Tillandsia recurvata

Rossado, Andrés J., Donadío, Sabina & Bonifacino, José M., 2024, A taxonomic revision of Tillandsia (Tillandsioideae, Bromeliaceae) from Uruguay, Phytotaxa 667 (1), pp. 1-90 : 1-90

11. Tillandsia recurvata (L.) Linnaeus (1762: 410). Figs. 16E , 32 , and 33.

≡ Diaphoranthema recurvata (L.) Beer ([1856] 1857: 156) ≡ Renealmia recurvata Linnaeus (1753: 287) . Type (lectotype designated here):—Without locality: s.d., s.leg. (Herb. Linn. 403.3 [LINN, online image!]).

Plants short caulescent, generally much branched from the base and forming subdense (sometimes not well-defined) semiglobose clumps, up to 16 cm in diameter. Roots to 0.6 mm diameter, only present at the base of the stem. Stems 0.8–5 cm long. Leaves distichously arranged, distributed along the stem, grayish to grayish-green; sheaths 5–22 × 3–5.5 mm, distinct from the blade, (narrowly) elliptic to elliptic-ovate, glabrous in the quarter or basal half, lepidote towards the apex; blades (15–)20–90(–130) × 0.7–1.7 mm, mostly spreading to divergent, linear, channelled, fleshy, densely tomentose-lepidote, trichomes asymmetric. Inflorescences simple; peduncles wholly or mostly exposed, (1.8–) 2.5–9 cm long, 0.4–0.7 mm in diameter, with trichomes all along; peduncle bracts 1 to 2 in number, erect, one always immediately below the basal floral bract, the other, when present, in the middle or apical part of the peduncle; upper peduncle bract with developed blade; sheath 3.6–11 × 2.2–3.4 mm, (narrowly) elliptic to elliptic-ovate, green to yellowish-green and somewhat grayish, subdensely to densely lepidote; blade 0.5–9(–16) mm long. Spike distichously and densely 1 to 2-flowered, 6–12 mm long, slightly complanate, narrowly elliptic in outline; rachis totally hidden by the floral bracts and flowers, green, glabrous. Floral bracts 3.3–8 × 2.3–4.3 mm, shorter than the sepals, densely distributed (about 1.5–3 times longer than the internodes), imbricate, clasping the basal half of the flower, bladeless or sometimes with a short blade (not longer than 2 mm), widely elliptic to widely elliptic-ovate, ecarinate, green to yellowish-green (sometimes with vinaceous tinges) and somewhat grayish, subdensely to densely lepidote. Flowers 7–10 mm long, scentless; sepals visible, (3.5–)4–6.5 × 1.8–2.8 mm, evenly short connate at the base for 0.4–0.8 mm, elliptic-obovate to oblong-obovate, or elliptic, ecarinate, vinaceous, sometimes somewhat reddish, glabrous or with scattered trichomes; petals 6–8.5 × 0.7–1.5 mm, lingulate; distal portion white with little fuzzy lilac spots, spreading, sometimes divergent, margins entire; stamens 3.2–4.2 mm long, deeply included, a little longer than the pistil; filaments 2.7–3.3 mm long, straight (not plicate); pollen yellow; pistil 2.5–3.3 mm long, deeply included; ovary 1.6–2.3 × ca. 1 mm, ellipsoid to cylindrical-ellipsoid, tapering into the style; style 0.7–0.8 mm long, much shorter than the ovary, whitish; stigmas simple-truncate. Capsules 15–30 × 1.1–1.8 mm, much exceeding its respective floral bract, cylindrical-prismatic, apex rounded (sometimes somewhat obtuse) and short-beaked.

Vernacular names:— Clavel del aire, clavelito ( Brito & Llano 2008).

Classification:— Tillandsia recurvata is the type species of T. subg. Diaphoranthema ( Smith & Downs 1977) .

Distribution and habitat:— Tillandsia recurvata is one of the most widely distributed species of Bromeliaceae , occurring from the southern United States to central-western Argentina (absent in the southernmost parts of Central America, Chile and in the Amazonian region). In Uruguay, it is the second most frequent species of Bromeliaceae , found throughout the country (with the exception of southern and eastern Rocha department; Fig. 33 ). It grows as epiphyte in all types of forests but it is usually more common in open or sparse forests (e.g. hydrophilous forest, open thorn forest, xeric hillside forest), edges of dense forests (forest-grassland ecotone), and isolated trees in grasslands. In addition, this species occasionally grows as epilithic on rocks associated with hillside forests. Tillandsia recurvata is well adapted to modified habitats; thus, it is very common to find it in disturbed forest sites, as well as in urban and suburban areas, growing on street trees, power lines, window bars, and even on walls.

Historically, the southern limits of Tillandsia recurvata distribution laid in southern Uruguay and central-eastern Argentina (northern Buenos Aires province). However, Apodaca & Guerrero (2019) reported the expansion of this species by 200 km towards the south. This recent expansion is probably attributed to climate change and lignification of the Pampas ( Apodaca & Guerrero 2019).

Phenology:— Flowering in early summer. The flowering period is short and very constant among years, always occurring from mid-December to mid-January ( Fig. 4 ). Fruiting takes place as of January. The fruits grow in length mainly through January and February and then remain without growth for several months. Tillandsia recurvata has a long period for seed dispersal that is also variable among years, depending on the climatic conditions of the year and the intrinsic characteristics of each plant. Generally the opening of the capsules occurs from July to October (winter and early spring), thus the plants are with fruits for six to 10 months. The development of the inflorescence takes approximately 30 to 40 days; thus, in each plant a period of eight to 12 months occurs between the inflorescence emergence and the opening of the capsules.

Reproductive observations:— Tillandsia recurvata is a self-compatible species that exhibits populations with cleistogamous and chasmogamous flowers ( Donadío 2013; Bianchi & Vesprini 2014). In Uruguay, all the plants observed have always chasmogamous flowers.

Conservation:— We consider Tillandsia recurvata as Not Threatened in Uruguay.The species is widely distributed and frequent across the country, inhabiting a wide range of habitats, including disturbed and urban areas.

Diagnostic characters:— Tillandsia recurvata can be distinguished from the rest of the species of Tillandsia occurring in Uruguay by the combination of distichous and imbricate leaves, and the presence of a peduncle bract immediately below the basal floral bract. Nevertheless, due to its general resemblance to T. bandensis subsp. bandensis , T. capillaris and T. virescens , T. recurvata is commonly confused with them, both in herbarium collections and in the field.

Background in Uruguay:— Tillandsia recurvata was first cited for the Uruguayan flora by André in 1905, without specifying reference material. Subsequently, several works ( Herter 1930, 1949 –1956; Smith 1933) mentioned this species for the country but it was Smith, in 1972, the first to cite a herbarium specimen from Uruguay.

Note: — Renealmia recurvata was published by Linnaeus in Species Plantarum (1753), who cited a synonym from Sloane (1707) under this name. Based on this information and in the fact that according to Smith (1935) Linnaeus had no material of his own when publishing this name, traditional taxonomic works on Bromeliaceae (e.g. Smith 1935; Smith & Downs 1977; and later works based on these) have consistently cited the collection of Sloane s.n. (BM) as the holotype (or just type) for Renealmia recurvata . Here, we discard the specimens of Sloane s.n. (BM000589694, BM000589695) as original materials for this species since, as stated by Turland & Jarvis (1997), Linnaeus never studied the collections cited in Sloane (1707; but he did study its illustrations). Therefore, in accordance with ICN (Arts. 9.1, 9.3 and 9.4; Turland et al. 2018), these cannot be taken as original material. In this sense, we found two elements that represent original material for R. recurvata . One corresponds to the illustration on tabula 121, figure 1 of Sloane (1707), as it is explicitly stated by Linnaeus (1753) “Sloan. jam. 77. hift. I. p. 190. t. 120[121] 1. f. 1”. The other original material is a sheet in Linnaean Herbarium with “ Renealmia recurvata ” and the number “403.3” written on it, and with material that matches the morphological information given in Linnaeus (1753) and the illustration in Sloane (1707). According to Turland & Jarvis (1997), the presence of a Species plantarum species number is a strong indication that Linnaeus had the material in his possession at the time he wrote the Species plantarum. Therefore, and in accordance with ICN Art. 9.12 ( Turland et al. 2018) we choose the specimen Herb. Linn. 403.3 as the lectotype of Renealmia recurvata .

It is worth mentioning that the illustration of Renealmia recurvata in Sloane (1707, original material) is undoubtedly based on the plant mounted on the upper part of the sheet of Sloane s.n. (BM000589695), and could be considered as a typotype.

Representative specimens examined:— URUGUAY. Artigas: Cuareim , 17 August 1901, Berro 1207 ( MVFA) . Canelones: Arroyo Solis Grande y ruta 9 ( Parque del MTOP, en isla del arroyo al sur del puente), 3 February 2013, Haretche 521 ( MVFA) . Cerro Largo: Aceguá (de Ruta 8 a Camino Internacional ), 12 December 1986, Armand-Ugón et al. s.n. ( MVFA 18887 ) . Colonia: Arroyo Tembetarí, cerca Playa Fomento , 29 January 1964, Marchesi 813 ( MVFA) . Durazno: Lago Rincón del Bonete, costa del Río Negro, entre brazo del Sauce y Brazo del Sarandí - Establecimiento Rincón del río, 12 March 2014, Haretche 725 ( MVFA) . Flores: Parque Centenario , 30 August 2013, Rossado 179 ( MVFA) . Florida: Estancia Los Cerros - monte sobre Santa Lucía Chico , 5 December 1971, Izaguirre s.n. ( MVFA 11031 ) . Lavalleja: Arroyo del Soldado y ruta 108, 22 March 1989, Ren s.n. ( MVFA 19574 ) . Maldonado: Sierra de las Ánimas , 16 November 1971, Izaguirre s.n. ( MVFA 10782 ) . Montevideo: Barrio Parque Batlle , 24 December 2014, Rossado 415 ( MVFA) . Paysandú: Cerros de Cambará ( Estancia “ El Cambará ”), 17 February 2013, Haretche 534 ( MVFA) . Rio Negro: Río Negro y Arroyo Yapeyú, campo Jabalí , 25 October 1993, Marchesi & Armand-Ugón s.n. ( MVFA 22444 ) . Rivera: Cuñapirú , May 1900, Berro 3162 ( MVFA) . Rocha: Entre el Río Cebollatí y la Laguna Merín - Frente a la Isla del Padre , 4 February 2019, Rossado et al. 566 ( MVFA) . Salto: Rambla de Salto, Arenitas Blancas , 28 March 2003, Brussa 1986 ( MVJB) . San José: Arazatí , April 1922, Hauman s.n. ( BA 22282 ) . Soriano: Playa de la Agraciada, 19 November 2012, Rossado et al. 42 ( MVFA) . Tacuarembó: Gruta de los Cuervos , 9 March 1966, Rosengurtt et al. s.n. ( MVFA 10047 ) . Treinta y Tres: Quebrada de los Cuervos , 16–18 December 1964, Brescia & Marchesi s.n. ( MVFA 3763 ) .

Figures

FIGURE 4. Phenology of Tillandsia species occurring in Uruguay. Divisions within each month indicate intervals of approximately 10 days, identified as the beginning, middle, and end of each month. Tillandsia loliacea is excluded due to lack of enough information. * Denotes species with cleistogamous flowers. ** Denotes species with insufficient data to ascertain the timing of fruiting or fruit dehiscence.

FIGURE 16. Illustrations of floral parts. A. Tillandsia capillaris. B. T. loliacea. C. T. virescens. D. T. tricholepis. E. T. recurvata. F. T. usneoides. From left to right: floral bract (stretched view), sepals (stretched view; adaxial sepals to the right), petal (stretched view), and pistil.A based on Rossado 404 (MVFA); B based on Villagrán et al. s.n. (MVJB 30693); C based on Rossado 362 (MVFA); D based on Rossado 363 (MVFA); E based on Rossado 415 (MVFA); F based on Rossado et al. 406 (MVFA).

FIGURE 32. Tillandsia recurvata. A. Habitat; image shows an individual of Sebastiania commersoniana densely covered with T. recurvata (Sierra de Mahoma, San José, Uruguay). B. Habit (San Miguel stream, Paysandú, Uruguay). C. Close up of an individual in bloom. D. Fruit-bearing plant (Botanical Garden of Montevideo, Uruguay). E. Inflorescence. F. Front view of the flower. G. Lateral view of the flower. Notice the floral bract and the upper peduncular bract clasping the basal part of the flower. H. Infructescence. Photos by Andrés Rossado.

FIGURE 33. Distribution map of Tillandsia recurvata in Uruguay and neighboring areas.