callus, with three yellow longitudinal lines near the apex; penicillate callus 5.1–5.5 × 3.8–4.2 mm, made by yellow-hyaline lacerate-laciniate scales and clusters of trichomes; lateral lobes not much evident, rounded, overlapping the column apex, margin undulate; midlobe deeply emarginated; margin undulate. Column 36–38 × 3.2–3.5 cm, trigonous, arched to the base, forming an angle ca 90 ° with the ovary, strait to the apex, ventral surface flat with white to yellowish hyaline trichomes over the distal half, attenuate to the base, dilated to the apex, with two lateral wings; lateral wings rounded, undulate. Stigma bilobed; rostellum 4–4.2 × 2.5–2.7 mm, trapezoidal, membranous, white. Anther 4.8–5.2 × 3.5–3.6 mm, rectangular to trapezoidal, white, versatile, apex truncate; pollen mass 2.9–3.2 × 2.9–3.1 mm, triangular, bipartite, whitish. Fruits 12–17 × 2.6–3.5 cm, oblong to clavate, arched, transversally subtrigonous, fleshy, brown indehiscent when mature, fragrant; pericarp soft; fruit cavity filled. Seeds ca 0.5 mm, ovoid, black.

Distribution and ecology

Vanilla chamissonis is endemic to the Brazilian Atlantic Forest (Fig. 4 View Figure 4 ). The species is a nomadic vine found both on tall trees in the restinga vegetation and on shrubs of the rocky outcrops close to the beach. The flowers are very fragrant and produce a small amount of nectar. The fruits have a bitter taste and an unpleasant odour. After 18 months from pollination, they turn brown and fall to the forest floor where they are consumed by agoutis.

Phenology

Vanilla chamissonis blooms from November to January. The fruits ripen from April to June, 18 months from pollination.

Preliminary IUCN conservation assessment

Endangered: EN B 2 ab (i, ii, iii). Vanilla chamissonis is distributed along the Brazilian coast. The species is particularly common in the Atlantic Rainforest of south and southeast Brazil. The populations are commonly composed of many individuals. The extent of occurrence (EOO) is estimated to be 229,334 km 2, which falls within the limits for Least Concern (LC) under criterion B 1, according to the IUCN Red List categories and criteria. The area of occupancy (AOO) is estimated to be 128 km 2, which falls within the limits for Endangered (EN) under criterion B 2. Considering that the Brazilian restingas have been reduced to scattered fragments due to the urban occupation, I project a continuing decline in (i) extent of occurrence, (ii) area of occupancy, and (iii) extent and / or quality of habitat of V. chamissonis . Based on these threats and the fact that the species is distributed in less than five locations, V. chamissonis is assessed as Endangered: EN B 2 ab (i, ii, iii).

Additional material examined

BRAZIL – Espírito Santo • Espírito Santo, Afonso Cláudio, Serra Pelada, comunidade de Palmital, propriedade da família Brandemburg, Pedra do Sol, Proximidades da Pedra da Lajinha. Inselberg ; 20°00’36”S, 41°04’30”W; 760 m; 21 Oct. 2019; C. N. Fraga & D. R. Couto 3912; MBML GoogleMaps • Guarapari, Village do Sol ; 20°33’16”S, 40°24’35”W; 27 Oct. 1984; B. Weinberg 645; MBML GoogleMaps • Guarapari, Parque Estadual Paulo César Vinha ; 20°36’19”S, 40°25’19”W; 17 Nov. 1994; C. N. de Fraga et al. 59; MBML GoogleMaps • Jaguaré ; 18°54’20”S, 40°04’33”W; 78 m; 30 Nov. 2012; D. A. Folli 6934; CVRD GoogleMaps • Santa Leopoldina, Rio das Farinhas, propriedade da Sra. Maria Knak Ule. Mata, dossel ; 20°01’21”S, 40°36’29”W; 827 m; 15 Apr. 2008; A. P. Fontana, L. Kollmann & K. A. Brahim 4969; MBML GoogleMaps • Vila Velha, Jacarenema ; 20°19’46”S, 40°17’32”W; 25 May 1990; J. M. L. Gomes & O. J. Pereira 1129; VIES GoogleMaps . – Paraná • Matinhos, ao pé do Morro do Tabaquara, perto de Matinhos ; 25°49’02”S, 48°32’34”W; 3 Jan. 1967; J. C. Lindeman et al. 3855; MBM GoogleMaps • Paranaguá, Ilha de Mel ; 25°31’13”S, 48°30’33”W; 3–5 m; 28 Nov. 1970; G. G. Hatschbach 25679; NY GoogleMaps • Paranaguá ; 25°31’11”S, 48°30’33”W; 3–5 m; 23 Nov. 1994; J. Cordeiro 1201; HUEFS GoogleMaps • Paranaguá, Ilha do Mel - Restinga da Praia Grande ; 25°31’11”S, 48°30’33”W; 8 Nov. 1986; W. S. Souza 388; UPCB GoogleMaps . – Rio de Janeiro • Angra dos Reis, Ilha Grande, Parcelas do Módulo Oeste do RAPELD Ilha Grande em Mata de restinga, Reserva Biológica Estadual da Praia do Sul ; 23°10’36”S, 44°17’52”W; 13 Oct. 2016; A. C. R. Cruz & R. G. Diniz 2; RBR GoogleMaps • Campos dos Goytacazes, Morro Itaoca - Morro do rato ; 21°47’49”S, 41°26’52”W, 201 m; 23 Nov. 2008; A. S. Pessanha & M. T. Nascimento 48; HUENF GoogleMaps • Maricá, Barra de Maricá, Área C 1 ; 22°55’09”S, 42°49’06”W; 25 Oct. 1988; Occhioni et al. 9286; RFA GoogleMaps • Macaé ; 22°22’14”S, 41°47’12”W; 18 Sep. 2008; I. E. Santo & M. F. Castilhori 174; R GoogleMaps • Rio de Janeiro, Restinga da Marambaia ; 23°03’11”S, 43°35’03”W; 20 May 2009; B. S. Haiad, C. Novaes & I. Soares s. n.; HUNI [ HUNI 4402 ] GoogleMaps . – Santa Catarina • Balneário Camboriú, Taquarinhas ; 26°59’26”S, 48°38’04”W; 17 Nov. 2002; C. Hering-Rinnert 244; JOI GoogleMaps • Bombinhas, Praia de Zimbros ; 27°08’16”S, 48°31’01”W; 35 m; 11 Nov. 2006; M. G. Caxambú 1270; HCF GoogleMaps • Garopaba, Ouvidor - Praia Vermelha ; 28°06’42”S, 48°38’02”W; 23 m; 7 Jul. 2018; A. Kassner-Filho 3056; FURB GoogleMaps • Florianópolis, Morro do Ribeirão, Ilha de Santa Catarina ; 27°35’48”S, 48°32’57”W; 100 m; 23 Sep. 1970; R. M. Klein & A. Bresolin 8773; FLOR GoogleMaps • Florianópolis, Lagoa Pequena, Rio Tavares ; 27°39’20”S, 48°28’18”W; 11 m; 2 Nov. 2017; G. D. S. Seger & E. Bach 843; ICN GoogleMaps • Imaruí, Forquilinha ; 28°09’52”S, 48°52’10”W; 666 m; 28 Jan. 2010; J. L. Schmitt et al. 1111; FURB GoogleMaps • Laguna, Praia do Sol, Dunas fixas ; 28°28’57”S, 48°46’50”W; 3–5 m; 13 Dec. 2000; G. Hatschbach, A. C. Cervi & E. Barbosa 71869; BHCB GoogleMaps • Paulo Lopes ; 28°01’21”S, 48°42’20”W; 123 m; 18 Nov. 2022; M. E. Engels 10283; UPCB GoogleMaps • Penha, J. B. World Entretenimentos S/A ; 26°48’06”S, 48°36’58”W; 7 m; 26 Dec. 2018; A. Kassner-Filho & F. L. V. Bones 4490; FURB GoogleMaps • São Francisco do Sul, Parque Estadual do Acaraí - Praia do Ervino ; 26°20’58”S, 48°33’47”W; 5 m; 29 Nov. 2010; A. Korte & A. L. de Gasper 5252; FURB GoogleMaps • São Francisco do Sul ; 26°14’35”S, 48°38’17”W; 5 m; 5 Apr. 2008; L. Sevegnani s. n.; FURB [ FURB 7432 About FURB ] GoogleMaps . – São Paulo • Cananéia, Parque Estadual da Ilha do Cardoso, Trilha morro das almas ; 25°00’36”S, 47°55’11”W; 19 Mar. 2003; T. B. Breier 953; UEC GoogleMaps • Cananéia, Ilha do Cardoso, Restinga do Pereirinha ; 25°00’36”S, 47°55’11”W; 2 Dec. 1990; F. Barros & P. Martuscelli 1989; SP GoogleMaps • Iguape, Estação Ecológica Juréia-Itatins, Estrada entre o rio Una do Prelado e o Rio Verde, 18 km do rio Una, restinga junto à desembocadura do rio Verde ; 24°42’00”S, 47°32’59”W; 9 Dec. 1995; I. Cordeiro et al. 1595; SP GoogleMaps • Peruíbe, Estação Ecológica Juréia-Itatins, Barra do Uma ; 24°19’12”S, 46°59’24”W; 20 Nov. 1990; E. L. M. Catharino, et al. 1503; SP GoogleMaps • Praia Grande ; 24°00’00”S, 46°24’00”W; 14 Nov. 1929; A. Gehrt s. n.; SP 24495 GoogleMaps • Praia Grande ; 24°00’00”S, 46°24’00”W; 14 Nov. 1929; A. Gehrt 24495; NY GoogleMaps • Ubatuba, Picinguaba ; 23°25’47”S, 45°04’12”W; 6 Nov. 1988; A. Furlan 598; HRCB GoogleMaps • Ubatuba, Picinguaba ; 23°25’47”S, 45°04’12”W; 28 Dec. 2024; E. R. Pansarin 1579; LBMBP GoogleMaps • Ubatuba, Estação Experimental ; 23°25’47”S, 45°04’12”W; 23 Nov. 1940; J. F. Cunha s. n.; SP [ SP 44817 ] GoogleMaps .

Notes

Differences between specimens identified as Vanilla chamissonis from coastal populations and from the interior of Brazil have been recognized. The plants that occur inland are commonly referred to as “ V. chamissonis mineira ”, in reference to their occurrence in Minas Gerais, a southeastern Brazilian state inserted in the Cerrado. Morphological differences among inland and coastal plants were formally recognized twice: Hoehne (1945) published the invalid name V. chamissonis var. longifolia based on a plant collected in the municipality of Itú, southeastern Brazil. Itú has areas typical of an open Cerrado vegetation with many rocky outcrops, while Hicken (1917) described V. argentina based on a plant collected in the Formosa region, Argentina. The region of Formosa, besides Corrientes and Misiones, is characterized as Arid Chaco or Dry Chaco. The Dry Chaco vegetation consists of a mosaic formed by xerophytic forests, gallery forests, and savannas. Here, V. chamissonis , which occurs in the Atlantic Forest, is considered as a distinct species from V. argentina , which is distributed in the Brazilian Cerrado and Dry Chaco. Some species described based on plant material collected in the state of the Rio de Janeiro, at the Brazilian coast, clearly are synonyms of V. chamissonis . This is the case for Vanilla vellozoi (as V. vellozii ). Although some authors consider V. vellozoi as an obscure taxon related to V. calyculata , both plant material collected in the Rio de Janeiro (hosted at K), i. e. A. Glaziou 11621 and A. Glaziou 14302 (the latter designated by Soto Arenas and Cribb (2010) as lectotype of V. vellozoi , as V. vellozii ), strongly agree with V. chamissonis . Apart from the small size of the flower structures, the shape of the leaves (symmetrically elliptic) and the short and robust inflorescence with patent triangular / rhomboidal bracts leave no doubt that both plant material are V. chamissonis . Soto Arenas and Cribb (2010) consider V. argentina to be synonymous with V. vellozoi . However, as presented here, V. vellozoi is related to V. chamissonis , not to V. argentina . The second taxon is V. carinata . The holotype of V. carinata has been considered as inadequate and conspecific with V. planifolia ( Hoehne 1945) . However, the material J. Miers s. n. ( K 000463750 ) was collected on the Atlantic Coast, in the state of Rio de Janeiro, where V. planifolia does not occur. Although I agree that the holotype is not in good condition, since it lacks vegetative structures, analysis of the reproductive characteristics, such as the presence of a calyculus, the robust inflorescence rachis with patent triangular / rhomboidal bracts, and the presence of an evident labellar keel leaves no doubt that V. carinata is conspecific with V. chamissonis . A third obscure taxon is V. gardneri . Vanilla gardneri was described based on several materials belonging to at least three Vanilla species ( Soto Arenas and Cribb 2010). However, the material G. Gardner 245 ( K), collected at Morro do Flamengo, Rio de Janeiro, has been designated as the lectotype of V. gardneri ( Soto Arenas and Cribb 2010) . This exsiccate appears to contain a mixture of plant elements from two distinct species. While the vegetative portion and the flowers appear to be V. phaeantha , the fruit longitudinally sectioned strongly agree with those of V. chamissonis . This is corroborated by the fact that both species are sympatric in this region. Furthermore, the fruiting period of V. phaeantha , whose fruits ripen in nine months, does not overlap with its flowering period. On the other hand, the fruiting period of V. chamissonis , whose fruits take 18 months to mature, overlaps with the flowering period of both sympatric species, V. phaeantha and V. chamissonis . Therefore, based on the presence of a fruit of the latter species in the material G. Gardner 245 ( K), V. gardneri was synonymized here under V. chamissonis .

Morphological affinities

While V. calyculata and V. argentina occur in dryer environments, V. chamissonis is distributed throughout the Atlantic Forest sensu stricto (Fig. 4 View Figure 4 ). Vanilla chamissonis is easily recognized by its elliptic to oblong and asymmetric leaf blades (Fig. 5 View Figure 5 ), sepals 4.9–6.1 cm long, oblong-elliptic petals 4.9–6.1 cm long, labellum 5.3–6.2 cm long with a nectar chamber 1.4–1.6 cm long, column 36–38 mm long, and fruits whose pericarp is soft (Figs 6 View Figure 6 – 7 View Figure 7 ). Vegetative and floral features of V. chamissonis suggest a close relationship with V. calyculata and V. argentina . All three species show robust climbing stems with long fleshy ascendant leaves and lateral inflorescences with whitish resupinate flowers. Flowers of V. chamissonis and related taxa also share several characteristics, such as an unguiculate labellum with an emarginated apical lobe, and a yellow central crest with a penicillate callus just below the anther. In addition, the inner surface of the labellum base of V. chamissonis and related taxa is yellow, while the distal portion is white. In particular, fruit features suggest a close relationship between V. argentina and V. chamissonis . Both species produce large brown and indehiscent fruits that take 18 months to mature. Fruits have an unpleasant fragrance and a bitter flavour. The close relationships among V. argentina , V. chamissonis , and V. calyculata is corroborated by molecular data (see further).