Tillandsia capillaris Ruiz & Pavón (1802: 42)

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

4. Tillandsia capillaris Ruiz & Pavón (1802: 42) . Figs. 16A , 17 , and 18.

≡ Diaphoranthema capillaris (Ruiz & Pav.) Beer ([1856] 1857: 153). Type (lectotype designated here):— PERU. Huánuco: without exact locality, 1778–1788, Ruiz & Pavón s.n. ( BM000923923 [online image!]; syntypes: B–W-06336-010 [online image!], BM000923922 [online image!], BM000923925 [online image!], F0361001F [online image!], MA810425 [online image!], MA810426 [online image!], MA810427 [online image!], MA810428 [online image!], P00753185 [online image!]) .

= Tillandsia capillaris Ruiz & Pavón (1802: 42) f. hieronymi (Mez) L.B. Smith (1935: 213) ≡ Tillandsia hieronymi Mez (1896: 876) . Type (lectotype first-step designated by Smith 1935: 213 [as “type”]):— ARGENTINA. Córdoba: without exact locality, 12 November 1876, Hieronymus 124 (B [one sheet with admixture] second-step lectotype designated here: B 10 0243353-a [digital image!]; isolectotypes: BM000923929 [online image!], CORD00002229 [digital image!], F0045282F [digital image!], G00098116 [digital image!], G00098147 [digital image!], GOET000429 [digital image!], M0111401 [digital image!], US 00089212 [digital image!], US 01101048 [digital image!]).

Plants caulescent, much branched from the base and forming very dense globose or semiglobose clumps, up to 11 cm in diameter. Roots less than 0.5 mm diameter, only present at the base of the stem. Stems up to 8 cm long. Leaves distichously arranged, distributed along the stem, grayish-green; sheaths 5.5–10 × 2.5–4 mm, distinct from the blade, elliptic, subdensely lepidote except for the glabrous base; blades 8–17 × 0.7–1.1 mm, mostly spreading, linear, not channelled, fleshy, densely lepidote, trichomes symmetric or generally slightly asymmetric. Inflorescences 1-flowered (seldom 2-flowered); peduncles bractless (sometimes bearing a foliaceous basal bract), (0.9–) 1.3–3.3 cm long, glabrous. Floral bracts with a short blade or sometimes bladeless, ecarinate; sheath 3.5–5.5 × 2.8–4 mm, shorter than the sepals, clasping the base of the flower, widely elliptic to widely elliptic-ovate, stramineous to greenish, glabrous; blade to 2(–3) mm long. Flowers 6–8.5 mm long, fragrant; sepals visible, 4–6 × 1.3–1.8 mm, unequally connate, narrowly elliptic to narrowly oblong, ecarinate, vinaceous to reddish all along or with the basal portion green and the apical portion yellowish, glabrous; abaxial sepal connate with the adaxial sepals for 0.6–1 mm; adaxial sepals connate to each other for 1.6–1.9 mm (less than ½ of their length); petals 7.2–8.2 × 0.9–1.3 mm, lingulate; distal portion yellow with vinaceous spots barely perceptible, spreading, margins entire; stamens 3.2–3.6 mm long, deeply included, longer than the pistil; filaments 2.2–2.5 mm long, straight (not plicate); pollen deep yellow; pistil 1.8–2.3 mm long, deeply included; ovary 1.1–1.4 × ca. 0.9 mm, subprismatic, abruptly contracted into the style; style 0.7–0.8 mm long, a little shorter than the ovary, whitish; stigmas simple-truncate. Capsules 12–22 × 0.7–1 mm, much exceeding its respective floral bract, cylindrical-prismatic, apex truncate or obtuse and short-beaked.

Vernacular names:— Not known. Possibly clavelito.

Classification:— Tillandsia capillaris belongs to T. subg. Diaphoranthema , based on its morphological characteristics ( Smith & Downs 1977), and the morphological and molecular phylogenies obtained by Donadío et al. (2015), Granados Mendoza et al. (2017), and Donadío et al. (2022).

Distribution and habitat: — Tillandsia capillaris s.s. is native to Southern Ecuador, Peru, Bolivia, Chile, and Argentina, growing mainly in arid environments associated to the Andean Mountains ( Castello et al. 2020). This species is exotic to Uruguay ( Rossado et al. 2018b), where it is found as epiphyte in Montevideo city (Montevideo department), growing subspontaneously in urban parks, street trees, gates, and power lines. Additionally, one specimen was found from Colonia department ( Fig. 18 ). In Montevideo, T. capillaris cohabits with T. virescens , a morphologically similar and closely related species, although the latter is more widely distributed throughout the city. In certain areas of Montevideo, T. capillaris is very abundant, surpassing T. virescens and/or T. recurvata in terms of number of individuals.

Phenology: —Flowering in November and December, fruiting in December ( Fig. 4 ). Seed dispersal mainly occurs in October and November of the following year. Therefore, sometimes the opening of the capsules generated the previous year coincides with the flowering period of the current year. The development of the inflorescences takes approximately 30 days, and the whole reproductive cycle (from the onset of the inflorescence emergence to the dehiscence of capsules) lasts about one year.

Reproductive observations: —According to Till (1989a), and Bianchi & Vesprini (2014), Tillandsia capillaris s.s. is a self-compatible species, exhibiting populations or individuals with both cleistogamous and chasmogamous flowers. In Uruguay, all observed plants always have chasmogamous flowers, emitting a sweet fragrance, especially noticeable at dusk.

Conservation: —Not applicable. Tillandsia capillaris is an exotic species to Uruguay.

Diagnostic characters:— The unique combination of characters that distinguishs this species from others within Uruguay are: distichous leaves, unequally connate sepals (adaxial sepals more connate between them than with the adaxial one), and glabrous floral bracts. Tillandsia capillaris is easily confused with T. virescens , as they share a similar gestalt. The main differences between these species are shown in the key.

Background in Uruguay:— Tillandsia capillaris s.s. was recently reported for the first time in Uruguay by Rossado et al. (2018b).

Note I: —Several specimens collected and identified by Ruiz & Pavón as Tillandsia capillaris exist. We designate BM000923923 as the lectotype for this species, as besides having both vegetative and reproductive structures and being a well-preserved specimen, it has the most complete label information of all. This designation is in agreement with W. Till (who has worked extensively on T. capillaris and related species) who in his PhD thesis (Till 1984) cited BM000923923 as the lectotype for T. capillaris . Further, in 1982, Till attached a label to this specimen (BM000923923) identifying it as a lectotype. Unfortunately, these lectotypifications attempts were not effectively published (ICN Arts. 30.1 and 30.9, Turland et al. 2018), and therefore, and in accordance with the ICN Art. 7.10 ( Turland et al. 2018), the designation is done here.

It is worth mentioning that Smith (1935) cited Ruiz & Pavón at BM as “ type ” for T. capillaris f. typica . According with ICN Art. 24.3 ( Turland et al. 2018), this designation is not validly published, rendering the inadvertent lectotypification by Smith ineffective.

Note II:— Tillandsia capillaris and related species conform an extremely morphologically variable complex with a complicated taxonomic history. Smith (1935) made significant contributions towards solving this conundrum by circumscribing all this diversity within a single species and recognizing five infraspecific entities [ Tillandsia capillaris f. capillaris , T. capillaris f. cordobensis ( Hieronymus 1885: 10) L.B. Smith (1935: 211) , T. capillaris f. hieronymi ( Mez 1896: 876) L.B. Smith (1935: 213) , T. capillaris f. incana ( Mez 1896: 879) L.B. Smith (1935: 211) , and T. capillaris f. virescens Ruiz & Pavón (1802: 43) L.B. Smith (1935: 212) ]. Later on, Till (1989a) based on morphological data, and more recently Castello et al. (2016) based on molecular evidence, reorganized these entities into two groups, associating each of these groups with a distinct species, T. capillaris s.s. and T. virescens , as it is followed here. It should be mentioned that the entity occurring in Uruguay, treated as T. capillaris in this work, corresponds to what Smith & Downs (1977) considered as Tillandsia capillaris f. hieronymi .

Representative specimens examined:— URUGUAY. Colonia: Riachuelo , October 1949, Fabris & Gebhard 67c ( LP) . Montevideo: Ciudad de Montevideo , 15 December 1972, Morey s.n. ( MVFA 11894 ) .

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 17. Tillandsia capillaris. A. Plants growing on an urban tree (Montevideo, Uruguay). B. Numerous globose individuals of T. capillaris on a dead tree (Montevideo, Uruguay). C. Habit; fruiting plant. D. Habit; flowering plant. E. Close up of a shoot bearing an inflorescence. F. Front view of the flower. G. Lateral view of the flower. H. Capsule. Photos by Andrés Rossado.

FIGURE 18. Distribution map of Tillandsia capillaris in Uruguay. All these populations are subspontaneous.