Tillandsia BROMELIACEAE

Gouda, E. J.

doi:10.1007/978-3-662-56324-3_96-1

E. J. Gouda⁵

Part of the book series: Illustrated Handbook of Succulent Plants ((SUCCPLANTS))

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Abstract

A diagnostic description of the genus is given with special emphasis on the occurrence of succulence amongst its species. The geographical distribution is outlined, together with a selection of important literature, and an explanation of the etymology of the name. This is followed by a short summary of its position in the phylogeny of the family and of the past and present classification in a phylogenetic context. The succulent features present amongst the species of the genus are shortly explained as to morphology and anatomy.

This is followed by a synoptical treatment of the succulent species of the genus, complete with typification details, full synonymy, geographical and ecological data, a diagnostic description, and, where applicable, notes on phylogenetic placement and relationships, as well as economic and/or horticultural importance.

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Tillandsia Linné (Spec. Pl. [ed. 1], 286, 1753). Type: Tillandsia utriculata Linné. — Tillandsioideae — Tillandsieae — Lit: Rauh (1970: ill. synopsis); Smith & Downs (1977: 665–1069, Fl. Neotropica); Rauh (1990: ill. synopsis); Shimizu (1992: ill. synopsis); Roguenant (2001: ill. synopsis); Isley (2009: ill. synopsis); Magalhães & Mariath (2012: seed anatomy); Donadío & al. (2015: morphological phylogeny subgen. Diaphoranthema); Barfuss & al. (2016: phylogeny/classification Tillandsioideae); Granados Mendoza & al. (2017: phylogeny). Distr: S USA to Chile and Argentina. Etym: For Elias Erici Tillandz (also Tillands or Til-Landz, later Tillander) (1640–1693), Swedish-born Finish physician and botanist at the Academy of Turku, and author of the first flora of Finland.

Incl. Renealmia Linné (1753) (nomen rejiciendum, Art. 56.1). Type: Renealmia paniculata Linné.
Incl. Caraguata Adanson (1763) (nom. illeg., Art. 52.1). Type: Tillandsia utriculata Linné.
Incl. Bonapartea Ruiz & Pavón (1802). Type: Bonapartea juncea Ruiz & Pavón.
Incl. Misandra Dietrich (1819) (nom. illeg., Art. 52.1). Type: Bonapartea juncea Ruiz & Pavón.
Incl. Acanthospora Sprengel (1825) (nom. illeg., Art. 52.1). Type: Bonapartea juncea Ruiz & Pavón.
Incl. Dendropogon Rafinesque (1825). Type: Renealmia usneoides Linné.
Incl. Amalia Endlicher (1837) (nom. inval., ICN Art. 36.1a). Type: not typified.
Incl. Buonapartea Sweet (1839) (nom. inval., Art. 61.1?). Type: Bonapartea juncea Ruiz & Pavón.
Incl. Strepsia Nuttall ex Steudel (1841) (nom. illeg., Art. 52.1). Type: Renealmia usneoides Linné.
Incl. Allardtia A. Dietrich (1852). Type: Allardtia cyanea A. Dietrich.
Incl. Anoplophytum Beer (1854). Type: Tillandsia stricta Solander.
Incl. Diaphoranthema Beer (1854). Type: Renealmia recurvata Linné.
Incl. Platystachys K. Koch (1854). Type: Allardtia cyanea A. Dietrich.
Incl. Phytarrhiza Visiani (1855). Type: Tillandsia duratii Visiani.
Incl. Pityrophyllum Beer (1857). Type: Tillandsia erubescens Schlechtendal.
Incl. Wallisia E. Morren (1870). Type: Tillandsia hamaleana E. Morren.
Incl. Viridantha Espejo (2002). Type: Tillandsia plumosa Baker.

Acaulescent or sometimes caulescent herbs of very variable habit; L rosulate or fasciculate or distributed along the stem, polystichous or less often distichous, entire, L lamina tongue-shaped to narrowly triangular or linear, leaf-scales symmetric to asymmetric and extended on one side; Inf usually with a distinct peduncle; floriferous Inf part various, usually consisting of distichously flowered spikes or sometimes reduced to a single polystichously flowered spike by the reduction of the spikes to single flowers, or rarely the whole inflorescence reduced to a single flower; floral Bra conspicuous to minute; Fl bisexual, ± perigynous, seemingly pedicellate but in fact with an expanded receptacle looking like a pedicel; Sep convolute, usually symmetrical, free or equally or posteriorly joined; Pet free, usually naked; St of various lengths relative to petals and pistil; Ov superior, glabrous, ovules usually many, caudate; Fr septicidal capsules; Se narrowly cylindrical or fusiform, with a basal, plumose, straight, white appendage.

Tillandsia is the largest genus in the family, with ±690 species (Gouda & al. 2012+). The genera Tillandsia and Vriesea are traditionally distinguished from each other by the absence (in the first) or presence (in the latter) of ligulae on the claw of the petals. This lead to the transfer of numerous species from Tillandsia to Vriesea, resulting in a growing group of Tillandsia-like Vrieseas. The character has been found to be marginally useful, however, to distinguish natural groups within the family (Brown & Terry 1992, Schulte & Zizka 2008). Moreover, in at least one species (T. incurva Griseb.), ligulae are present in some populations, but absent from others (Gouda 1987). Recently, this group has been transferred back to Tillandsia (Grant 1993). According to Magalhães & Mariath (2012), the structure of the plumose seed appendages could provide a better character to separate Tillandsia from Vriesea, since seeds of the former have a double superimposed parachute-like plumose appendage derived from the exo- and mesostesta, as well as from the endotesta, while Vriesea seeds have a single such structure without participation of the endotesta. The derivation of the plumose appendage from the testa was probably first described by Morra & al. (2002).

In the last monograph of the family (Smith & Downs 1977), 7 subgenera were recognized. Subgenus Pseudo-catopsis was subsequently elevated to genus level as Racinaea by Smith & Spencer (1993). Recent molecular DNA work by Barfuss & al. (2016) supports this move and shows that the traditional subgeneric classification is not fully supported. As a consequence, subgenus Aerobia (Mez 1896) was resurrected for 50 species that were previously placed either in subgen. Allardtia or subgenus Anoplophytum. The revised subgeneric classification is as follows:

[1]
Subgen. Allardtia (A. Dietrich) Baker 1888: St included, exceeding the claw of the petals; Sty slender, much longer than the ovary; Fil straight. — 200 species, 12 of them succulent.
[2]
Subgen. Anoplophytum (Beer) Baker 1887: St included, equalling the claw of the petals; Sty slender, much longer than the ovary; Fil strongly plicate. — 104 species, 13 of them succulent.
[3]
Subgen. Phytarrhiza (Visiani) Baker 1887: Pet blades broad, conspicuous; St deeply included; Sty short and stout; L lamina flat or terete. — 39 species, 7 of them succulent.
[4]
Subgen. Diaphoranthema (Beer) Baker 1878: Pet blades narrow, inconspicuous; St deeply included; Sty short and stout; L lamina terete, densely cinereous-lepidote. — 30 mostly tiny to small species, 22 of them succulent. Donadío & al. (2015) have recently published a phylogeny of the subgenus based on morphology.
[5]
Subgen. Tillandsia: St exserted, exceeding the petals; Pet erect at anthesis or nearly so, relatively firm; L lamina narrowly triangular in most species. — 246 species, 29 of them succulent.
[6]
Subgen. Pseudalcantarea Mez 1934: St exserted, exceeding the petals; Pet spreading at anthesis, rapidly becoming flaccid; L lamina tongue-shapped to narrowly triangular. — 5 species, none of them succulent.
[7]
Subgen. Aerobia Mez 1896: St included, often emerging from the throat of the corolla; Sty slender, many times longer than the ovary; Fil straight or plicate; Pet spreading at anthesis, often with distinct blade; L lamina narrowly triangular. — 50 species, most of them succulent, but not treated here for space reasons.

Succulence: Most subgenera contain species that can be variously described as possessing succulent or semi-succulent features at least to some degree, i.e. showing considerable amounts of water-storing tissues in the leaves. Most mesophytic species have funnel-shaped rosettes with inflated leaf sheaths compounding water, and these are — though superficially appearing to have water-storage capacity — not succulent. All xerophytic species are succulent to some extent, but only 83 species are selected here because they have obviously fleshy leaves. They are all from semi-arid to arid areas, where they grow as epiphytes on cacti and small trees, or are saxicolous on rocky surfaces. In the coastal deserts of Peru and Chile, some species grow on the sand dunes in the so-called Loma vegetation belt, without any other vegetation.

Pollination: Many species, esp. from the largest subgen. Tillandsia, with tubular corolla and exserted stamens and stigma are pollinated by hummingbirds. Birds are attracted by the brightly coloured bracts (often red and yellow) and scentless flowers. Flowers are available over a longer period of time and individual flowers are open for only one or a few days. This is ideal for “trap-line” pollination by birds, where a bird flies a certain route, probably several times a day, from one plant to another. Another group of species, from subgen. Phytarrhiza, have open flowers with broad coloured petal blades, and are often strongly fragrant. The bracts are mostly uncoloured and do not have a role in attracting pollinators. The petals have a narrow claw that together form a narrow tube-like structure, accessible only to the long and very slender tongues of moths and butterflies.

Most species of Tillandsia are self-incompatible and are dependent on pollinating animals for reproduction. The flowers of monocarpic species, however, are autogamous when not pollinated. Also the small-flowering plants from subgen. Diaphoranthema are (partly) autogamous, which explains the pronounced variety of forms within a species and within an area. The flowers of some species of this group are fragrant, e.g. T. usneoides, fragrant during the night only, and probably pollinated by small moths according to Gouda (1995).

Horticulture : Tillandsia is the subject of considerable horticultural interest. Esp. the xerophytic species are frequently seen in cultivation, no doubt in part due to the availability of synoptical literature aimed at hobby collectors, such as Rauh (1990). Several profusely illustrated accounts (e.g. Isley (1987), Isley (2009) or Shimizu (1992)) have further added to the popularity of the genus.

T. aizoides Mez (in A. & C. de Candolle, Monogr. Phan. 9: 866, 1896). Type: Argentina, La Rioja (Hieronymus & Niederlein 850 [B, CORD]). — Distr: SW and S Bolivia (Potosí, Tarija), Argentina (widespread); epiphytic on twigs, or sometimes saxicolous, 120–2800 m.

[4] Plant small, long-caulescent, forming dense clusters or few-branched; stem to 5 cm, many from a single point; L densely rosulate, usually ±1 (−2) cm × 2 mm; L sheaths 0.5 cm, broadly ovate, scarious, with broad nerveless margins, 4- to many-veined, glabrous except at the distal end, distinct from the lamina; L lamina stout, rigid, densely cinereous-lepidote, erect to somewhat divergent, sometimes slightly secund, angular-subulate, convex below, with a furrow adaxially, apex pungent; Inf 1-flowered, often with a reduced flower at the apex, peduncle naked except for 1 or 2 bracts below the flower or with a single bract midway, from very short to 2 cm, angled, glabrous; upper peduncular Bra like the floral bracts, elliptic, often carinate, sulcate and strongly veined, glabrous, acute; floral Bra erect, shorter than the sepals, 5.5 × 3.5 mm, elliptic, membranous, at least 3-veined, ecarinate, glabrous, green, broadly rounded or subapiculate; Sep 7–8 × 3 mm, elliptic or obovate-oblong, scarious, 5- to 7-veined, all connate for 1–2.5 mm, ecarinate, green tinged red, glabrous, broadly rounded or subapiculate; Pet slightly longer than the sepals, spatulate, 13 × 3 mm, cream-coloured with purple-red speckles, brownish when dry, blade obscure or distinct and then suborbicular; St all of equal length, 6 mm, deeply included, exceeding the style; Fil flaccid; Anth basifixed, 1.4 mm; Ov 4 mm, subovoid; Sty slender, shorter than the ovary; Sti capitellate.

T. albertiana Vervoorst (Bromeliad Soc. Bull. 19: 121, 128, ills., 1969). Type: Argentina, Salta (Vervoorst 7255 [LIL, US]). — Distr: NE Argentina (Salta); lithophytic on rocks and cliffs, altitude not recorded. – Fig. 1.

[2] Plant short- to long-caulescent, forming dense clusters or laxly pulvinate; stem much branched, 3 mm ∅; L distichous; L sheaths imbricate, making the stem appear 6 mm thick, 2–2.5 × 1.4 cm, ovate, clasping the stem with overlapping margins, with broad membranous margins, veined or slightly rugose, densely appressed-lepidote abaxially and glabrous adaxially except at the distal end, green or brown, tapering into the lamina; L lamina linear-triangular, fleshy, at the base with extending trichomes at the margins, 5–9 (−17) × 0.4–0.8 cm, dark green flushed with red or red-striated at the base, appressed-lepidote, divergent to less often spreading, veined at the base, strongly canaliculate, convex or involute-subulate when dry, apex abruptly obtuse; Inf 1-flowered (with a short extended rachis), erect; peduncle ±2 cm, concealed by the leaves; peduncular Bra 2, excl. lamina 1.8 × 0.85 cm, the sheathing part glabrous except abaxially towards the apex, with foliaceous lamina, obtuse; floral Bra (excl. acumen) 17–20 (−23) × 9 mm, ovate-oblong, clasping the flower with slightly overlapping margins, margins hyaline, veined, glabrous at the base and adaxially to subdensely appressed-lepidote abaxially, green or with reddish apical margins, rounded and fleshy-acuminate; Fl subsessile; Sep 17–24 × 4–6 mm, narrowly oblong, slightly contracted in the centre, thin, with membranous margins, faintly veined, but distinctly veined when dry, free, ecarinate, green, tinged wine-red at the apex, glabrous or with a few trichomes abaxially, rounded; Pet claw narrow with a broad suborbicular blade or spatulate, 28–44 × to 11 mm, margins slightly crenulate, wine-red to bright red, rounded or obtuse; St all of equal length, 17–22 mm, included and longer than the style; Fil ribbon-like, flat at the base but upwards fleshy, not plicate, green towards the apex; Anth subbasifixed, 4 mm, linear, apex obtuse, yellow; Ov 5 × 2.5 mm, triangular-ovoid (broadest near the base) to prismatic, gradually contracted into the style; Sty slender, 8–17 mm; Sti linear, erect, weakly conduplicate.

T. andicola Gillies ex Baker (J. Bot. 16: 239, 1878). Type: Argentina, Mendoza (Gillies s.n. [K, CGE, GH]). — Distr: Argentina (Catamarca, Buenos Aires, San Juan, Mendoza, Río Negro); epiphytic, altitude not recorded.

[4] Plant flowering to 30 cm tall; stem to >20 cm, branched esp. near the apex; L lax, weakly rosulate or distichous, 4 (−6) cm, cinereous-lepidote with subappressed basally produced scales; L sheaths densely imbricate, ±5 mm apart on the stem, making it appear 3–4 mm ∅, much broader than the lamina, suborbicular, occasionally with glabrous broad veinless margins, ciliate with long scales, several-veined, usually lepidote except at the extreme base or wholly glabrous in age; L lamina 2 mm wide, recurved, veined when dry, subulate, apex acuminate, pungent; Inf 1- or 2-flowered (then internode 8 mm long and rachis extended behind the terminal flower), densely lepidote; peduncle ±0–6 cm, bractless or with 1 bract, densely lepidote; peduncular Bra lanceolate, involute, lepidote, acuminate; floral Bra exceeding the sepals in the first flower to shorter than the sepals in the second one, to 14 × 7 mm, ovate to lanceolate, clasping and closely enveloping the flower, veined, ecarinate, 2× as long as the internodes, densely appressed-lepidote, cinereous to green, acuminate or apiculate; Fl subsessile; Sep 10–12 × 3.5 mm, lanceolate-oblong, with broad hyaline margins, veined, adaxial ones connate for 6 mm, the others less connate, adaxial ones with a strong midvein, green, lepidote from the middle to the apex, apiculate; Pet tongue-shaped, 14 × 3 mm, pale white, the blade narrow, rounded; St deeply included, exceeding the style; Anth basifixed, 1.5 mm, red-orange; Ov 3.5 mm, ovoid, contracted into the style; Sty stout, 3 mm; Sti shortly lobed.

T. angulosa Mez (in A. & C. de Candolle, Monogr. Phan. 9: 868, 1896). Type: Argentina, La Rioja (Hieronymus & Niederlein 851 [B, CORD, F [photo]]). — Distr: C Argentina (La Rioja, San Juan, Mendoza); epiphytic in xeric environments; 1160–1200 m.

[4] Plant small, pulvinate, stem to 3 cm, richly branched at the base; L weakly polystichous or somewhat distichous (to spirally distichous), to 1.5 × ±0.2 cm, coarsely densely cinereous-lepidote; L sheaths suborbicular, thin, with broad nerveless margins, many-veined, densely lepidote to glabrous at the base; L lamina subtriangular, spreading to recurved, subulate, angled, lower ½ canaliculate or grooved, apex mucronate; Inf 1- (rarely 2–) flowered; peduncle almost none or short and concealed by the leaves, with a single bract at the base or bractless; peduncular Bra ovate, involute, prominently veined, densely lepidote, acute; floral Bra slightly to distinctly shorter than the sepals, 12 × 7 mm, ovate, clasping the flower, 5- to 9-veined, sparsely lepidote, acute; Sep 11 × 4 mm, triangular-ovate, 7- to 9-veined, almost free or evenly shortly connate, lepidote; Pet elliptic, 10–13 × 3.5 mm, brown to yellow, obtuse; St included, slightly exceeding the style with part of the anthers; Fil 4–5 × 0.2 mm, whitish; Anth basifixed, ±3 × to 0.5 mm; Ov 3.5 × 2–3 mm, obovoid-pyriform, gradually contracted into the style; Sty slender, 1.5 mm; Sti disc-shaped, shortly lobed.

T. araujei Mez (in Martius, Fl. Bras. 3(3): 600, t. 112, fig. 2, 1894). Type [lecto]: Brazil, Rio de Janeiro (Glaziou 16457 [B, B [photo], P]). — Distr: SE & S Brazil (Rio de Janeiro, São Paulo, Rio Grande do Sul), Paraguay; saxicolous or very rarely epiphytic, 0–850 m. – Fig. 2.

Incl. Tillandsia araujei var. minima E. Pereira & I. A. Penna (1980).

[2] Plant caulescent, flowering 15–30 cm tall; stem simple or few-branched, often curved; L densely polystichous along the stem, 3–7 cm; L sheaths short, broadly triangular, glabrous at the base, white; L lamina rigid, appressed-lepidote, all upwardly secund, keeled, angled-subulate, trichomes with a brown centre, apex attenuate and pungent; Inf simple; peduncle completely covered by the bracts or partly visible, erect or ascending, much exceeding the leaves, slender, glabrous, exposed part red; peduncular Bra with thick linear lepidote lamina, imbricate, elliptic or obovate, membranous, rose-coloured, Inf spikes sublax, 3–5 cm, 5- to 12-flowered; floral Bra exceeding the sepals, 20–23 × 10–12 mm, ovate, membranous, ecarinate, glabrous except the apex in the lower ones, dark pink-red, broadly acute or the lower ones apiculate; Fl sessile, polystichous, erect to slightly divergent, 30 mm; Sep (12–) 15–17 × 4.5 mm, narrowly ovate to lanceolate, with broad hyaline margins, adaxial ones highly connate for ±11 mm, adaxial ones carinate, rose-coloured with green base, acute; Pet spatulate, 20–32 × 8 mm, white, blade tapering into the cuneate claw, spreading, rounded or obtuse; St 17 mm, included; Fil inflated towards the apex, flaccid, lightly plicate near the apex to strongly plicate in the middle; Anth basifixed, 2 mm, yellow; Ov 3 × 2 mm, obovoid (to pyriform), abruptly contracted into the style; Sty slender, 16 mm, exceeding the stamens; Sti weakly conduplicate, erect.

There is some variation in succulence between different populations; the small, short-leaved form (var. minima) is more succulent than the larger ones .

T. argentina C. H. Wright (Bull. Misc. Inform. Kew 1907: 60, 1907). Type: Argentina, Córdoba (Stuckert s.n. [K, GH [photo]]). — Distr: S Bolivia (Chuquisaca, Tarija), N Argentina (Catamarca, Córdoba, Jujuy, Salta, Tucumán, La Rioja); saxicolous or epiphytic in dry habitats, 450–1950 m. – Fig. 3.

Incl. Tillandsia unca Baker (1887) (nom. illeg., Art. 53.1).

[2] Plant distinctly short-stemmed, flowering 7–13 cm tall, rosette solitary or few-branched; L densely rosulate, the outer greatly reduced, 5.5–10 (−13) cm; L sheaths 8–11 × 8–16 mm, broadly triangular-ovate, thin, lustrous, at least the lower ½ glabrous, passing almost imperceptibly into the lamina; L lamina very narrowly triangular, linear, rigid, 3–6 mm wide, appressed-lepidote, divergent and often secundly curved upward, often pulvinate, canaliculate, obtusely keeled below or angular-subulate, apex abruptly acute and pungent; Inf always simple, shorter than to exceeding the leaves; peduncle hidden by the inner leaves and bracts, erect or ascending, shorter than the leaves, ±0–6 cm, glabrous; lowest peduncular Bra foliaceous, imbricate, lanceolate, chartaceous, strongly nerved, glabrous or sparsely pale-lepidote near the apex, stramineous, acuminate; Inf spikes (2–) 3.5–4.5 × (0.7–) 1–1.5 cm, broadly (ob-)lanceolate, strongly complanate, (2- to) 4- to 7-flowered, rachis strongly 4-angled, nearly straight, glabrous, partly exposed; floral Bra imbricate, much exceeding the sepals, 19–25 × 7–10 mm, lanceolate or triangular-ovate, subcoriaceous, even or somewhat veined, sublustrous, ecarinate or carinate towards the apex, so narrow as to expose the rachis in part, glabrous throughout, rose-coloured or green, acuminate; Fl subsessile; Sep 12–18 × 4–5.5 mm, narrowly ovate or elliptic, subequal, free, adaxial ones carinate, green with reddish apex, glabrous, apiculate or broadly obtuse; Pet linear or subspatulate, ±30–34 × 6–7.5 mm, margins entire, bright rose-red except at the base, blade scarcely distinct, divergent to strongly recurved, obtuse or rounded-emarginate; St ±20 mm, included; Fil broadly ribbon-like, flaccid, strongly plicate near the apex, white; Anth basifixed, 4–5 mm, linear, cream-coloured to yellow; Ov 2.5 mm, subprismatic or subglobose, abruptly contracted into the style; Sty slender, 19–24 mm, exceeding the stamens; Sti spreading or recurved, conduplicate.

T. arhiza Mez (in A. & C. de Candolle, Monogr. Phan. 9: 855, 1896). Type: Paraguay, Paraguarí (Balansa 4747 [P, GH [photo]]). — Distr: E Brazil (Minas Gerais), S Paraguay (Paraguari, Guairá); saxicolous, ±540 m.

Incl. Tillandsia rupestris Mez (1896) ≡ Tillandsia arhiza var. rupestris (Mez) Hassler (1919); incl. Tillandsia rupestris var. pendens Chodat & Vischer (1916) (nom. inval., ICN Art. 32.1c); incl. Tillandsia breweri hort. ex D. Butcher (1996) (nom. inval., ICN Art. 29.1).

[3] Plant stout, caulescent; stem to 60 cm; L spirally arranged, 20 cm, white-tomentose-lepidote; L sheaths elongate, broadly elliptic, amplexicaul, glabrous inside and below the middle outside and densely lepidote upwards; L lamina very narrowly triangular, 7–9 mm wide, densely ferrugineous or cinereous -lepidote or tomentose-lepidote, erect or divergent and recurved towards the apex, canaliculate or involute-subulate above the middle, apex filiform attenuate; Inf often simple or depauperate-compound, distichous; peduncle erect, exceeding the leaves, slender, glabrous; peduncular Bra erect, equalling or exceeding the internodes, sublanceolate, tubular-involute, densely lepidote, acute, apiculate; primary Bra like the peduncular bracts, shorter than the spikes, lepidote; Inf spikes densely flowered, to 7.5 cm, 1 cm wide, lanceolate, (single/principal one) to linear, 6- to 12-flowered, rachis angled, slightly flexuous; floral Bra erect, subimbricate, from distinctly shorter than to slightly exceeding the sepals, 15 mm, narrowly elliptic, clasping the flower, chartaceous, veined, glabrous or the lowest sparsely lepidote, apiculate; Sep 12 × 5 mm, elliptic, coriaceous, even, subevenly short-connate for 3 mm, glabrous, acute, obtuse; Pet claw narrow with suborbicular blade, 23 mm long, violet; St deeply included, exceeding the style; Anth linear, apex acute; Ov elongate, prismatic.

T. ariza-juliae L. B. Smith & J. Jiménez Almonte (Phytologia 6: 433, t. 1, figs. 1–2, 1959). Type: Dominican Republic, La Vega (Ariza Julia s.n. [US, Herb. J. J. Jiménez]). — Distr: Dominican Republic (Santiago Rodríguez, La Vega, Santiago Rodríguez); epiphyte in pine forest or on shrubs, 200–900 m.

[5] Plant stemless, flowering 19 cm tall; L ±12, densely cinereous-lepidote; L sheaths forming an ovoid pseudobulb 3–4 cm long, orbicular, greatly inflated, green tinged with purple near the margin, abruptly contracted into the lamina; L lamina 3–4× as long as the sheath, erect to convergent, 9− ± 20 cm × 2–4 mm, involute-subulate and nearly straight, narrowly canaliculate, apex attenuate; Inf simple or rarely branched; peduncle erect; lowest peduncular Bra subfoliaceous, imbricate, the uppermost broadly ovate, subchartaceous, apiculate; primary Bra shorter than the axillary branches; Inf spikes rather laxly to densely flowered, 8 cm, lanceolate, complanate, 2- to 5- (to 8–) flowered, acute, rachis slightly alate, slender; floral Bra imbricate at anthesis, 30 mm, ovate, subchartaceous, veined, ecarinate, >3× as long as the internodes, densely appressed-lepidote, roseate, acute; Fl subsessile; Sep 20 mm, oblong, adaxial ones shortly connate, lepidote, apiculate; Pet forming an erect tube, size unknown, purple; St exserted.

T. baileyi Rose ex Small (Fl. Southeast. US, 246, 1903). Type: USA, Texas (Bailey “26” [226] [US]). — Distr: S USA (Texas); epiphytic in scattered oak forest, 0–1080 m.

[5] Plant stemless, flowering 20–40 cm tall, forming dense clusters; L several, to 40 cm, densely appressed-cinereous lepidote throughout; L sheaths forming a pseudobulb 2–5 cm long, ample, ovate, often conspicuously ciliate-lepidote, merging into the lamina; L lamina linear, at the base 5 mm wide, contorted or involute-subulate, apex attenuate; Inf simple, shorter than to as long as the leaves; peduncle nearly fully concealed by the bracts, erect or ascending, shorter than the leaves, often flexuous, internodes 2–4 mm, upwards decreasing in size, glabrous to cinereous-lepidote upwards at least near the nodes, green; peduncular Bra foliaceous but smaller than the leaves, suberect, few (±4), basal part (excluding lamina) slightly or not amplexicaulous, ± as long as the internodes, upper ones 2.3–2.7 × 1.1 cm, broadly ovate, abaxially densely pale-lepidote, green, shortly and weakly acuminate; Inf spikes, often ascending, (4–) 8–10 × 0.8–1.2 cm, linear, complanate, subdensely 6- to 17-flowered; floral Bra erect, loosely imbricate, exceeding the sepals, 20–23 × 8–10 mm, narrowly ovate-triangular, evenly incurved, closely enveloping the flowers, subchartaceous, margins membranous and with extended scales, smooth or veined, ecarinate, narrow and exposing the rachis in part, <2–3× as long as the internodes, appressed-cinereous-lepidote, glabrous inside except towards the apex, roseate or green, acutish or fleshy-apiculate; Fl with a short pedicel-like receptacle; Sep slightly incurved, 15–18 × 4.5–5 mm, very narrowly elliptic, chartaceous, with hyaline margins, smooth or veined when dry, evenly short-connate for 1–1.5 mm, adaxial ones carinate, green, subdensely cinereous-lepidote, obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 30–38 × 6 mm near the apex, 2.5 mm wide at the base, lower ½ hyaline, purple to pale lavender, triangular-acute; St in 2 whorls of unequal length, ±40 mm, exserted; Fil thread-like in the lower ½ and broadened towards the apex, flat, straight; Anth dorsifixed just below the centre, 4 × 1.2 mm before dehiscence, oblong, base bilobed, apex obtuse, yellow; Ov 4 × 2 mm, ellipsoid, gradually contracted into the style; Sty slender, 36 mm; Sti shortly conduplicate-spiralized.

T. brealitoensis L. Hromadnik (Pl. Syst. Evol. 147(3–4): 280, ill., 1984). Type: Argentina (Hromadnik & Hromadnik 7152b [WU]). — Distr: NE Argentina (Salta); epiphytic in bushes, ±2350 m.

[4] Plant caulescent, few-branched; stem to 10 cm; L densely rosulate, erect, 1.5 cm; L sheaths 5 × 7 mm at the base, amplexicaul, veined, lepidote to glabrous towards the base, passing almost imperceptibly into the lamina; L lamina narrowly triangular, 1 cm, thickly succulent, cinereous-green, densely appressed-lepidote, apex acute; Inf peduncle 1.5–3 cm, glabrous; peduncular Bra 1 only, at the base, membranous, lepidote; spikes 1- or 2-flowered; floral Bra 7.6–11.6 × 4.8–6.8 mm, ovate to triangular, membranous, 9- to 12-veined, lepidote; Sep 7.6–10 × 2.8–3.9 mm, membranous, 6- to 8-veined, the 2 innermost veins joining the central vein towards the tip of the sepal, evenly shortly connate for 2–3 mm, lepidote; Pet tongue-shaped, 9.9–12 × 2.4–3 mm, violet to dark violet, rounded; St included; Fil 3.5–4.8 mm, 1-nerved, pale white; Anth 1.5–2.1 × 0.35–0.55 mm, orange; Ov 2.8–4 × 1.9–2.5 mm; Sty 1 mm; Sti conspicuous.

T. bulbosa Hooker (Exot. Fl. 3: t. 173, 1826). Type: [icono]: l.c. t. 173. — Distr: SE Mexico, Belize, Guatemala, N & E Honduras, Nicaragua, Costa Rica, Panama, Bahamas, Cuba, Jamaica, Dominican Republic, Puerto Rico, Leeward Islands, Trinidad & Tobago, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador; epiphytic, 0–1210 m.

≡ Platystachys bulbosa (Hooker) Beer (1856); incl. Tillandsia bulbosa var. brasiliensis Schultes fil. (1830); incl. Tillandsia bulbosa var. picta Hooker (1847); incl. Pourretia hanisiana Morren ex E. Morren (1847) (nom. inval., ICN Art. 32.1c); incl. Tillandsia erythraea Lindley & Paxton (1850) ≡ Platystachys erythraea (Lindley & Paxton) Beer (1856); incl. Tillandsia pumila Lindley & Paxton (1850); incl. Tillandsia pumila Grisebach (1865) (nom. illeg., ICN Art. 53.1).

[5] Plant stemless, flowering 10–30 cm tall, very variable in size and colour, often in clumps; L 8–15, mostly exceeding the inflorescence, 8–30 cm, outer leaves often reduced and sheath-like; L sheaths forming a small ovoid pseudobulb, ample, 1.5–3.5 cm wide, orbicular, greatly inflated, stiffly coriaceous, densely appressed-lepidote towards the apex, brownish to silvery green at the apex, often purple-red at the margins, abruptly constricted into the lamina; L lamina fleshy-coriaceous, 10–25 cm × 1–5 mm, pale to very dark lustrous-green, minutely appressed-lepidote, reflexed or often spreading, involute-subulate or mostly flexuous, apex attenuate, pungent; Inf simple or subdigitately compound with 2–5 spikes, polystichous, subdense, 7.5–23 cm, green or orange-red; peduncle completely covered by the bracts, erect, 5–15 cm; peduncular Bra foliaceous, exceeding the inflorescence, the upper ones often orange-red in part, with foliaceous lamina, axes short but exposed between the spreading bracts, angled, lepidote, often pale-rose; primary Bra like the upper peduncular bracts, the upper ones more ovate, the lower ones often exceeding the spikes with the foliaceous lamina, acuminate, clasping the spikes; Inf spikes short-stipitate, with a sterile bract at the base, spreading or suberect, dense, 2.5–5 × 0.7–1 cm, lanceolate, complanate, distichously 3- to 6-flowered, acute, with sterile bracts at the apex, rachis sharply angled when dry, nearly straight, lepidote, partly exposed; floral Bra erect, imbricate, much exceeding the sepals, 11–19 mm, ovate, somewhat incurved at the apex, subcoriaceous, (obscurely) carinate, 2–4× as long as the internodes, densely lepidote, acute; Fl sessile, contiguous with each other and/or the rachis; Sep 10–12 mm, oblong, thin or rigid, with broad veinless margins, veined (only when dry), adaxial ones connate for up to 3 mm, adaxial ones bluntly carinate at the base, glabrous, obtuse or obscurely apiculate; Pet forming an erect narrow tube clasping the filaments at the apex, linear, to 40 mm, blue-violet or rarely white, acute; St exserted, the 3 longest ± equalling or shorter than the style; Fil flat or subterete towards the apex, not plicate; Anth dorsifixed at 1/3 from the base, 2.5 mm; Ov 4 mm, ovoid, tapering into the slender style; Sti slender, exceeding the stamens.

T. burle-marxii Ehlers (Bromelie 1994(2): 75, ill., 1994). Type: Brazil, Bahia (Burle-Marx s.n. [WU]). — Distr: NE Brazil (Bahia); ecology unknown.

[2] Plant long-caulescent, to 30 cm long and 1.5 cm ∅, forming clumps, repeatedly branched from the bottom; L spirally arranged, numerous, erect or bent over to the side, densely upright, 2–4 × 0.4–0.5 cm, very thick and succulent, margins with few asymmetric trichomes, both faces densely cinereous-lepidote, grey; L sheaths hiding the stem, 3–6 × to 5 mm, triangular, indistinctly merging into the lamina; L lamina erectly imbricate, slightly keeled, 1.5–3.5 cm, subulate, margins involute, outer face convex, apex triangular-acuminate; Inf simple, green, glabrous; peduncle partly visible, erect, decurved, exceeding the leaves, to 5 cm, slender, glabrous, light green; peduncular Bra few, laxly imbricate leaving the peduncle visible, 2 cm, 1.5× as long as the internodes, elliptic, densely lepidote, acute; Inf spikes arching, sublax, 3–4 × 1.5 cm, with 3–5 polystichously arranged flowers, rachis partly exposed; floral Bra exceeding the sepals, 11–15 (−19) × 7–10 mm, elliptic or suborbicular, membranous, margins hyaline, veined, ecarinate or carinate towards the (fleshy) apex, cinereously lepidote at the apex at both faces, yellow-green, cuspidate at the acute apex; Sep strongly incurved at the apex, 10–14 × 3–5 mm, narrowly elliptic, membranous, smooth or veined (when dry), adaxial ones highly connate for 5–9 mm and carinate, yellow-green, glabrous, acute or narrowly rounded-obtuse; Pet tongue-shaped, 20–23 × 5–6.5 mm, base narrowed to 3 mm, margins erose-dentate, white, the blade tip margins divergent, almost rounded or rounded-emarginate; St 12 mm, included; Fil 10–12 × 1–1.2 mm, broadly ribbon-like, the upper ½ slightly plicate, white; Anth basifixed, 2 × 0.3 mm, linear, cream-coloured; Ov 4–5 × 2.5 mm, ovoid, contracted into the style; Sty slender, to 9 × 0.4–0.5 mm; Sti shortly linear, hardly wider than the style, lobes erect.

T. butzii Mez (in Engler, A. (ed.), Pflanzenr. IV(32): 636, 1934). Type: Mexico, Veracruz (Schiede s.n. [B]). — Distr: SE Mexico, S Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, W Panama; epiphytic in mostly dry open habitats, 16–2300 m.

≡ Tillandsia variegata Schlechtendal (1844) (nom. illeg., Art. 53.1); incl. Tillandsia inanis Lindley & Paxton (1850) (nomen rejiciendum, Art. 56.1) ≡ Platystachys inanis (Lindley & Paxton) Beer (1856) (nomen rejiciendum, Art. 56.1).

[5] Plant stemless, flowering 20–30 cm tall; L few, to 50 cm, margins at first ciliate with coarse scales, wholly finely appressed-lepidote; L sheaths forming a pseudobulb of 2–5 cm ∅, suborbicular, inflated, green and dark brown, green with minute violet spots or wavy patterns; L lamina linear, 3 mm wide, often contorted and involute-subulate, apex filiform-attenuate, Inf digitately compound with 3–5 subequal spikes, or rarely simple; peduncle erect, slender; peduncular Bra leaf-like, imbricate; primary Bra almost leaf-like, the broadly ovate sheath much shorter than the subtended branch which is much exceeded by the linear lamina, concolorous; Inf spikes with 1–2 sterile bracts at the base, erect to spreading, 6–8 × 1 cm (at anthesis), linear, strongly complanate, 5- to 8-flowered, acute; floral Bra erect, imbricate, much exceeding the sepals, 20–28 × 9–10 mm, ovate, sometimes incurved, stiffly subcoriaceous, prominently veined, obscurely carinate towards the apex, 3–4× as long as the internodes, densely pale appressed-lepidote, reddish with green spots or cinereous-green, acute; Fl subsessile, appressed to the rachis; Sep 12–15 × 4.5 mm, narrowly elliptic or obovate, coriaceous, adaxial ones connate for 4 mm, adaxial ones bluntly carinate, green, glabrous, narrowly obtuse; Pet forming an erect narrow tube clasping the filaments, with revolute apex, 30–35 × 7 mm, purple or pale violet, rounded-emarginate; St ±40 mm, unequal, exserted; Fil exposed part slightly inflated and (sub-) terete, not plicate; Anth dorsifixed at 1/3 from the base, 2.3 × 0.7 mm, ellipsoid, purple-blue in the upper part; Ov 5 mm, narrowly ovoid and broadest near the base, tapering into the style; Sty slender, 30 mm, exserted.

T. caliginosa W. Till (Pl. Syst. Evol. 147: 282–284, figs. 6a-4, 7-af, 1984). Type: Argentina, Jujuy (Muhr s.n. [HEID 30925]). — Distr: Bolivia, N Argentina; epiphytic, 1150–3000 m.

Incl. Tillandsia myosura Grisebach (1879) (nom. illeg., ICN Art. 53.1); incl. Tillandsia crocata var. tristis Rauh (1983); incl. Tillandsia crocata fa. major Rauh (1983) (nom. inval., ICN Art. 39.1).

[4] Plant short stemmed, flowering 20–40 cm tall, branched; stem ±8 cm, 0.5 cm ∅, simple; L distichous, ±10; L sheaths densely imbricate and clasping each other and the stem, to 2 × 2.4 cm, strongly veined, glabrous and partly visible, vividly green; L lamina 12–18 × 0.2–0.5 cm, linear-subulate with long-attenuate apex, flexuous, horizontally spreading to reflexed, densely cinereous-lepidote, trichomes eccentric; Inf always simple; peduncle naked except for 1 or 2 bracts at the base, 10–16 cm, slender, straight, subdensely cinereous-lepidote; peduncular Bra leaf-like, remote; spikes erect, densely strongly lepidote, to 6 × 5–10 mm, complanate, 2- to 4-flowered, rachis flexuous, exposed; floral Bra erect, equalling or shorter than the sepals, 12–14 (−20) mm, ovate, enveloping the rachis at the base, margins with extended scales at the margins, veined esp. on the inside, ecarinate, densely cinereous-lepidote only abaxially, brownish-green, obscurely acuminate or subattenuate-acute; Fl with a short pedicel-like receptacle, distichous, erect and contiguous with each other and/or the rachis, fragrant; Sep 14 × 3.5–4 mm, obovate, 5- to 7-veined, evenly short-connate (adaxially slightly more) for 1–2 mm, adaxial one carinate, green and somewhat fleshy, flushed red upwards, then more hyaline, densely grey-scaled at least towards the apex and upper margins (with extended marginal scales) to glabrescent towards the base, acutish; Pet 10 (−15) mm, small, spatulate, 2 mm wide, almost blackish-brown marbled with yellow (and often with a yellow midvein), blade recurved; St and Ov with Sty deeply included.

T. capillaris Ruiz & Pavón (Fl. Peruv. 3: 42, t. 271: fig. c, 1802). Type: Peru (Pavón s.n. [BM, GH [photo], P]). — Lit: Till (1989); Castello & Galetto (2013). Distr: E Brazil (Bahia), S Ecuador (Azuay), Peru, Bolivia, N Chile, Argentina, S Uruguay; saxicolous and epiphytic, 0–4200 m.

≡ Diaphoranthema capillaris (Ruiz & Pavón) Beer (1856); incl. Tillandsia incana Gillies ex Baker (1878) (nom. inval., ICN Art. 36.1c); incl. Tillandsia pusilla Gillies ex Baker (1878) (nom. inval., ICN Art. 36.1c); incl. Tillandsia propinqua Grisebach (1879) (nom. illeg., ICN Art. 53.1); incl. Tillandsia lichenoides Hieronymus (1885) ≡ Tillandsia hieronymi var. lichenoides (Hieronymus) A. Castellanos (1945); incl. Tillandsia propinqua var. saxicola Hieronymus (1885); incl. Tillandsia capillaris var. incana Mez (1896) ≡ Tillandsia capillaris fa. incana (Mez) L. B. Smith (1935); incl. Tillandsia capillaris var. lanuginosa Mez (1896); incl. Tillandsia hieronymi Mez (1896) ≡ Tillandsia capillaris fa. hieronymi (Mez) L. B. Smith (1935); incl. Tillandsia capillaris fa. typica L. B. Smith (1935) (nom. inval., ICN Art. 24.3); incl. Tillandsia permutata A. Castellanos (1945).

[4] Very variable in size and form, flowering to 16 cm tall, forming dense clusters; stem simple or branched, many from a single point; L distichous, mostly 1–4 (−9) cm, rarely shorter, densely and finely punctulate-lepidote with cinereous to ferruginous scales; L sheaths usually elliptic, thin, several-veined, densely lepidote except where covered by the next below; L lamina linear, narrowly triangular, to 2 mm wide, erect to spreading or rarely contorted, straight; Inf 1- (rarely 2–) flowered; peduncle always naked, from almost none to 8 cm, developping almost wholly after anthesis, mostly slender, glabrous to sparsely lepidote towards the apex; floral Bra of the first flower exceeding or shorter than the sepals, that of the upper flower (if any) shorter than the sepals, almost orbicular, thin and soon drying, strongly 3- to 5-veined, densely lepidote to glabrous on both faces, stramineous; Fl subsessile or with a short pedicel-like receptacle; Sep ±8 × 3 mm, veined with strongly developed vein branches, adaxial ones for ±1/3 connate, ecarinate, green tinged red-purple, glabrous on both faces, obtuse or almost round; Pet linearly tongue-shaped to lanceolate, 12 × 2 mm, cream-coloured (sometimes flushed red) or yellow or brown, blade obscure, divergent to recurved, rounded or acutish; St 3.5–4.5 mm, deeply included, exceeding the style; Fil ribbon-like, flaccid, not plicate; Anth basifixed, ±1 mm, oblong-ellipsoid (to slightly tapering from base to apex), yellow; Ov 1.5–2 × >1 mm, obovoid (to pyriform), abruptly contracted into the style; Sty stout, shorter than to about as long as the ovary; Sti capitellate, lobes short, concave and erect.

A very variable species including many forms, some of them more succulent than others. Infraspecific taxa are no longer recognized (Till 1989, 1991), with the exception of fa. virescens, which is recognized as separate species, T. virescens. T. virescens is very similar, however, intermediate forms occur and separation is probably not warranted.

T. caput-medusae E. Morren (Belgique Hort. 30: 90, 1880). Type: K [Morren icono]. — Distr: Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, W & C Panama; epiphytic, 15–2400 m.

Incl. Tillandsia langlassei Poisson & Menet (1908).

[5] Plant stemless, flowering 15–25 (−40) cm tall; L often exceeding the inflorescence, densely subappressed-lepidote; L sheaths forming an ovoid pseudobulb, (broadly) ovate to triangular-ovate, strongly inflated, with membranous margins, adaxially densely ferruginous-lepidote with dark-centered scales, brown to dark brown inside, upwards with purple-red margins, merging into the lamina; L lamina linear-triangular, 10–25 cm, to 1.5 cm wide, involute-subulate or usually much contorted, apex attenuate; Inf simple or digitately composed of 2–6 spikes; peduncle erect, ascending, slender; peduncular Bra leaf-like, densely imbricate; primary Bra usually smaller than the floral bracts, broadly ovate, with a little lamina or without, lepidote; Inf spikes bearing several sterile bracts at the base, suberect to spreading, often curved, to 18 cm, linear-lanceolate, 6- to 12-flowered, acute, rachis angled, slender, nearly straight, glabrous; floral Bra suberect or divergent, imbricate, shorter than to slightly exceeding the sepals, 12–15 (−20) × 8–9 mm, ovate-lanceolate, incurved esp. at the apex, chartaceous, even or veined when dry, lustrous, carinate but sometimes obscurely so (not thickened), exposing most of the rachis, barely more than 2× as long as the internodes, nearly or quite glabrous or obscurely sparsely lepidote on both faces, red or pink or sometimes green, rounded or broadly obtuse; Fl subsessile; Sep incurved especially at the apex, 13–14 × 5 mm, oblong, thin-coriaceous to chartaceous, veined only when dry, lustrous, adaxial ones connate for 5–6.5 mm, adaxial ones sharply carinate, glabrous or very sparsely lepidote adaxially only, rounded or broadly obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 28–44 × 7 mm with narrow base only 2 mm wide, violet (rarely white), with cuneate claw, rounded and then abruptly obtuse; St unequal in length, 30 mm exserted; Fil 53–57 mm, thin and slender and flattened at the base and terete at the distal end; Anth dorsifixed just below the middle, (fresh) 6 × 1 mm, linear, base bilobed, apex apiculate; Ov 4 × 2 mm, ovoid, contracted into the style; Sty exceeding the stamens.

T. castellanii L. B. Smith (Contr. Gray Herb. 104: 80, t. 3, figs. 17–19, 1934). Type: Argentina, Córdoba (Castellanos s.n. in BA 1576 [GH, BA]). — Distr: NW & C Argentina; epiphytic, 1000–3000 m.

[4] Plant flowering to 15 cm tall, forming dense clusters; stem simple, few-branched, many from a single point; L dense, distichous, to 4–5 mm; L sheaths broadly ovate, strongly veined, densely lepidote with a ciliate margin of elongate scales; L lamina sublinear, 2–3 mm wide, spreading, recurved, subulate with pungent apex, cinereous-lepidote with spreading scales; Inf 1- to 2-flowered; peduncle bractless, pseudo-axillary, evident, to 3 cm, slender, glabrous; floral Bra shorter than the sepals, (5.5–) 6.5–9.5 (−10.5) × (5.5–) 5.8–6.8 (−7) mm, ovate, strongly 9- to 11-veined, glabrous or upper part lepidote, broadly acute or short-laminate; Fl subsessile, not fragrant; Sep 7.5–8.5 (−9) × 2.9–3.3 (−3.5) mm, obovate-lanceolate, strongly 7- to 9-veined, evenly short-connate or abaxial one free, glabrous, rounded or obtuse; Pet ovate-lanceolate, 7–8 (−10) × 1.5–2 mm, veined, yellowish, rounded or rarely obtuse; St deeply included, exceeding the style; Fil 3–3.7 × 0.2 mm; Anth basifixed, 2.1–2.2 × ±0.3 mm, whitish; Ov obovoid (to pyriform), 3.2–3.4 mm, abruptly contracted into the style; Sty stout, 1.6–1.7 mm, 1/3 as wide as the ovary; Sti obscurely lobed.

T. chiapensis C. S. Gardner (Selbyana 2(4): 338, ill., 1978). Type: Mexico, Chiapas (Gardner 211 [SEL]). — Distr: SE Mexico (Chiapas); saxicolous, 600–1000 m. – Fig. 4.

[5] Plant stemless, offsetting from the base, eventually forming clumps; L forming a spreading rosette, numerous, 30 cm; L sheaths 4.5 cm wide, suborbicular, often brown-lepidote in the distal part, pale castaneous; L lamina narrowly triangular, fleshy, 2.5 cm wide at mid-blade, coarsely and densely cinereous-lepidote, curved to recurved, strongly canaliculate or the margins involute towards the base, apex attenuate, trichomes ±0.5 mm long, suborbicular; Inf simple, ± equalling or exceeding the leaves; peduncle short and concealed by the leaves and bracts; peduncular Bra polystichous, the lowest leaf-like, densely imbricate, tinged or wholly rose-coloured; spikes suberect or becoming decumbent after flowering, lanceolate; floral Bra distichous, densely imbricate, >2× as long as the sepals, ±40 × 26 mm, ovate-elliptic, incurved at the apex, finely veined when dry, ecarinate or slightly so at the apex, concealing the rachis, 3–4× as long as the internodes, subdensely or densely lepidote, pink, acute or obtuse; Fl distichous, suberect (contiguous with the bracts in the lower part); Sep incurved, 15 × 7.5 mm, elliptic or obovate, coriaceous esp. at the base, veined when dry, free or nearly so, adaxial ones carinate, acute or obtuse; Pet forming an erect narrow tube clasping the filaments, with revolute apex, narrowly tongue-shaped, 75 mm, 8.5 mm wide in the uppermost ¼, 3 mm wide at the base, blue-violet, obtuse; St in 2 whorls of unequal length, 75–80 mm, shorter than the style; Fil subterete towards the apex, purple; Anth dorsifixed at ¼ from the base, 3.5 × 1 mm, oblong-ellipsoid, apex obtuse; Ov narrowly ovoid, merging into the style; Sty slender; Sti conduplicate-spiral, white.

T. circinnatoides Matuda (Cact. Succ. J. (US) 45: 187, figs. 4, 4a, 5, 1973). Type: Mexico, Guerrero (Matuda 38432 [MEXU, US]). — Distr: S & E Mexico, NW Costa Rica; epiphytic on cacti, trees, and bushes in dry areas, 600–2200 m. – Fig. 5.

[5] Plant stemless, flowering 10–20 cm tall, often forming dense clusters; L few, the outer ones reduced and without lamina, the others 7–16 cm, appearing veined by trichomes in longitudinal rows, covered with appressed cinereous scales; L sheaths forming an elongate pseudobulb, large, ovate, merging into the lamina; L lamina narrowly triangular, erect or curved, grooved, involute-subulate; Inf simple; peduncle erect, 7.5 cm, concealed by the leaves; lowest peduncular Bra leaf-like, the upper like the floral bracts or slightly larger, densely imbricate, acuminate; spikes curved or erect, 5–10 cm, 1.4–2 cm wide, lanceolate to linear, complanate, attenuate, rachis hidden or (slightly) partly exposed; floral Bra densely imbricate, exceeding the sepals, 20–30 × 9.5 mm, elliptic, thin, finely veined when dry, scarcely or not at all carinate towards the apex, 3–4× as long as the internodes, densely cinereous-lepidote except the base and margins and less dense on the inside, pale roseate, broadly acute or apiculate; Fl subsessile, contiguous with each other and/or the rachis; Sep 19–25 × 4–6 mm, lanceolate, abaxial one chartaceous and adaxial ones with a rigid subcoriaceous keel, subfree or adaxial ones connate for up to 5 mm, adaxial ones carinate from base to apex, lepidote with few large scales abaxially, or glabrescent; Pet forming an erect narrow tube clasping the filaments with revolute apex, tongue-shaped, 40–45 × 5 mm, pale blue to dark violet in the upper ½, rounded; St long-exserted; Fil twisting at the base, flat and exposed part slightly inflated and (sub-) terete, bluish in the upper ½; Ov ovoid, abruptly contracted into the style; Sty slender, exceeding the stamens; Sti conduplicate-spiralized, long-papillose.

T. colganii Ehlers (J. Bromeliad Soc. 47: 103, ill., 1997). Type: Bolivia, Santa Cruz (Ehlers EB951002 [WU]). — Distr: E Bolivia (Santa Cruz); saxicolous, 2700–3200 m.

[2] Plant short stemmed or with a rhizomatous stem, flowering 10–15 cm tall, often branched; L rosulate, 12–14, 3–7 cm long (outer ones reduced with short acuminate lamina), coriaceous to subsucculent, margins with asymmetric trichomes, covered with appressed cinereous scales, green-grey looking striped; L sheaths erect or suberect, 2–3 × 1–1.8 cm, ovate, passing almost imperceptibly into the lamina; L lamina narrowly triangular, 1–1.5 cm wide at the base, sometimes slightly secund or spreading or recurved, abaxially with a median keel and strongly veined, subulate or canaliculate, apex attenuate; Inf simple, exceeding the leaves; peduncle completely covered by bracts, erect, 0–2 cm and concealed by the leaves, stout, lepidote; lowest peduncular Bra almost leaf-like, imbricate, the apical ones like the floral bracts, becoming glabrous, red, attenuate or acute; spikes with a sterile bract at the base, 3–4 cm, 5–8 (−10) mm wide, lanceolate, subterete, 2- or 3-flowered, acute, rachis 4-angled, nearly straight, glabrous, hidden at anthesis, green; floral Bra imbricate, exceeding the sepals, 16–25 (−28) × 8–10 mm, ovate-triangular, submembranous, adaxially veined, ecarinate, glabrous or the apical 4–6 mm finely grey-lepidote, sublustrous wine-red, acute or cuspidate; Fl sessile, distichous, not fragrant; Sep 14–16 × 4.5–6 mm, narrowly elliptic, membranous, thickened along the keels, free, adaxial ones carinate, greenish with pink tips, acute; Pet erect and throat open, 36–40 × 5 mm, at the base 3 mm wide, margins entire, violet to white at the base, blade spreading to recurved, obtuse; St just visible in the throat of the corolla; Fil 25–30 × 0.5 mm, flat, thin towards the base, once transversely plicate or straight; Anth basifixed, 4–5 mm, linear, apex acute, yellow; Ov 4 × 2.5 mm at the base, ovoid; Sty surpassing the stamens, 25–30 mm, 1 mm wide at the base, tapering to 0.7 mm at the apex; Sti 4 × 3 mm, exserted for 2–12 mm, narrow, spreading, papillose, yellowish.

T. copanensis Rauh & Rutschmann (J. Bromeliad Soc. 38: 7–9, ill., 1988). Type: Honduras, Copán (Kamm s.n. in BG HEID 68087 [HEID]). — Distr: W Honduras (Copán); saxicolous on rock cliffs and epiphytic on pine trees, elevation not recorded. – Fig. 6.

[5] Plant stemless, flowering to 80 cm tall; L numerous; L sheaths 9 × 6 cm, narrowly ovate, brown-lepidote on both faces, distinct from the lamina; L lamina very narrowly triangular, somewhat succulent, to 35 × ±4 cm at the base, pale green, abaxially densely white-lepidote (trichomes in rows with a green centre), recurved, canaliculate, apex attenuate and pungent; Inf mostly laxly 1× branched or rarely simple, composed of up to 6 spikes, often decurved and then ascending, lax, to 30 cm (including the terminal spike), 20 (−30) cm wide; peduncle curved, shorter than the leaves, 15 mm ∅; lower peduncular Bra somewhat leaf-like, the upper ones like the primary bracts, imbricate, spike axes angled, 10 mm ∅, glabrous, bright carmine-red; primary Bra triangular, the upper ones to 9 × 3 cm, acute; spikes ±20 × 2 cm, shortly stipitate for 1.5–3 cm, with 1 or 2 sterile bracts at the base that are sharply keeled, sub-pendulous and then ascending or curved, linear-lanceolate, complanate, acute; floral Bra imbricate, >2× as long as the sepals, 47–50 × 20–23 mm, elliptic, strongly convex, bluntly subcarinate towards the apex and ecarinate below, densely lepidote becoming glabrous at the base, pale green to cinereous-white at the apex, acute or obtuse; Fl with a stout pedicel-like receptacle, contiguous with each other and/or the rachis; Sep ±22 × 5–8.5 mm, oblong-lanceolate, fleshy at the base, with broad hyaline margins and apex, finely veined adaxially, evenly short-connate for 1–2 mm, adaxial ones inconspicuously bluntly carinate, green, glabrous on both faces, obtuse; Pet forming an erect narrow tube clasping the filaments, with revolute apex, tongue-shaped, to 66 × 7.5 mm, broadest at ¼ from the apex, base 4 mm wide, dark violet with hyaline margins and apex, covered part white, blade rounded-emarginate; St in 2 whorls of unequal length, 72 and 78 mm, exserted; Fil exposed part slightly inflated and (sub-) terete, violet; Anth dorsifixed at 1/3 from the base, 4 × 1 mm, elliptic in outline, apex rounded, yellowish-brown; Ov 7 × 3 mm, attenuate from near the base, gradually contracted into the style; Sty slender, to 73 mm, exceeding the stamens; Sti 3 mm, conduplicate-spiral.

T. cotagaitensis L. Hromadnik (Pl. Syst. Evol. 147: 285–285, 1984). Type: Bolivia, Potosí (Hromadnik & Hromadnik 5093 [WU]). — Distr: SW Bolivia (Potosí); epiphytic on Oreocereus in association with T. caliginosa, ±3000 m.

[4] Plant caulescent, flowering to 14 cm tall, branched; L densely distichous, divergent to recurving; L sheaths to 1 × 0.8 cm, amplexicaul, scarious, with broad veinless margins, veined or lustrous, glabrous, becoming greenish; L lamina flexible, to 4 × 0.2–0.3 cm, densely tomentose-lepidote, subulate, apex subulate-acuminate; Inf simple; peduncle 2–3 cm, 1 mm ∅, densely lepidote; spike dense, 2 cm, distichously 2- to 3-flowered, rachis lepidote; floral Bra equalling or shorter than the sepals, (8.2–) 11.8 × 5.4–6.8 mm, ovate, 11- to 12-veined, lepidote, with a very short lamina; Fl subsessile, fragrant; Sep 8.2–10.2 × 2.4–2.6 mm, lanceolate, 5- to 7-veined, adaxial ones connate for 4–6 mm and slightly carinate, abaxial one connate for 1 mm, green, glabrescent; Pet tongue-shaped, 14.2–15.4 × 2.7–3.1 mm, brown-violet to black-violet or rarely ochraceous, blade recurved, moderately broadened, tip rounded; St included; Fil 4.4–5.1 mm, white, 1-nerved; Anth basifixed, 1.9–2.3 × 0.3 mm, golden-yellow; Ov 2–2.3 × 1.5 mm, narrowly ovoid; Sty slender, 4–4.6 mm, included.

T. crocata (E. Morren) Baker (J. Bot. 25: 214, 1887). Type: Brazil, Rio de Janeiro (Lietze s.n. [K [icono: Morren drawing], GH [photo]]). — Distr: SE & S Brazil (Rio de Janeiro, Paraná, Rio Grande do Sul), NE Bolivia (La Paz), NE Argentina (Entre Ríos), SW Uruguay (Soriano); saxicolous, 875–2650 m.

≡ Phytarrhiza crocata E. Morren (1880).

[3] Plant short-stemmed, leafy stem part often shorter than the leaves, flowering 15–35 cm tall, forming dense clusters; stem to 20 cm, L distichous, 10–30 cm, densely white-tomentose-lepidote with asymmetric elongated scales; L sheaths broadly ovate, glabrous except for the upper ½ outside; L lamina linear, 2–5 mm wide, spreading and recurved, involute-subulate, apex long-attenuate; Inf always simple; peduncle bractless or with a single leaf-like bract at the base, erect or nearly so, 5–15 cm, slightly >1 mm ∅, slender, retrorse-tomentose like the leaves; spike densely 2- to 6-flowered, 1–4 cm (excl. petals or capsules), lanceolate, elliptic, acute, with a reduced flower at the apex; floral Bra imbricate, ± equalling the sepals, 9–10 (−20) (including blade like apex) × 6.5–7.5 mm, ovate or elliptic, 2–5× as long as the internodes, densely tomentose-lepidote abaxially only, green, tinged purple-red, acuminate, the lowest with a thick blade-like apex; Fl with a ±1 mm pedicel-like receptacle or subsessile, distichous, fragrant; Sep 10 × 4.5 mm, elliptic or sublanceolate, thin, with veinless broad hyaline margins, faintly veined in the centre, unequally short-connate for 1–2 mm, adaxial ones obscurely carinate esp. towards the apex, lower ½ green and then stramineous speckled purple-red, densely appressed-lepidote abaxially on the centre ridge towards the apex, obtuse or acutish; Pet claw narrow with suborbicular blade or spatulate with elliptic blade, tapering into the cuneate claw, 19–20 mm, blade 6–8 mm wide, spreading, obtuse, margins slightly crenulate, bright yellow (paler towards the base), claw 1 mm wide at the base to 3 mm distally; St 7 mm, deeply included, much exceeding the style; Fil slender-attenuate from near the base, flaccid, straight; Anth basifixed, 1.5 mm, apex broadly obtuse, orange-yellow; Ov 3 × 2 mm, obovoid (pyriform), abruptly contracted into the style; Sty 1.5 mm; Sti erect, shortly lobed.

T. curvifolia (Ehlers & Rauh) Ehlers (Bromelie Sonderheft 6: 126–134, ills., 2009). Type: Mexico, Guanajuato (Ehlers M850201 [WU]). — Distr: C Mexico (Guanajuato); mostly saxicolous, sporadically epiphytic, 1650–3000 m. – Fig. 7.

≡ Tillandsia tortilis ssp. curvifolia Ehlers & Rauh (1990) ≡ Viridantha curvifolia (Ehlers & Rauh) Lopez-Ferrari & Espejo (2009).

[1] Plant stemless or short-stemmed, flowering 5–15 cm tall; L spirally arranged, 8–12, 4–16 cm, white-tomentose-lepidote; L sheaths 1.5 × 1.2 cm, ovate, somewhat succulent, with translucent margins, densely tomentose and glabrous only at the base, conspicuously offset from the lamina; L lamina narrowly triangular, succulent esp. at the base, to 14 cm, 7 mm wide at the base, mostly upwardly secund or erect, silvery-white, subulate and canaliculate, margins involute, apex filiform-attenuate, adaxially densely appressed-lepidote, abaxially tomentose-lepidote, trichomes eccentric; Inf simple or very rarely with a smaller lateral spike, erect, shorter than or slightly exceeding the leaves; peduncle concealed within the leafy rosette or extended and hidden by the inner leaves and bracts, to 5 (−14) cm, 2 mm ∅, glabrous or sparsely lepidote; lowest peduncular Bra leaf-like, upper ones like the floral bracts, erect, few (3–5 only), imbricate, 3–5× as long as the internodes, with setaceous lamina to 7 cm; spikes with a sterile bract at the base, 2–3 × to 1 cm, (narrowly) ellipsoid, terete, (1- to) 2- to 5-flowered, rachis angled, stout, densely lepidote, hidden; floral Bra erect, densely imbricate, slightly exceeding the sepals, 10–20 × 10 mm, ovate to lanceolate, thin, margins hyaline, finely veined or distinctly veined towards the apex, 2–3× as long as the internodes, glabrous or sparsely lepidote, roseate or sometimes green, acuminate; Fl polystichous or subdistichous; Sep to 15 × 4 mm, lanceolate-oblong, membranous, with hyaline margins, free, adaxial ones distinctly carinate, glabrescent or sparsely lepidote at the apex; Pet for most of the length tubular-erect, tongue-shaped, 20–27 × 4 mm, green with white base, blade apex recurved, rounded; St all of equal length, deeply included; Fil to 15 mm, filiform, flat, white; Anth basifixed, 3–5 mm, sagittate, pale brown; Ov 4 × 2 mm; Sty slender, 6 mm; Sti lobes short, erect or somewhat divergent at maturity.

T. diaguitensis A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 36: 55, t. 10, 1929). Type: Argentina, Jujuy (Castillon 7224 [BA]). — Distr: N Argentina (Jujuy, Formosa, Salta, Tucumán), C Paraguay; epiphytic, ±90– 2050 m. – Fig. 8.

[2] Plant long caulescent, slender; stem 60 cm, simple or few-branched, 5 mm ∅; L laxly polystichous along the stem, 8–10 cm, densely cinereous-lepidote (furfuraceous); L sheaths ±1.3 cm wide, elliptic, amplexicaul, imbricate, making the stem appear 7–10 mm thick; L lamina narrowly triangular, 0.6 cm wide, erect to recurved, strongly veined at least when dry, canaliculate, apex attenuate; Inf always simple; peduncle completely covered by the bracts, conspicuous, to 8 cm, 3 mm ∅; peduncular Bra imbricate, elliptic, thin, strongly veined, lepidote towards the apex, stramineous, acute; spike dense, 4–9 cm, 1.3–1.4 cm wide, lanceolate, 3- to 6-flowered, acute, rachis slender, nearly straight, glabrous, sulcate; floral Bra imbricate to divergent at anthesis, much exceeding the sepals, 40–50 × 10 mm, lanceolate, chartaceous, margins scarious and veinless, finely to strongly veined when dry and waxy, ecarinate, 2–3× as long as the internodes, glabrous, roseate or stramineous, narrowly acute; Fl subsessile, fragrant; Sep 26 (−32) × 4.5–6 mm (abaxial one smaller), lanceolate-oblong, with membranous margins, veined, free, adaxial ones carinate, glabrous, acute; Pet spatulate, 60–70 × 13 mm, margins minutely denticulate or slightly crenulate, white or bluish, claw linear, forming a tube well beyond the sepals, blade obovate, rounded; St all of equal length, 38–51 mm, included and visible in the throat; Anth subbasifixed, 5–8 mm, linear; Ov 5–7 mm, tapering into the style; Sty slender, 37 mm, exceeding the stamens; Sti 3 mm, spreading, weakly conduplicate.

T. dorisdaltoniae Ibisch & al. (Revista Soc. Boliv. Bot. 4(1): 45, 2003). Type: Bolivia, Cochabamba (Vargas 6355 [WU]). — Distr: C Bolivia (Cochabamba); epiphyte, 950–1050 m.

[2] Plant short-stemmed, flowering to 20 cm tall or more, forming dense clusters; stem richly branched at the base; L forming a dense rosette, numerous, erect to divergent, exceeding or about equalling the inflorescence; L sheaths to 1.5 × 1 cm, abaxially lepidote, conspicuously offset from the lamina; L lamina rigid, to 10 cm, 0.6–0.7 cm wide at the base, erect or divergent, subulate, apex attenuate, acute, intensely green, sparsely lepidote, adaxially glabrescent towards the apex; Inf simple; peduncle erect, much shorter than the leaves, to 3 cm, slender, green; peduncular Bra imbricate, upper ones reduced to ovate sheaths, lepidote to glabrous on the sheath, whitish, the lower ones with leaf-like lamina; spike erect, 4–5 cm, complanate, densely ±8-flowered, rachis completely covered by the bracts at anthesis but slightly exposed when dry as the margins of the bracts bend outwards, internodes 3.5 mm; floral Bra erect, distichous, densely imbricate, exceeding the sepals, 12–14 (−20) × 5–8 mm (2.5–3 mm wide in lateral view), ovate, faintly veined towards the apex esp. when dry, ecarinate, covering >½ of the flowers, glabrous, bright red, acute; Fl subsessile; Sep 11 × 3.5 mm, membranous, completely free, subadaxial ones alate-carinate, whitish, glabrous, subobtuse; Pet suberect, tongue-shaped, 16 × 3 mm, white, blade slightly divergent, apex rounded; St all of equal length, 12 mm, included; Fil 8.5 mm, flat, slightly plicate in the upper 1/3; Anth subbasifixed, 4 mm; Ov 2 × 1.2 mm, ovoid; Sty slender, ±9 mm; Sti 0.5 mm, erect, papillose.

T. ×dorotheae Rauh pro sp. (Trop. subtrop. Pfl.-welt 60: 59, ill., 1987). Type: Argentina, Salta (BG Heidelberg 31306 [HEID, MO, WU]). — Distr: NE Argentina (Salta); saxicolous in bare lands, in crevices and almost inaccessible crags and bluffs, altitude not recorded. – Fig. 9.

[2] Plant ± caulescent, forming dense clusters; stem 5–10 cm, laxly branched at the base; L densely spirally arranged but sometimes showing a tendency to distichous, erect to spreading; L sheaths distinct, to 1.5 × 1 cm, elongate, with membranous margins, base glabrous, upper ½ densely cinereous-lepidote, whitish or occasionally pinkish-violet; L lamina very narrowly triangular, rigid, 8–10 × 0.3 cm, cinereous-green, densely lepidote, veined and distinctly rugose at the base, adaxially deeply and narrowly canaliculate but not keeled, apex abruptly obtuse; Inf simple, decurved, shorter than the leaves; peduncle mostly decurved, sometimes erect, short and concealed by the leaves, 4–6 cm, 2 mm ∅, terete, green; peduncular Bra few (3–4), the lower leaf-like, the upper 1 or 2 with shorter lamina, densely imbricate; spike decurved, sublax or subdensely flowering, to 3 × 0.8 cm, (1- to) 2- (to 4–) flowered, with a reduced flower at the apex, rachis flat on one side, slender, flexuous, glabrous; floral Bra erect, slightly shorter than the sepals, 20 × 5–8 mm, lanceolate, coriaceous, margins hyaline, shiny at the base, strongly veined, carinate towards the apex, lower part glabrous, scatteredly white-lepidote upwards, green to wine-red, acuminate or fleshy-apiculate; Fl subsessile, distichous; Sep 19–22 × 4 mm, lanceolate, thin, even, free, adaxial ones carinate, green and flushed red, acuminate or abruptly acute; Pet spatulate, 33–35 × 6–7 mm (base 2.5 mm wide) reddish or dark pink, blade obscure, recurved and often twisting, acuminate; St (9–) 13–17 mm, deeply included, shorter than the style; Fil ribbon-like, not plicate; Anth dorsifixed near the base, 5–6 mm, yellow; Ov 5 × 2.5 mm, cylindrical, abruptly contracted into the style; Sty slender, 15 mm; Sti erect, weakly conduplicate.

Considered to be the hybrid T. argentina × T. albertiana.

T. duratii Visiani (Nuovi Saggi Imp. Regia. Accad. Sci. Padova 5: 271, t. 29, 1840). Type: PAD?; [icono]: l.c. t. 29. — Distr: Brazil, N Peru, Bolivia, Argentina, Paraguay, NE & NW Uruguay; epiphyte or saxicolous, 80–2750 m.

≡ Phytarrhiza duratii (Visiani) Visiani (1854) ≡ Anoplophytum duratii (Visiani) Beer (1856); incl. Tillandsia floribunda Durat ex Visiani (1840) (nom. illeg., ICN Art. 53.1); incl. Tillandsia circinalis Grisebach (1874) ≡ Phytarrhiza circinalis (Grisebach) E. Morren ex Baker (1889); incl. Tillandsia gigantea Ruchinger (1876) (nom. inval., ICN Art. 32.1c); incl. Tillandsia decomposita Baker (1889); incl. Tillandsia weddellii Baker (1889); incl. Tillandsia revoluta Burbidge ex Baker (1889) (nom. inval., ICN Art. 32.1c); incl. Tillandsia tomentosa N. E. Brown (1894); incl. Tillandsia confusa Hassler (1919) ≡ Tillandsia duratii var. confusa (Hassler) L. B. Smith (1968); incl. Tillandsia confusa var. minor Hassler (1919); incl. Tillandsia confusa var. saxatilis Hassler (1919) ≡ Tillandsia duratii var. saxatilis (Hassler) L. B. Smith (1968).

[3] Plant caulescent, flowering 20–100 cm tall or more; stem to 30 cm, simple, stout, curved; L laxly polystichous, densely cinereously lepidote with subappressed coarse scales; L sheaths ±2 cm, broadly ovate, distinct from the lamina; L lamina very narrowly triangular, rigid, thick, 15–20 (−40) × 1–2 cm, involute-subulate to often contorted towards the apex that is then twining around tree branchlets, apex pungent; Inf 1× branched or 2× branched in the lower part, 6–60 cm; peduncle erect, elongate, stout, glabrous; lowest or all peduncular Bra almost leaf-like, densely imbricate, elliptic, densely cinereous-lepidote, the upper apiculate; primary Bra erect, like the peduncular bracts, enveloping the bracteate sterile bases of the axillary branches; Br erect or spreading or recurving, lower Inf spikes long-stipitate for up to 10 cm, spikes complanate, 4.5 cm long, 1.7 cm wide, with several sterile bracts at the base, lanceolate or linear; floral Bra erect, slightly longer or shorter than the sepals, to 17 × 8 mm, ovate to oblong, even, ecarinate, subdensely to densely lepidote, greenish but soon stramineous, obtuse; Fl subsessile, fragrant; Sep to 14 mm, ovate, elliptic, coriaceous at the base, with membranous margins and apex, even, evenly or unequally short-connate for 2 mm, ecarinate, green or brown, glabrous, subobtuse; Pet 25–33 mm, claw narrow with suborbicular blade, margins slightly crenulate, usually (purple-) blue-speckled, claw linear, white, blade subrhombic to orbicular, spreading; St all of nearly equal length, 11 mm, deeply included but exceeding the style; Fil thin but not flaccid, not plicate; Anth basifixed, 2.5 mm, green; Ov 7.5 mm, bottle-shaped, ribbed; Sty short; Sti shortly lobed.

This is a very varible species growing in a range of habitats. It can grow on rock faces mixed with T. streptocarpa, looking very different in habit, but in other habitats intermediate forms occur. Intermediate forms between these 2 species, but also with T. paleacea, are common and are difficult to place under one of the 3 species, mainly because the flowers and spikes are not very different if at all from each other. The infraspecific taxa recognized by some authors are not accepted here.

T. ehlersiana Rauh (J. Bromeliad Soc. 34(4): 166–169, ills., 1984). Type: Mexico, Chiapas (Ehlers 83/404 [HEID]). — Distr: SE Mexico (Chiapas); on steep granitic rocks in deciduous forest, ±700 m. – Fig. 10.

[5] Plant stemless, flowering to 20 cm tall, forming clumps; L many, exceeding the inflorescence, silver-grey; L sheaths forming a pseudobulb to 8 × 10 cm, individual sheaths to 7 × 6–7 cm, suborbicular, strongly inflated, densely furfuraceously lepidote, adaxially light leather-brown, abaxially whitish, distinct from the lamina; L lamina narrowly triangular to linear, with asymmetric trichomes at the margins, above the sheath to 2.5 cm wide, both faces densely white-lepidote, upper part often recurved, strongly involute-subulate and canaliculate; Inf 1× branched, composed of ±4 spikes, ovoid in outline, much shorter than the leaves; peduncle short and concealed by the leaves, to 7 cm, 1 cm ∅, terete, appressed-lepidote; peduncular Bra leaf-like, few (±3), similar to the upper rosette leaves, silver-greyish; axes compressed, 7 cm, straight; primary Bra erect or divergent with the branches, the basal ones with broadly ovate sheaths, somewhat leaf-like, 2 × 1 cm, exceeding the spikes or inflorescence and the upper shorter than the spikes and acute, densely furfuraceously lepidote; spikes shortly stipitate for 0.5 cm, erect or divergent, 3 × 1.8 cm, complanate, 2- to 3-flowered; floral Bra distichous, imbricate, much exceeding the sepals, 19–22 (−28) × 10–13 mm, triangular-ovate, membranous, margins even, carinate with a thick midvein, concealing the rachis, cinereously lepidote, brownish-red or roseate, acute or obscurely apiculate; Fl subsessile, 30–65 mm; Sep 13–19 × 6–7 mm, (narrowly) elliptic, membranous, even, free or shortly connate for up to 3 mm, adaxial ones carinate (sharply so when dry), greenish-red, (very) sparsely lepidote especially on the keel, acute or broadly obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, linearly tongue-shaped, (33–) 42–53 × 8 mm, upper ½ blue-violet, white towards the base, obtuse; St in 2 whorls of unequal length, 57–60 mm, exserted, shorter than or as long as the style; Fil subterete towards the apex, straight, upper 1/3 violet; Anth dorsifixed at 1/3 from the base, 2.5 mm, blackish; Ov 5 mm, subprismatic, gradually contracted into the slender style.

T. erecta Gillies ex Baker (J. Bot. 16: 239, 1878). Type: Argentina, Mendoza (Gillies s.n. [K]). — Distr: C Bolivia (Cochabamba), Argentina (Mendoza, La Rioja); saxicolous and epiphytic, 1300–2610 m.

Incl. Tillandsia rigida Gillies ex Baker (1878) (nom. inval., ICN Art. 32.1c) ≡ Tillandsia erecta var. rigida (Gillies ex Baker) Baker (1889).

[4] Plant to 11 cm tall; stem 3–6 cm, simple or few-branched; L laxly polystichous, to 5 cm, densely cinereous-lepidote; L sheaths 0.5–1 cm, broadly ovate, glabrous; L lamina very narrowly triangular, rigid and succulent, ±0.4 cm wide at the base, erect to divergent, veined or obviously grooved, involute-subulate, angled towards the acuminate apex; Inf 1-flowered with the rachis extended behind the flower; peduncle to 4.5 cm, elongating in fruit, slender, strongly sulcate or strongly furrowed when dry, glabrous; peduncular Bra 0–2, ±1.5 cm, elliptic, glabrous; floral Bra equalling or shorter than the sepals, (8–) 12–13.5 × 5.5–6 mm, triangular-ovate to ovate-lanceolate, not tapering at the base, distinctly 6- to 10-veined, glabrous or upper part sparsely lepidote, stramineous or brown-reddish in the lower part, acute or obtuse; Fl subsessile, erect, weakly fragrant; Sep 10–11.5 × 3–4 mm, narrowly elliptic or narrowly lanceolate-triangular, 5-veined, evenly short-connate or adaxial ones somewhat more connate, glabrous or sparsely lepidote, acute or obtuse; Pet broadly tongue-shaped, 9–12.5 × 2.5–3 mm, dirty greenish-yellow or greenish-punctate, blade narrowly elliptic, rounded; St deeply included, exceeding the style; Fil 3–5 × ±0.2 mm, whitish; Anth (sub-) basifixed, 2–2.5 × 0.5 mm; Ov ±2 mm, obconical or subcylindrical, abruptly contracted into the style; Sty stout, only 1/5–1/3 as long as the ovary; Sti capitellate, flat.

T. erici Ehlers (Bromelie 1998(1): 18–21, ills., 1998). Type: Bolivia, Tarija (Haugg 11,247 [WU]). — Distr: S Bolivia (Tarija); epiphyte, 1800–2200 m. – Fig. 11.

[2] Plant stemless, flowering 12–22 cm tall, in small clusters or solitary; L 12–20, forming an erect rosette to 25 cm ∅, almost erect, fleshy to coriaceous, strongly veined (prominently when dry), finely appressed-lepidote, reddish-brown; L sheaths 2.5–4 × 1.4–2 cm at the base, triangular-ovate, inconspicuously offset from the lamina; L lamina very narrowly triangular, 6–18 cm (outer ones reduced and only 10–12 cm), 0.8–1 cm wide at the base, very finely appressed-lepidote (adaxially less), margins strongly incurved, apex acute and pungent; Inf simple, shorter than the leaves; peduncle completely covered by the bracts, decurved, only 3–4 cm, ±4 mm ∅; lowest peduncular Bra leaf-like, the upper like the floral bracts, densely imbricate, 3.5 cm, coriaceous, lower ones caudate, the upper acute; spike with 1–2 sterile bracts at the base, often ± decumbent, 4–7 cm, 0.8–2 (after flowering to 3) cm wide, lanceolate or elliptic, complanate, (1–) 3- (to 7–) flowered, acuminate; rachis 4-angled, almost straight, glabrous, green; floral Bra imbricate, exceeding the sepals, 18–35 × 8–13 mm, narrowly ovate, rigid and coriaceous, margins broadly hyaline, veined (strongly when dry), shiny, ecarinate, concealing the rachis, glabrous, stramineous or green tinged purple, acute; Fl sessile or with a short bicarinate pedicel-like receptacle, distichous, not fragrant; Sep 18–26 × 6–9 mm, elliptic or oblong, membranous, with broad hyaline margins, slightly finely veined, free or shortly connate for 1–2 mm (abaxial one less if at all), adaxial ones bluntly carinate, pale green or brown tinged purplish-red, glabrous, acute; Pet spatulate, throat opening, 40–60 × 9–12 mm, claw 3 mm wide, margins crenate-serrate and slightly undulate, bright orange-yellow, blade spreading and recurved; St included or ± equalling the petal claw; Fil 25 mm, ribbon-like, filiform, thin, straight, yellowish-white, upwards yellow; Anth basifixed, 4–5 × 0.4 mm, linear, yellow; Ov 5 × 2.2 mm, slenderly prismatic; Sty slender, 30 mm; Sti lobes 14 mm, spreading or recurved, apex somewhat papillose for ±4 mm, yellow.

T. ferrisiana L. B. Smith (Bromeliad Soc. Bull. 10: 92, ills., 1960). Type: Mexico, Sinaloa (Ferris & Mexia 5121-A [DS]). — Distr: Mexico (Sinaloa, Baja California); epiphytic in dry woods and scrub, ±630 m.

[5] Plant forming dense clusters or few-branched; stem 2–6 (−10) cm; L laxly spirally arranged, much exceeding the inflorescence, to 15 cm, densely cinereous-lepidote; L sheath ±1 cm, suborbicular to narrowly ovate, densely cinereous-lepidote but glabrous where overlapping; L lamina very narrowly triangular to linear, involute-subulate, apex filiform-attenuate; Inf simple, erect, to 3 cm; peduncle short and concealed by the leaves; peduncular Bra somewhat leaf-like, densely imbricate; spike dense, terete, (1- to) 2- to 3-flowered; floral Bra exceeding the sepals, 20 mm, ovate, thin, veined towards the apex, ecarinate, densely cinereously-lepidote, apiculate; Fl subsessile, distichous, ±50 mm; Sep 12–20 × 6 mm, elliptic to lanceolate, membranous, smooth, free, adaxial ones carinate, appressed-lepidote or becoming glabrous, obtuse; Pet erect with recurving apex margins, 50 mm, bicoloured, the base red-violet to purple, the upper 20 mm yellowish or cream-coloured (Rauh 1979); St and Sty long exserted.

T. fresnilloensis W. Weber & Ehlers (Feddes Repert. 94(9–10): 609–611, fig. 9, 1983). Type: Mexico, Zacatecas (Ehlers s.n. [HAL (Herb. Weber)]). — Distr: C Mexico (Zacatecas); ecology and altitude not recorded. – Fig. 12.

[5] Plant flowering to 22 cm tall; stem richly branched at the base; L forming an erect rosette, numerous, white-lepidote; L sheaths to 2.5 × 2 cm, triangular to ovate, strongly wrinkled when dry, indistinctly merging into the lamina; L lamina narrowly triangular, to 9 cm, in the middle to 1 cm wide, erect and secund, strongly grooved, apex acuminate; Inf simple; peduncle to 12 cm with internodes 2–2.5 cm, 2 mm ∅, terete, smooth, with weak purplish tint; peduncular Bra erect, just exceeding the internodes, lanceolate, veined, sparsely appressed-lepidote to almost glabrous, stramineous, finely apiculate; spike 3- to 6-flowered, rachis strongly geniculate; floral Bra shorter than the sepals, 20–23 × 10–12 mm, ovate or lanceolate, margins broadly hyaline, faintly veined or even, ecarinate, glabrous on both faces, green and tinged red at the base, rounded and apiculate; Fl with a stout pedicel-like receptacle, spreading out, ±40 mm; Sep to 25 × 9 mm, narrowly obovate or lanceolate, with broad hyaline margins, finely veined adaxially, connate for 2–3 mm, ecarinate but with very fleshy midvein, green, flushed red at the apex, glabrous on both faces, rounded or obscurely apiculate; Pet tongue-shaped, 42 × 9 mm (3 mm at the base), greenish-white or green, acute; St in 2 unequal series of 46 and 51 mm, exserted but shorter then the style; Fil fleshy throughout, slender and flattened at the base and dilated-terete at the distal end, green with paler base; Anth dorsifixed near the middle, 2.5 mm, apex and base both truncate, green; Ov 4 mm, ovoid, gradually contracted into the style; Sty slender, 51 mm, twisting; Sti shortly conduplicate-spiral and slightly dilated, green.

T. funebris A. Castellanos (Anales Mus. Nac. Hist. Nat. Buenos Aires, ser. 3, 37: 502, 1933). Type: Argentina, Tucumán (Schreiter 1689 [BA, B, GH, LIL]). — Distr: Venezuela, Bolivia, N to E & C Argentina, Paraguay; epiphytic in dry forest, 200–2100 m.

[4] Plant small, flowering rarely over 10 cm tall, forming dense clusters; stem 2–5 cm, few- to richly branched at the base but otherwise simple; L laxly polystichous, to 5 cm; L sheaths making the stem appear stout, 0.5–0.8 cm, suborbicular to reniform, densely imbricate, subcoriaceous, with broad veinless margins, many-veined, lepidote and glabrous below, distinct from the lamina; L lamina narrowly triangular, densely cinereous-lepidote with small nearly symmetrical subappressed scales, divergent to reflexed, often distinctly keeled below, ± contorted, triangular-subulate, strongly angled but not at all sulcate-striate, with a furrow above that does not reach the attenuate apex; Inf simple, 1- or 2-flowered, glabrous; peduncle distinct, nearly naked, erect or curved, much exceeding the leaves, to 6 cm, 0.5 mm ∅, slender, glabrous; peduncular Bra 1–2 just below the flowers, 1.3 (−1.7) cm, oblong-lanceolate, subcoriaceous, even or many-veined, glabrous, acute; spike rachis like the peduncle and nearly as thick, swollen at the nodes; floral Bra slightly shorter or longer than the sepals, 9–15 × 4.5–5 mm, like the upper peduncular bract but more ovate or elliptic, progressively smaller upwards, even, ecarinate, subacute; Fl subsessile, erect; Sep 9.5–10 × ±3 mm, narrowly elliptic, even or finely veined (many-veined), evenly shortly connate for <1–2 mm, adaxial ones with a strong midvein but ecarinate, acute; Pet spatulate, to 14 × 3 mm, yellowish or coffee-brown when fresh and dark orange-brown when dry, blade distinct, subrhombic, spreading or near the tip margins recurved to helicoid, obtuse; St slighly unequal in length, 6–7 mm, deeply included; Fil slightly shorter than the style; Anth basifixed, 1.5–2 mm, linear, apex acute; Ov 2.5 mm, ellipsoid or shortly cylindrical, obtusely angled, contracted into the thick style; Sty short, 2 mm; Sti broadly capitate.

T. gilliesii Baker (J. Bot. 16: 240, 1878). Type: Argentina, Mendoza (Gillies s.n. [K, CGE, GH]). — Distr: S Peru (Arequipa), Bolivia, Chile, Argentina; epiphytic, 310–3200 m.

Incl. Tillandsia compressa Gillies ex Baker (1878) (nom. inval., ICN Art. 36.1c); incl. Tillandsia andicola Wittmack (1887) (nom. illeg., ICN Art. 53.1).

[4] Plant caulescent, flowering 4–6 cm tall, forming dense clusters; stem to 8 cm; L very densely arranged, distichous, 2–8 cm, densely cinereous-lepidote; L sheaths densely imbricate making the stem appear 5–7 mm ∅, broadly oblong, margins not overlapping behind the stem except at the extreme base, many-veined, glabrous where covered or lepidote upwards, passing gradually into the lamina; L lamina narrowly triangular, arching-recurved and often somewhat contorted, subulate and narrowly triangular, canaliculate above and much compressed laterally, apex acuminate and pungent; Inf peduncle distinct to almost absent, bractless or with 1 lanceolate involute bract at the very base, 5–13 cm, appressed-lepidote; Inf proper mostly 2-flowered, to 3.8 cm, densely cinereous-lepidote; spike rachis flexuous or straight, lepidote; floral Bra erect, equalling or exceeding the sepals, (11–) 14–16 (−18.5) × (7–) 9–10.5 mm, the uppermost sometimes slightly shorter, ovate-lanceolate to broadly triangular, thin, many-veined (±15 veins), distinctly so at the apex, ecarinate, slightly to 2× as long as the internodes, lepidote, acute or obtuse; Fl subsessile, erect, subdensely arranged; Sep (11.5–) 12.5–14.5 × 4–6 mm, ovate-lanceolate or narrowly ovate-triangular, thin, many-veined (±10 veins), evenly shortly connate, adaxial ones carinate, lepidote towards the apex or glabrous, apiculate or obtuse; Pet tongue-shaped, 12–15 × 3–4 mm, pale yellow, blade narrow, broadly rounded; St deeply included, exceeding the style; Fil 5–6 × ±0.3 mm, whitish; Anth subbasifixed, 2.5–4 × to 0.9 mm; Ov 3.5–5.5 × 2.5–3.5 mm, subglobose or obovoid (pear-shaped), abruptly contracted into the style; Sty stout, if at all slightly tapering, shorter than the ovary, ±1.5 mm; Sti erect, slightly wider than the style, flat-capitate.

T. glabrior (L. B. Smith) López-Ferrari & al. (Selbyana 25(1): 60, 2004). Type: Mexico, Oaxaca (Foster & Van Hyning 2937 [US]). — Distr: S Mexico (Oaxaca); saxicolous on canyon walls, 910–1220 m. – Fig. 13.

≡ Tillandsia pueblensis var. glabrior L. B. Smith (1958) ≡ Tillandsia schiedeana ssp. glabrior (L. B. Smith) Gardner (1984).

[5] Plant caulescent, flowering to 35 cm tall; L spirally arranged, exceeding the peduncle and much shorter than the inflorescence, densely appressed-lepidote; L sheaths 1.5 × 1 cm, broadly ovate, sulcate towards the base, moderately contracted into the lamina; L lamina narrowly triangular, distinctly fleshy-succulent, whitish-cinereous-green, upwards secund, arcuate-recurved, to 11 cm and to 0.7 cm wide above the sheath, subulate and canaliculate; Inf simple; peduncle completely covered by its bracts, suberect, terete, 6–8.5 cm, 2 mm ∅; lower peduncular Bra somewhat leaf-like and laminate, erect, lamina gradually reduced in size distally, imbricate, longer than the internodes, densely enveloping the peduncle, appressed-lepidote; spike to 7 × 0.7 cm, lanceolate, subterete, ±5-flowered, acute, rachis partly exposed when dry; floral Bra erect, laxly imbricate, much exceeding the sepals, to 30 × 10 mm, lanceolate, veined, ecarinate, concealing the rachis or so narrow as to partially expose the rachis, ±3× as long as the internodes, scatteredly appressed-lepidote to subglabrous; Fl distichous; Sep to 18 × 6 mm, ovate-lanceolate, adaxial ones connate for 8 mm, carinate, glabrous; Pet forming an erect narrow tube clasping the filaments at the apex, size not recorded, (rose-)red or partly yellow towards the apex; St exserted; Anth dorsifixed near the middle; Ov not described; Sty just exceeding the stamens; Sti lobes slightly divergent, green.

T. guerreroensis Rauh (Trop. subtrop. Pfl.-welt 60: 65–67, ills., 1987). Type: Mexico, Guerrero (Koide s.n. in HEID 66721 [HEID]). — Distr: S Mexico (Guerrero); saxicolous, ±1000 m.

[1] Plant stemless or with a short rhizome, flowering to 15 cm tall; Ros 12 × 18 cm; L numerous; L sheaths distinct, 2 × 1.5 cm, broadly ovate, both faces densely leather-brown-lepidote; L lamina very narrowly triangular, to 8 × basally 0.8 cm, spreading to strongly recurved, fleshy, with extending trichomes at the margins, strongly canaliculate, both faces densely cinereous-lepidote, margins ± involute, apex attenuate and abruptly pungent; Inf simple or rarely digitate with 2 spikes; peduncle erect, very short, largely concealed by the leaves and bracts, mostly <5 cm; lowest peduncular Bra somewhat leaf-like, the upper like the floral bracts, erect, densely imbricate; spikes decumbent, to 7 × 1.5 cm, lanceolate, complanate with convex sides, 5- to 6-flowered, acuminate; floral Bra suberect and contiguous with each other, densely imbricate, much exceeding the sepals, 25 × 12 mm, ovate, margins membranous, veined esp. when dry, ecarinate, concealing the rachis, abaxially subdensely lepidote masking the colour in part, rose-red, with pale green edges, broadly rounded; Fl distichous, erect appearing at the upper side of the curving spike; Sep 15 × 5 mm, membranous, free, adaxial ones sharply carinate, whitish with reddish midvein, glabrous, acuminate; Pet forming an erect narrow tube clasping the filaments at the apex, the tips spreading, tongue-shaped, to 30 × 8 mm, deep dark violet; St exserted; Ov not described; Sty shorter than the stamens.

T. hasei Ehlers & L. Hromadnik (Bromelie 1996(3): 76, ill., 1996). Type: Bolivia, Chuquisaca (Hase & al. s.n. [EB 951501, WU]). — Distr: S Bolivia (Chuquisaca); on rocks in a river bed, ±3100 m. – Fig. 14.

[2] Plant strongly rooted on rocks, short-stemmed, flowering rosette 6–10 × 2–2.5 cm, forming large mats; L 10–20, densely rosulate, erect or spreading, the outer ones reduced and to 6 cm, robust, very thick or stiffly succulent, margins with asymmetric trichomes, densely cinereous-lepidote, grey; L sheaths 1–2 × basally 0.7–1.2 cm, elliptic to triangular, lepidote with fine grey scales on both faces but adaxial base glabrous for 1–2 mm, slightly distinct from the lamina; L lamina very narrowly triangular, 2–4 cm, base 0.4–0.5 cm wide, strongly secund, densely lepidote on both faces with large grey scales, strongly veined particularly abaxially, adaxially keeled, margins involute, subulate, apex attenuate or long-caudate; Inf always simple; peduncle erect or decurved, very short and concealed by the leaves, 1–2 cm, ±2 mm ∅, terete, glabrous; lowest peduncular Bra somewhat leaf-like and 2 cm, the upper shorter and like the floral bracts, densely imbricate, lepidote; spike 2–2.5 cm, 3–6 mm wide, linear-lanceolate, complanate, distichously 1- to 3-flowered, acute, with 1 sterile bract at the apex, rachis 4-angled, sinuous or almost straight, glabrous, exposed; floral Bra imbricate, exceeding the sepals for 5–8 mm, 15–23 × 5–7 mm at the base, triangular, coriaceous, strongly veined, towards the base glabrous but lepidote in the upper 1/3 and esp. at the tip with large grey scales, red, cuspidate; Sep 13–15 × 3 mm, lanceolate, very thinly coriaceous, veined, free, adaxial ones strongly carinate, light green with a red tip, glabrous, acuminate; Pet erect, linear without a distinct blade, 25–29 × 2.5 mm, shiny carmine-red, white at the base, blade tip margins recurving or helicoid, rounded; St 15 mm, included; Fil thin, ribbon-like, plicate in the middle, white; Anth basifixed, 3.5–4 × 0.4 mm, linear, olive-green; Ov 2 × 1 mm, ellipsoid; Sty 15 mm; Sti lobes 0.8 mm, inconspicuous, spreading.

T. hegeri Ehlers (Bromelie 1994(2): 33, ill., 1994). Type: Bolivia, Tarija (Heger s.n. [WU]). — Distr: S Bolivia (Chuquisaca, Tarija); saxicolous, 2700–2720 m. – Fig. 15.

[1] Plant stemless or very short-stemmed, flowering 10–14 cm tall; Ros broad, basally offsetting; L ±12, erect to becoming spreading horizontally, to 12 cm, very thick-leathery or succulent, margins with asymmetrical trichomes, densely cinereous-lepidote, somewhat reddish-grey; L sheaths 3–4 × 2–2.3 cm, ovate, passing gradually into the lamina; L lamina narrowly triangular, canaliculate or subulate towards the apex, with a central keel, outer face very strongly veined, apex attenuate; Inf simple, terminal, exceeding the leaves; peduncle very short and concealed by the leaves; lowest peduncular Bra somewhat leaf-like and similar to the rosette leaves, densely imbricate, 1–1.5 cm, upper ones similar to the floral bracts, ovate, coriaceous, lepidote to glabrous, wine-red, acute; spike with 1 sterile bract at the base, 6 × 2 cm, elliptic, ±10-flowered, acuminate, rachis almost straight, glabrous; floral Bra imbricate, exceeding the sepals, 14–16 mm wide, ovate, thinly coriaceous, veined on the inner face, shiny, often slightly waxy, carinate towards the apex, concealing the rachis at anthesis, glabrous on the outer face, dark wine-red, acute or cuspidate; Fl sessile, distichous, not fragrant; Sep 20–24 mm, elliptic, thinly coriaceous, evenly shortly connate for 2 mm, adaxial ones strongly carinate, red with strong thicker green centre, glabrous, acute; Pet forming an erect narrow tube clasping the filaments at the apex and the tips spreading with opening throat, almost tongue-shaped, 36–39 × 7 mm, base 4 mm wide, margins finely serrulate, blue-violet, white towards the base, blade slighly if at all divergent, obtuse; St included; Fil 20–25 × 1.2 mm in the basal 1/3, ribbon-like, almost straight, wavy at the base, white; Ov ovoid; Sty as long as the petal tube, narrowing towards the tip; Sti 4 mm, lobes strongly spreading, papillose, yellowish.

T. helmutii L. Hromadnik (Bromelie 1990(3): 63, ill., 1990). Type: Bolivia, Chuquisaca (Hromadnik 13,009 [WU]). — Distr: S Bolivia (Chuquisaca); saxicolous on vertical cliffs, 1700–1800 m. – Fig. 16.

[1] Plant short-stemmed, flowering 50 cm tall; Ros solitary or few-branched; L densely rosulate, the upper ones erect and those below spreading, 20–30 cm, becoming stiff, margins fimbriate; L sheaths ±2 × 4 cm, broadly triangular, thickly leathery, with membranous lobes at the margins, finely veined, lower 1 cm of the adaxial face glabrous, upper ½ and abaxially densely appressed-lepidote with radial trichomes, greenish-yellow; L lamina narrowly triangular, thick, succulent, 1.7–2.5 cm wide at the base, densely lepidote with prominent dorsiventral silvery scales, smooth and upper ½ strongly grooved to finely veined, apex apiculate; Inf simple or 1× branched and subdigitate, exceeding the leaves; peduncle not wholly hidden by its bracts, erect, ± equalling the leaves, 5 mm ∅, smooth towards the base, internodes 2–3 cm, glabrous, green; lowest peduncular Bra somewhat leaf-like, 4.5–6 cm, narrowly ovate, the top ones convex, thickly leathery, veined, densely lepidote, green, apiculate; primary Bra like the peduncular bracts or narrowly ovate with distinct tip, 4–4.5 × 1.5 cm, thickly leathery, veined, apiculate, abaxially densely appressed-lepidote, adaxially glabrous except at the tip, greenish to pale pinkish-orange; spikes up to 5, almost sessile, erect, 8 × 1.9 cm, strongly complanate, long acuminate, rachis straight, sparsely lepidote with scattered punctulate scales, partly exposed, with cavities, green, internodes 0.7 cm; floral Bra densely imbricate, 25–40 × 12–15 mm, narrowly or triangularly ovate, convex, fleshy-coriaceous, margins membranous, strongly veined when dry, obscurely carinate towards the apex and bicarinate at the base, lower ones lepidote, upper ones glabrous, green or pale pink-orange, apiculate; Fl densely distichous, 55 mm, not fragrant; Sep 20–30 × 9.5 mm, ovate, fleshy in the lower ½ to membranous upwards, veined, evenly short-connate or at least the adaxial ones connate for 1 mm and bluntly carinate, green, glabrous, slightly acuminate or attenuately acute; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 41–47 × 5–6 mm, margins slightly undulate at the apex, white and tinged pale violet on the blade, blade indistinct, divergent to spreading, obtuse; St 28–32 mm, included; Fil thin and ribbon-like, flaccid, straight, basally white, near the tip pale violet; Anth basifixed, 3.5–4 mm, linear, pale yellow; Ov 6–7 × 2 mm, shortly cylindrical or slenderly ovoid, merging into the style; Sty slender, 23 mm, exceeding the stamens and emerging from the corolla throat; Sti spreading or erect, linear, papillose, lobes white.

T. hirta W. Till & H. Hromadnik (Pl. Syst. Evol. 147: 285–286, 1984). Type: Bolivia, Cochabamba (Hromadnik & Hromadnik 5001 [WU]). — Distr: S Peru (Arequipa), Bolivia, NE Argentina; epiphytic on trees, bushes or cacti, 1800–3700 m.

[4] Plant short-stemmed, flowering 10–20 cm tall, few-branched; L distichous, 2–6 cm, densely silvery-white tomentose-lepidote; L sheaths 0.8 × 0.6 cm, ovate, clasping the stem, membranous, veined, densely tomentose but glabrous towards the base; L lamina 1–5 × 0.2–0.3 cm, ± recurved with the tips spreading and bending outwards, subulate, apex acuminate; Inf simple; peduncle naked or rarely with 1 bract just below the flower, 1–6 cm, 1 mm ∅, subglabrous, spike 1–2.5 cm, 1- to 3-flowered; floral Bra erect, equalling or exceeding the sepals, 7–15 × 5 mm, ovate, lanceolate, veined, ecarinate, densely lepidote, acute; Fl subsessile, distichous; Sep 10–12 mm, lanceolate, thin, veined, evenly shortly connate, densely lepidote; Pet erect, forming a tube, 30–40 mm, yellowish, blade narrow; St included; Fil 1-veined, (3.2–) 3.8 (−4.9) mm, white; Anth basifixed, (1.4–) 1.7 (−2.1) × ±0.3 mm, orange; Ov (3.8–) 4.3 (−5.2) × (1.8–) 2.2 (−2.7) mm; Sty much shorter than the ovary; Sti inconspicuous.

T. intermedia Mez (Bull. Herb. Boissier, sér. 2, 3: 141, 1903). Type: Mexico, Guerrero (Langlassé 370 [G, F, GH, US]). — Distr: Mexico (Guerrero, Jalisco, Sinaloa, Colima, Yucatán); epiphytic, 0–290 m.

Incl. Tillandsia paucifolia var. schubertii F. Ebel & Röth (1988).

[5] Plant flowering to 40 cm tall or much longer when growing from shoots from the inflorescence; L to 35 cm; L sheaths forming an elongate pseudobulb, 2.5 cm wide, inflated, densely cinereous-lepidote, merging gradually into the lamina; L lamina linear, rigid, covered with dense pale appressed lepidote trichomes for most part, erect or irregularly curved, involute-subulate or more canaliculate at the base, apex subpungent; Inf simple or 1× branched, composed of up to 4 almost equal-sized spikes; peduncle shorter than or much exceeding the leaves, slender; peduncular Bra somewhat leaf-like with conspicuous decurved involute lamina, shorter than the internodes; primary Bra like the upper peduncular bracts but with a shorter lamina, decurved, densely lepidote abaxially; spikes subsessile, strictly erect, sublax, 4–10 × 1.2 cm, elliptic, 3- to 12-flowered, acute, rachis lepidote, partly exposed, green-brown; floral Bra distichous, becoming further apart but laxly imbricate, slightly exceeding the sepals, 18–20 × 8 mm, triangular-ovate, convex, coriaceous, distinctly veined, ecarinate (but bi-angled at the base), so narrow as to partly expose the rachis, abaxially densely to sparsely lepidote with small appressed scales, green and tinged reddish at the apex and margins or roseate, narrowly rounded; Fl with a short pedicel-like receptacle, distichous, suberect, densely arranged, 50 mm; Sep elliptic, thinly coriaceous, even or prominently veined near the apex, adaxial ones connate for up to 3 mm, adaxial ones carinate at the base, green and tinged red, glabrous, broadly acute; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 38 × 5.5 mm, basally 2 mm wide, purple, blade tip margins recurving, acutish or obtuse; St 47 mm, exserted; Fil exposed part slightly inflated and (sub-) terete; Anth dorsifixed at ¼ from the base, 3 mm, apex obtuse, brownish; Ov 5 × 2.5 mm, ovoid, gradually contracted into the style; Sty slender, exceeding or ± equalling the stamens.

T. ionantha Planchon (Fl. Serres Jard. Eur. 10: 101, t. 1006, 1855). Type: [icono]: l.c. t. 1006. — Distr: SE & S Mexico, Guatemala, C & S Honduras, NW El Salvador, W & C Nicaragua, NW & W Costa Rica, W Panama, S Peru; epiphyte, 0–3050 m. – Fig. 17.

Incl. Tillandsia erubescens H. Wendland (1854) (nom. illeg., ICN Art. 53.1) ≡ Pityrophyllum erubescens (H. Wendland) Beer (1856); incl. Pourretia stricta hort. ex Beer (1856); incl. Tillandsia quesneliana hort. ex Beer (1856); incl. Tillandsia scopus Hooker fil. (1871) (nom. inval., ICN Art. 32.1c); incl. Tillandsia rubentifolia Poisson & Menet (1908); incl. Tillandsia ionantha var. zebrina B. T. Foster (1982).

[5] Plant stemless or rarely shortly caulescent, usually forming dense clusters; stem stout and fleshy; L rarely >6 cm, densely cinereous-lepidote with spreading scales; L sheath ±½ as long as the lamina, elliptic; L lamina narrowly triangular, 5 mm wide at the base, green or the inner ones (or all) becoming deep red at anthesis, densely cinereous-lepidote to tomentose-lepidote, divergent esp. upwards, subulate and canaliculate except the attenuate apex; Inf appearing to be a simple abbreviated spike with polystichous flowers, but actually a reduced compound panicle; peduncle nearly none, axes abortive; primary Bra narrowly ovate, ±23 × 5 mm, equalling or exceeding the sepals, membranous, acute, lepidote towards the apex; spikes reduced to a single flower each; floral Bra like the primary bracts but shorter than to slightly exceeding the sepals, 13–20 × 4–8 mm, triangular-ovate, faintly veined, obtuse; Fl sessile, erect; Sep 16–18 × 4–5 mm, lanceolate to oblong-lanceolate, membranous, with broad hyaline margins, evenly short-connate for 1 mm, or free, adaxial ones carinate, glabrous, acutish; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped, 40–48 × 7 mm, base 3 mm wide, violet, claw linear-cuneate, blade tip margins recurving, subacute or rounded; St in 2 whorls of unequal length, 45–56 mm, long-exserted; Fil exposed part slightly inflated and (sub-) terete, straight, exposed part violet; Anth dorsifixed at 1/3 from the base, apex obtuse or rounded, yellow; Ov 6 × 2–2.5 mm, narrowly ovoid, tapering into the style; Sty slender, 42–52 mm, exceeding the stamens, dilatated in the upper part; Sti spreading, white.

A very variable species with several published varieties and forms. The most different and succulent is var. vanhyningii M. B. Foster that grows on steep rock surfaces and has a more cinereous appearance.

T. kirschnekii Rauh & W. Till ex Rauh (Trop. subtrop. Pfl.-welt 43: 13–14, ill., 1983). Type: Peru, Apurímac (Kirschnek s.n. in BG Heidelberg 39,770 [HEID]). — Distr: S Peru (Apurímac); rock walls, ±2500 m. – Fig. 18.

[3] Plant long caulescent, richly branching from the base forming big cushions; leafy stem part to 15 cm, 3–5 mm ∅; L densely rosulate, numerous, erect; L sheaths ±1 × 1.5 cm, very broadly ovate, tightly clasping the stem, membranous, glabrous in the lower 2/3 to densely cinereous-lepidote abaxially in the uncovered upper part; L lamina narrowly triangular, succulent, 2–3.5 (−4) × 0.4 cm, cinereous-green, densely lepidote on both faces with brown-centred trichomes, suberect or slightly divergent and then curved inwards, angled subulate and shallowly canaliculate in the lower ½ and with a furrow in the centre, apex acute or filiform-attenuate; Inf simple, short to much exceeding the leaves; peduncle completely covered by its bracts, erect, 1–3 cm, mostly short and concealed by the leaves, glabrous; lowest peduncular Bra somewhat leaf-like, the upper like the floral bracts, few (2–4), imbricate, cinereous-lepidote, the upper ones only acute without lamina; spikes 1–3 × 1 cm, elliptic to lanceolate, strongly complanate, (1- to) 3- to 5-flowered, apex irregular with sterile bracts, rachis alate-excavated, slightly flexuous, partly exposed or hidden, green, internodes 2–5 mm; floral Bra imbricate, slightly exceeding the sepals, 10–16.5 × 4–7 mm, narrowly ovate or elliptic, closely enveloping the flowers, chartaceous, margins scarious and veinless, strongly veined when dry, ecarinate, so narrow as to expose the rachis in part, ±3× as long as the internodes, abaxially lepidote but sparsely so towards the base, green, obtuse or obtuse-apiculate; Fl with a short pedicel-like receptacle, distichous, contiguous with each other and/or the rachis, densely arranged; Sep 9–14 × 5–5.5 mm, obovate, slightly asymmetrical, membranous, with broad hyaline margins, finely veined (strongly so when dry), evenly shortly connate (obscure when fresh), adaxial ones carinate, green and rose or tinged purplish towards apex and margins, glabrous on both faces, obtuse or acutish; Pet claw narrow with suborbicular (to subdeltoid) blade, 22–25 × 12–15 mm, blue-violet with white claw, blade abruptly broadened, spreading, broadly rounded; St 7 mm, deeply included, exceeding the style; Fil flat, lanceolate, tapering from near the base, thin; Anth basifixed (?), 1.5 mm, yellow; Ov 3 × 2 mm, pyriform or ellipsoid, abruptly contracted into the style; Sty stout, 1 mm; Sti capitate.

T. klausii R. Ehlers (J. Bromeliad Soc. 38(6): 257–260, ills., 1988). Type: Mexico, Chiapas (Ehlers & Ehlers EM851801 [WU, HAL, WU]). — Distr: SE Mexico (Chiapas); saxicolous on steep rocks. – Fig. 19.

[5] Plant stemless, flowering 15 cm tall, forming clumps; L forming a subbulbous or a spreading rosette, few to many, to 17 cm, succulent and soft or brittle, green to rose or red in bright sun; L sheaths suberect, to 4 × 3.5 cm, broadly ovate, ± inflated, densely cinereous-lepidote on both faces, both faces (pale) brown or abaxially green (in cultivation), conspicuously offset from the lamina; L lamina narrowly triangular to linear, to 13 × 2 cm wide, densely cinereous-lepidote with spreading scales on both faces, spreading to recurved, trichomes with elongated wings, margins involute-subulate and nearly touching each other when dehydrated; Inf simple, erect, ± equalling to much exceeding the leaves; peduncle erect, short and concealed by the leaves, 1–5 cm, 7 mm ∅; peduncular Bra similar to the floral bracts but smaller, erect, few, densely imbricate, to 3.5 cm, elliptic, densely lepidote, green-rose, shortly acuminate; spike (3–) 5–9 × to 2.2 cm, elliptic or lanceolate, complanate, 2- to 4- (to 7-) flowered; floral Bra erect, ± imbricate, >2× as long as the sepals, 35–45 × 15–20 mm, narrowly ovate, faintly veined, ecarinate, mostly concealing the rachis, 2–3× as long as the internodes, densely lepidote except towards the base, pink or roseate, acute; Fl with a short pedicel-like receptacle, not fragrant; Sep 22–24 × 4–5 mm, lanceolate to narrowly oblong, subcoriaceous, with broad hyaline margins, veined, connate for ±1 mm, fleshy, adaxial ones carinate at the base (sharply so when dry), green at the base but centre part red, glabrous, acutish; Pet erect, forming a tube, tongue-shaped, 50–70 × 9–10 mm, basally 4 mm wide and white, upper ½ blue-violet, blade tip margins recurving, rounded; St 70–75 mm, exserted; Fil flat, thicker towards the apex but still complanate, thin and flaccid at the base, sometimes slightly twisted, white at the base and violet upwards; Anth dorsifixed at 1/3 from the base, 3–4 × 1 mm, pale brown; Ov 7 × 2.5 mm, cone-shaped with flat sides, contracted into the style; Sty slender, more robust in the upper 1/3, ±70 mm, exceeding the anthers for 8 mm; Sti 3 × 2 mm, ± spreading, papillose, white.

T. kuehhasii W. Till (Bromelie 1995(2): 33, ill., 1995). Type: Bolivia, Chuquisaca/Potosí (Kühhas & Kirschnek s.n. [WU]). — Distr: S Bolivia (Chuquisaca, Potosí); saxicolous, 3600 m.

[4] Plant forming dense clusters; stem to 25 cm but usually only terminal 10 cm with living leaves, branched at several nodes along the stem; L densely distichous, distally straight, flexible, white; L sheaths ±1.5 cm, broadly ovate, amplexicaul, densely lepidote, merging gradually into the lamina; L lamina subulate, (2.5–) 5–7.5 cm, 2 mm wide at the base, densely cinereous-lepidote, suberect or irregularly spreading, upper face with a furrow; Inf 1-flowered, erect, ±5 cm, shorter than to as long as the leaves; peduncle with a single bract at the base, partly visible, very short or slightly elongate, 2–3.5 cm, elongating after flowering, 1 mm ∅, grooved when dry, sparsely lepidote to almost glabrous towards the base; peduncular Bra involute around the peduncle with overlapping margins, cinereous-lepidote, with leaf-like lamina; floral Bra shorter than the sepals, 13 × 10 mm, broadly ovate to ovate-oblong, distinctly 10- to 12-veined, abaxially densely lepidote only, cinereous-green or tinged brown towards the margins, rounded and apiculate or caudate with a 2 mm long apiculus; Sep (10–) 13.5–16 × 3.5 mm, lanceolate, veined, adaxial ones connate for 5–7 mm, adaxial ones bluntly if at all carinate or narrowly convex, green or tinged brown-red at the apex, abaxially sparsely to subdensely appressed-lepidote towards the apex, apiculate or obtuse; Pet tongue-shaped, 15–25 × 4 mm, yellow tinged brown to chestnut-brown with yellow claw, apical part recurved to recoiled; St ±10 mm, deeply included, much exceeding the pistil; Fil flat, attenuate from near the base, straight; Anth basifixed, slightly <2 mm, narrowly oblong, yellow; Ov 2.5 mm, obconical, abruptly contracted into the style; Sty stout, ± as long as the ovary; Sti erect, short and capitellate.

T. latifolia Meyen (Reise um die Erde 2: 45, 1835). Type: Peru, Arequipa (Meyen s.n. [B]). — Lit: Rauh & Bismarck (1996). Distr: C Venezuela (Aragua), Ecuador (widespread), Peru (widespread); terrestrial or epiphyte, 0–3500 m. – Fig. 20.

≡ Platystachys latifolia (Meyen) K. Koch (1874); incl. Tillandsia kunthiana Gaudichaud (1842) ≡ Platystachys kunthiana (Gaudichaud) Beer (1856); incl. Tillandsia grisea Baker (1887); incl. Tillandsia oxysepala Baker (1888); incl. Tillandsia lanata Mez (1905); incl. Tillandsia murorum Mez (1913).

[1] Usually caulescent, flowering to 60 cm high; stem often branched, prostrate; L to 20 (to 30 and more) cm, covered with appressed cinereous scales; L sheath slightly distinct from the lamina; L lamina narrowly triangular, to 3 cm wide at the base, usually spreading to recurved, canaliculate, apex acute or long attenuate; Inf 1× branched or rarely simple, dense or lax, sometimes viviparous; peduncle (almost) completely covered by the bracts, shorter than or exceeding the leaves but always distinct, from short to ±40 cm, 3.5 mm ∅, stout, glabrous; lowest peduncular Bra leaf-like, slender with spreading to recurved blades, imbricate, the upper more lanceolate, cinereous-lepidote; primary Bra like the upper peduncular bracts, suberect, shorter than the spikes, apiculate or acute; spikes subsessile, erect or mostly spreading sidewards (from between the bract and axis), 1 cm wide, lanceolate, complanate, 6- to 12-flowered; floral Bra densely imbricate, slighly shorter than to exceeding the sepals, 10–13 (−23) mm, broadly ovate, ± incurved, strongly convex, stiffly subcoriaceous, even or nearly so, upper ½ finely or obscurely carinate, subdensely cinereous-lepidote with large trichomes or generally becoming glabrous with age, base often green to wholly red, tip acute or finely apiculate; Fl subsessile; Sep incurved, 10.5–12 (−20) × 3.5–4 mm, narrowly ovate or lanceolate, nearly even, adaxial ones connate for 5.5–8 mm, adaxial ones sharply carinate from base to apex, green and with red upper ½ or apex, sparsely lepidote or glabrescent, acutish or obtuse; Pet erect, (ob-)lanceolate, 18–19 (−27) × 3–4 mm, blades to 7 mm longer than the sepals (according to Mez), upper ½ (deep) pink, blade narrow, suberect or slightly divergent, narrowly rounded to obtuse or acutish; St 13 mm, included, exceeding the style; Fil broadly ribbon-like, flaccid, plicate in the upper ¼; Anth dorsifixed near the base or at 1/5 from the base, 3.5 mm, linear, obtuse, yellow; Ov obovoid (to pyriform), 3 mm, gradually contracted into the style; Sty slender, 8 mm, attenuate from the base upwards; Sti erect, shortly conduplicate with papillae only near the apex, pink at the base.

Very variable in habitus, esp. as to leaf size and succulence. The variety from the Peruvian desert forming adventitious shoots has short fleshy leaves. The status of the published varieties and the relationships with closely similar species have not yet been fully investigated.

Plants depicted on Mochica pottery from the Río Moche area near Trujillo dating from 600 BCE to 1000 CE are likely this species and/or T. purpurea, making them the earliest-figured species of the genus (Rauh & Bismarck 1996).

T. lepidosepala L. B. Smith (Proc. Amer. Acad. Arts 70: 155, t. 2, figs. 2–3, 1935). Type: Mexico, Michoacán (Pringle 5323 [GH]). — Distr: Mexico (Aguascalientes, Guanajuato, Michoacán); saxicolous and epiphytic, dry slopes, 1460–2500 m.

≡ Viridantha longisepala (L. B. Smith) Espejo (2002).

[1] Plant stemless or short-stemmed, flowering to 15 cm tall; Ros 18 cm ∅, irregular and open, often in groups or forming clumps; L laxly arranged, few to many, spreading, to 16 cm, fairly succulent, densely cinereous-lepidote; L sheath 1–1.5 × to 1.5 cm, ovate or suborbicular, somewhat inflated, lepidote but glabrous towards the base; L lamina narrowly triangular, to 15 cm, 0.7 cm wide at the base, silvery cinereous-lepidote, spreading to recurved, slightly canaliculate, apex acuminate; Inf simple or more rarely with a small second lateral spike, shorter than the leaves; peduncle short and concealed by the leaves, slender; peduncular Bra leaf-like, the lower roughly as long as the inflorescence, the upper like the floral bracts, erect, densely imbricate, much exceeding the internodes; spikes 3–5 × (1–) 1.2–1.8 (−2) cm, complanate with ± flat sides, 2- to 5-flowered; floral Bra densely imbricate, exceeding the sepals, 19–21 (−35) × 6–9 mm, narrowly or triangular-ovate, almost membranous, finely veined when dry, ecarinate, 3–4× as long as the internodes, densely lepidote, cinereous to dark green at the base, shortly fleshy-acuminate; Fl sessile, densely distichous; Sep 12–15 (−20) × 4–5.5 mm, widest just above the base, lanceolate or narrowly ovate, with broad hyaline margins, strongly veined when dry, free or unequally short-connate for up to 1.5 mm, adaxial ones carinate, green, subsparsely lepidote on both faces, attenuately acute or obscurely apiculate; Pet erect with slightly divergent apex, linearly tongue-shaped, 20–25 × 2.5–3.5 mm, basally only 2.5 mm wide, green, narrowly rounded; St 15 mm, included, longer than the style; Fil 15–18 mm, thread-like, thin, straight, white; Anth dorsifixed near the base, 2–3 mm, linear, apex obtuse, olive-green; Ov 4–5 × 3.5 mm, ellipsoid, gradually contracted into the style; Sty slender, incl. stigma 8–10 mm; Sti shortly conduplicate, small, erect or the green lobes somewhat divergent, papillose at the apex only.

T. lithophila L. Hromadnik (Bromelie 5: 48–51, ills., 2005). Type: Peru, La Libertad (Hromadnik & Hromadnik 23,115 [USM, WU]). — Distr: NW Peru (La Libertad); saxicolous on steep rocks, ±2200 m.

[1] Plant with very short stem, flowering to 20 cm tall, growing in groups; L forming a narrow tuft-like rosette, polystichous, erect, 6–12 cm; L sheath 1.5 cm wide at the base, triangular, subamplexicaul, appressed-lepidote, indistinctly merging into the lamina; L lamina narrowly triangular to linear, rigid and somewhat succulent, 0.7 cm wide at the base, silvery and densely lepidote, secund-arcuate, involute-subulate, apex attenuate and subpungent; Inf 1× branched or rarely simple, composed of up to 4 spikes, 5.5 × 1.5 cm; peduncle erect or decurved, much exceeding the leaves, to 12 cm, 1–2 mm ∅, densely white-lepidote; peduncular Bra densely imbricate, longer than the internodes, to 2.5 cm, coriaceous, veined, abaxially densely lepidote, adaxially glabrous, purplish-red, with an awl-like tip, keeled; primary Bra like the upper peduncular bracts, 1.5–1.7 cm, caudate-acuminate; spikes short-stipitate, stipe 2–3 cm, bearing 2–3 small sterile bracts at the base, erect to somewhat divergent, 3–3.5 cm, 6–9 mm wide, lanceolate, strongly complanate, 4- to 7-flowered, rachis lepidote, exposed, internodes 3–4 mm; floral Bra distichous, densely imbricate, 13 × 6 mm, triangular to ovate, coriaceous, veined, ecarinate, adaxially glabrous, abaxially densely lepidote, purplish-roseate, white-caudate; Fl sessile, to 20 mm; Sep 12 × 2 mm, lanceolate, chartaceous, 3-veined, connate for 3 mm, adaxial ones alate-carinate, rose, abaxially lepidote only along the midvein; Pet tongue-shaped, to 18 × 2 mm, white, with a blue band above the sepals (changing to pale violet immediately after flowering), blade tip margins divergent, obtuse; St 15 mm, included and visible in the throat, as long as the style; Fil filiform, straight, white; Anth dark brown; Ov 3 × 1 mm; Sti lobes suberect and very thin.

T. loliacea Martius ex Schultes fil. (in Roemer & Schultes, Syst. Veg. 7(2): 1204, 1830). Type: Brazil, Bahia (Martius s.n. [M, GH [photo]]). — Lit: Till (1989). Distr: Brazil, Bolivia, Argentina, Paraguay, Uruguay; saxicolous and epiphytic in semi-arid habitats, 0–2400 m.

Incl. Tillandsia undulata Baker (1887); incl. Tillandsia quadriflora Baker (1889); incl. Tillandsia atrichoides S. Moore (1895).

[4] Plant with a rhizomatous stem, flowering to 17 cm tall but usually much less, stem rarely >4 cm, most of it leafless, simple or few-branched; L densely rosulate, mostly 2–3 (−4) cm, densely cinereous-lepidote; L sheath ±0.3 cm, only slightly broader than the lamina, subcoriaceous, with broad hyaline margins, veined, glabrous, whitish; L lamina very narrowly triangular, rigid, 0.3–0.5 cm wide, densely cinereous-lepidote with ± spreading scales, erect to suberect or arching-secund, subulate, apex long-attenuate but not filiform; Inf simple or very rarely forked; peduncle (nearly) completely covered by its bracts, curved, mostly much exceeding the leaves, to 10 cm but often much less, ±1 mm ∅, straight, lepidote near the nodes, dark green; peduncular Bra like the floral bracts, many, ± equalling the internodes, elliptic or obovate, tubularly amplexicaul, chartaceous, veined, densely lepidote, acutish; spikes to 4 cm, linear, sinuous, very much like those of Lolium, to 16-flowered but often with very few flowers or reduced to a single flower at the apex, rachis excavated next to the flowers, ± flexuous, lepidote with large scales; floral Bra not at all imbricate, shorter than the sepals, 3–6 (−8) × 3.5–4 mm, (broadly) ovate, closely enveloping the flowers, thin, margins hyaline, veined, ecarinate, small and exposing most of the rachis, just exceeding the internodes, only abaxially subdensely lepidote with large green centered scales, cinereous-green, acute or apiculate; Fl with a short pedicel-like receptacle, distichous, erect, appressed to the rachis except the upper part of the sepals and petals caused by the flexuous rachis; Sep 5.5–6 (−9) × 3.5–4 mm, (oblong) lanceolate, faintly 7-veined, evenly short-connate for >1 mm, green or spotted brown-red, glabrous, acutish or obtuse; Pet 10 mm, claw sublinear, yellow, blade narrow, spreading or recurved, obtuse or narrowly rounded; St 4 mm, deeply included, exceeding the style with much of the anthers; Fil filamentous, thin; Anth basifixed, ±1.5 mm, yellow; Ov shortly cylindrical, obovoid or pyriform, abruptly contracted into the style; Sty stout, shorter than the ovary.

T. lydiae Ehlers (Bromelie 2000(2): 55–59, ills., 2000). Type: Mexico, Guerrero (Köhres & Köhres s.n. [MEXU, WU]). — Distr: S Mexico (Guerrero); epiphytic, 350–1000 m.

[5] Plant flowering 20–30 cm tall; Ros 3–4 cm ∅; L 5–8, forming a bulbous rosette, 25–30 cm, the outer greatly reduced, very thick and succulent, stiffly leathery, margins very strongly grooved, densely appressed-lepidote, silver-grey but lustrous; L sheaths erect, 6–10 × 3–4 cm, narrowly ovate, firmly pressed together, inflated, appressed-lepidote, glabrous towards the base, inside brown, outside dark green, conspicuously offset from the lamina; L lamina narrowly triangular, 10–20 cm, 2–2.5 cm wide at the base, red-brown, with striking longitudinal brown-red stripes, almost erect or slightly divergent, in dry conditions appearing grooved or strongly veined, subulate, margins involute, apex acuminate tapering to a sharp tip; Inf 1× branched or simple, composed of (1–) 2–5 spikes; peduncle with a single bract similar to the inner leaves, short and concealed by the leaves, 4 mm ∅, sparsely lepidote; primary Bra with a basal flat part like the floral bracts, 1–1.8 cm, but with an acute short lamina, densely grey-lepidote; spikes short-stipitate, stipe 2–4 cm, with 2–3 (bi-) carinate sterile bracts at the base, spreading at an angle of 45–90°, 5–12 cm, 8–10 (−12) mm wide, lanceolate, complanate, 3- to 11-flowered, rachis with internodes 8–10 mm; floral Bra 3–5 mm shorter than the sepals, 13–15 × 6–8 mm, ovate, veined, towards the apex keeled, densely appressed-lepidote, pink, acute; Fl distichous, contiguous with each other and/or the rachis; Sep 20–22 × 6–7 mm, elliptic, with membranous margins, subfree, ecarinate or adaxial ones slightly carinate at the base, raspberry-red except the base, glabrous, acute; Pet erect, forming a tube, spatulate, 35 × 6 mm (base 4 mm wide), pale violet with white base, blade tip margins divergent; St in 2 whorls of unequal length, exserted; Fil 34–36 mm, broadened towards the apex and 0.8 mm wide, the upper portion pale lilac tapering to the white base; Anth dorsifixed at 1/3 from the base, 3 × 0.75 mm (5 mm before dehiscence), linear or somewhat sagittate, dark yellow; Ov 4 × 2 mm, cone-shaped; Sty slender, to 35 mm; Sti 1.5 × 1–2 mm, narrow, small, erect, lobes green.

T. mauryana L. B. Smith (Contr. Gray Herb. 117: 31, t. 2, figs. 32–33, 1937). Type: Mexico, Hidalgo (Maury 5747 [GH, F]). — Distr: C Mexico (Hidalgo); saxicolous, 1200–2000 m. – Fig. 21.

≡ Viridantha mauryana (L. B. Smith) Espejo (2002).

[1] Pants often vertically on steep rockwalls, stemless or short-stemmed, flowering 5–15 cm tall; Ros 4–11 × 5–14 cm, usually in groups; L densely rosulate, to 30, 5–10 (−12) cm, thick-fleshy, densely cinereous-lepidote, silver-grey; L sheath 1.5–2 × 0.8–1.2 cm at the base, triangular, grey to brown-lepidote, glabrous for the lower 2 mm, both faces light brown, inconspicuously offset from the lamina; L lamina narrowly triangular, 4–10 × 0.5–1 cm wide at the base, densely cinereous-lepidote, spreading or reflexed, flat, apex attenuate; Inf composed of 3–9 spikes, longer or shorter than the leaves; peduncle short and concealed by the leaves, 1–2 (−3.5) cm, with a collar of bladeless acute bracts below the inflorescence; lowest peduncular Bra leaf-like, the upper like the primary bracts, erect or bent outward, densely imbricate, (2–) 5–8 cm, narrow-triangular; primary Bra like the floral bracts, lanceolate, 2.5–3 × 1–1.5 cm, shorter then the spikes, membranous, acuminate, densely lepidote; spikes with 1 or 2 sterile bracts at the base, erect or divergent, to 30 × 10 (−15) mm, complanate and with slightly convex sides, 3- to 8-flowered, rachis hardly visible at anthesis; floral Bra densely imbricate, slightly exceeding the sepals, 13–20 (−25) × 10–15 mm, (triangular-) ovate, thin, faintly veined, finely to sharply carinate esp. at the apex, adaxially glabrous, abaxially cinereously lepidote with large grey spreading scales, roseate, acuminate; Fl subsessile, distichous, densely arranged; Sep 12–15 (−17) × 4–5 mm, narrowly ovate, membranous, evenly short-connate for 1 mm or subfree, alate, adaxial ones carinate, sparsely to subdensely lepidote, acute; Pet almost erect, linear, 14–24 × 3–4 mm (base 2–2.5 mm wide), dark emerald-green and yellow-green at the base, blade suberect or slightly divergent, rounded or obtuse; St all of equal length, 18–20 mm, included; Fil 12–14 mm, ribbon-like, thin and flaccid, white; Anth subbasifixed, 2.5–3 × 0.4 mm, linear, base bilobed, apex apiculate, yellow-olive to light brown; Ov 3 × 2 mm, ovoid, tapering into the style; Sty 11 mm, reaching or surpassing the anthers; Sti erect, slightly wider than the style, linear, lobes green.

T. mitlaensis W. Weber & Ehlers (Feddes Repert. 94: 617–618, ill., 1983). Type: Mexico, Oaxaca (Ehlers s.n. [Herb. Weber 283]). — Distr: S Mexico (Oaxaca); saxicolous, ±1800 m. – Fig. 22.

[5] Plant flowering ±16 cm tall; L suberect, the outer ones reduced, densely coarsely white-tomentose-lepidote; L sheath to 3 × 2 cm, ovate, indistinctly merging into the lamina; L lamina narrowly triangular, ± to strongly secund, finely veined, 7–9 cm, stout, canaliculate, apex subulate, subobtuse or acute, not pungent; Inf simple; peduncle suberect, short and concealed by to ± equalling the leaves, stout; peduncular Bra with short somewhat leaf-like blades, erect or slightly bent upwards, densely imbricate, enveloping the peduncle, densely lepidote; spike 7–12 × 1.6–1.8 cm, lanceolate, subterete to distinctly complanate, ±5-flowered, attenuately acute; floral Bra suberect, densely imbricate, much exceeding the sepals, 32–35 (−44) × 16 mm, ovate-lanceolate, submembranous, margins hyaline, strongly veined, ecarinate, densely concealing the rachis, on both faces (sub-) densely lepidote, reddish, obtuse or obscurely acuminate; Fl with a short pedicel-like receptacle, distichous; Sep 23–24 × 5 mm, lanceolate, membranous, with broad hyaline margins, veined, adaxial ones connate for 4–5 mm, abaxial one subfree, adaxial ones distinctly bluntly carinate, whitish-green, glabrous or abaxially sparsely lepidote with large trichomes, acute; Pet forming an erect narrow tube clasping the filaments at the apex, 50 × 7.5 mm, pale to dark violet, whitish below, blade tip margins recurved, rounded-emarginate; St in 2 whorls of unequal length, ± exserted; Fil 50–54 mm, slender and flattened at the base and dilated-terete at the distal end, white below, exserted part violet; Anth dorsifixed at 1/3 from the base, 2 mm, elliptic, brownish-green; Ov 5 mm, ovoid, contracted into the style; Sty ±55 mm, slender, slightly dilated upwards; Sti conduplicate-spiral, unequal, 2.5 mm, whitish-green.

T. mollis H. Hromadnik & W. Till (Pl. Syst. Evol. 142: 123–128, ills., 1983). Type: Bolivia, Tarija (Hromadnik 9088 [WU, B, NY, W, Herb. W. Till]). — Distr: S Bolivia (Tarija); saxicolous on S-facing cliffs, 1770–2700 m.

[4] Plant flowering (9–) 16–24 (−26) cm tall and 3.5 – 4 cm wide, forming dense clusters ±40–50 cm ∅; stem simple or few-branched, internodes (0.7–) 0.8–1 cm; L distichous, spreading, (2–) 3–5 (−6) cm, soft, flexible; L sheath amplexicaul, completely covering the stem and making it firm, 0.9–1.1 × 0.8–1.1 cm, broadly obovate, with translucent margins near the middle, margins connate to each other in the lower ½, ±9- to 10-veined, adaxially glabrous, abaxially densely lepidote with eccentric spreading scales, pale stramineous; L lamina subulate-terete, 1.9–4 × 0.3–0.5 cm wide at the base, silvery-white, densely lepidote, trichomes eccentric and spreading, apex subacuminate, abruptly flattened and obtuse; Inf 1-flowered, rachis thread-like extended above the flower but hidden between the bract; peduncle nearly none or very short, to 0.8 cm, soon laterally displaced after anthesis, densely lepidote; peduncular Bra 1, placed approximately beneath the floral bract, 1.2–1.3 cm, densely lepidote, short-caudate; floral Bra ± equalling the sepals, 6–9 × 2–5.5 mm, suborbicular, 9-veined, ecarinate, shortly laminate or obtuse-apiculate; Fl sessile, distinctly terminal; Sep (5–) 6–7.5 × 2.5–3.5 mm, (narrowly) ovate, thin-membranous, with overlapping broad hyaline margins, (3- to) 4- to 5-veined, the outer veins joined to the midvein near the apex, subfree, ecarinate, abaxially densely lepidote except the bottom 1/3, adaxially glabrous, rounded; Pet tongue-shaped, 10–15 × 2–2.5 mm, base 2 mm wide, dirty yellow-brown or olive-green, blade recurved, rounded-emarginate; St all of equal length; Fil 7 × 0.5 mm, flat and attenuate, flaccid, straight or rarely plicate, white; Anth basifixed, 1.8–2.5 × 0.5 mm, apex obtuse, greenish-yellow; Ov 1.5–2 × 1.5 mm, ovoid to shortly cylindrical, abruptly contracted into the style; Sty 3 mm, ±0.4 mm ∅; Sti short, lobes crested, capitellate, not wider than the style.

T. myosura Grisebach ex Baker (J. Bot. 16: 240, 1878). Type: Argentina, Córdoba (Lorentz 122 [BM, B, G, MO, NY, P]). — Distr: S Peru (Arequipa), Bolivia, Argentina, Uruguay; epiphytic in dry habitats, 700–3200 m.

≡ Tillandsia nappii Lorentz & Niederlein (1881) (nom. illeg., ICN Art. 52.1); incl. Tillandsia mandonii E. Morren ex Mez (1896).

[4] Plant flowering to 30 cm high or sometimes more, forming dense clusters; stem to 6 (−10) cm, simple, few-branched or richly branched at the base; L subdensely distichous, 5–17 cm, stiff; L sheaths densely imbricate and margins overlapping behind the stem for most of their length, making the stem to appear stout (5–8 mm ∅), 1–1.5 cm, suborbicular to reniform, normally with the upper ½ densely lepidote like the lamina or sometimes glabrous except a fringe of narrow marginal scales, distinct from the lamina; L lamina narrowly triangular or linear, 0.3–0.5 cm wide, densely cinereous-lepidote with small basally produced subappressed to reflexed pruinose scales, strongly recurved, becoming strongly sulcate when dry, ± contorted, subulate, with a furrow, apex acuminate and pungent; Inf simple; peduncle bractless or with 1 bract well removed from the flowers, erect, 4–20 cm, lepidote or glabrous; peduncular Bra lanceolate, involute, lepidote; spikes lax, to 8 cm but often much shorter, linear, (1- to) 2- to 5- (to 8-) flowered, rachis slender, flexuous in the few-flowered specimens or usually geniculate in the larger ones, ± lepidote; floral Bra slightly or not at all imbricate, the lower ones exceeding or equalling or the upper ones often distinctly shorter than the sepals, (13–) 17.5–30 × 9–9.5 mm, broadly ovate or ovate-lanceolate, clasping the flower, chartaceous, many-veined, not concealing the rachis except by their extreme base, rarely >2× as long as the internodes (but much less esp. in the many-flowered specimens), cinereously lepidote, acuminate with a short tip; Fl subsessile, closely appressed to the rachis, fragrant; Sep 11–13.5 × 5–5.8 mm, oblong-lanceolate, thin, many-veined, evenly short-connate, adaxial ones bluntly carinate, usually sparsely lepidote or glabrescent, rounded or obtuse; Pet linear to tongue-shaped, ±13 (−30) × 3 mm, at the base slightly narrowing, pale yellow; St deeply included, exceeding the style; Fil to 5.5 mm; Anth (sub-)basifixed, ±3.5 × 0.5 mm; Ov ±4 × 2 mm, subcylindrical, tapering into the style; Sty ± as long as or shorter than the ovary; Sti almost flat and disc-shaped (gemmate).

T. neglecta E. Pereira (Bradea 1: 78, t. 2, 1971). Type: Brazil, Rio de Janeiro (Sucre s.n. [HB]). — Distr: SE Brazil (Rio de Janeiro); saxicolous on granitic rock walls. – Fig. 23.

[2] Plant caulescent, flowering to 20 cm tall, often forming dense clusters; stem few-branched at the base, curved; L rosulate, somewhat curved, 4–7 cm, stiff and fleshy-coriaceous, subappressed-lepidote on both faces, green or tinged reddish or silver-grey at the base because of spreading trichomes; L sheaths 1.5 cm wide, whitish; L lamina narrowly triangular, not or slightly secund, subulate towards the apex, apex abruptly acute and pungent; Inf simple, mostly much exceeding the leaves; peduncle completely covered by its bracts or partly visible, erect, equalling or exceeding the leaves, glabrous, green; peduncular Bra erect, laxly imbricate, much (3–4× longer than) to slightly exceeding the internodes, ovate-oblong, sparsely pale-lepidote at the margins and apex, long-acuminate; spike 3–4 cm, corymbiform, 6- to 10-flowered; floral Bra equalling to much exceeding the sepals, 17 × 9.5 mm, ovate or elliptic, margins narrowly hyaline, darker veined, subglabrous or lepidote towards the apex (esp. the lower ones), green to reddish esp. at the margins, acutish or obscurely acuminate; Fl sessile, polystichous, suberect or divergent, densely arranged, 22–30 mm; Sep 15–16 × 3 mm, lanceolate, with broad hyaline margins, connate for 2–2.5 mm, ecarinate, glabrous, narrowly acute; Pet spatulate or tongue-shaped with long cuneate base, 22–30 × 5 mm (base 1.5 mm wide), blue, blade slightly divergent, rounded-emarginate; St 11–16 mm, included, visible in the throat but shorter than the style; Fil slender and narrow at the distal end or ribbon-like, towards the middle slightly plicate, 17.5 mm long; Anth basifixed, 2 × 0.5 mm, linear, base bilobed, apex obtuse; Ov 3–4 × 1.5 mm, trigonous or ellipsoid, the top 1/3 sterile, tapering into the style; Sty ±14 mm, slender; Sti erect, linear, unequal.

T. paleacea C. Presl (Reliq. Haenk. 1: 125, 1827). Type: Chile (Haenke s.n. [PR, GH [photo]]). — Distr: C Colombia (Cauca, Huila, Tolima), Peru (widespread), NE Bolivia (La Paz), N Chile, S Paraguay; on rocks and desert sands, or on trees in dry areas, 0–3900 m. – Fig. 24.

Incl. Tillandsia fusca Baker (1878); incl. Tillandsia scalarifolia Baker (1887); incl. Tillandsia chilensis Baker (1889); incl. Tillandsia schenckiana Wittmack (1889); incl. Tillandsia favillosa Mez (1906).

[3] Plant variable in habit, caulescent, flowering 10–70 cm tall, often much branched; stem 10–35 cm, appearing stout because of the densely imbricate leaf sheaths, brittle; L laxly spirally arranged, densely tomentose-lepidote, silvery-grey often becoming fuscous with age; L sheaths large, broadly ovate, elliptic, glabrous, the upper ½ cleaerly visible, outside lepidote; L lamina narrowly triangular, to 12 cm, 4–6 mm wide, abruptly spreading, usually slighly flexuous, for most part canaliculate to subulate in the upper part, apex attenuate; Inf simple; peduncle mostly completely covered by its bracts, erect, shorter than to much exceeding the leaves, to 15 cm or sometimes much more, slender, glabrous or nearly so; lowest peduncular Bra somewhat leaf-like with short lamina, soon erect, laxly imbricate, exceeding or about equalling the internodes, narrowly elliptic, tightly enveloping the peduncle, lepidote, apiculate; spike dense, to 5 cm, linear-lanceolate, complanate, (1- to) 2- to 5- (to 12-) flowered, acute, rachis angled, straight or flexuous, glabrous, partly exposed; floral Bra distichous, slightly shorter than or slightly exceeding the sepals, 13–16 × 6 mm, ovate or elliptic, finely veined, ecarinate, so narrow as to expose the rachis in part, ±2–3× as long as the internodes, soon glabrous or in the upper part subdensely lepidote, green, sometimes tinged reddish; Fl with a short pedicel-like receptacle; Sep 10–17 × 5 mm, elliptic or obovate-lanceolate, with hyaline margins, free, ecarinate but slightly fleshy at the base, tinged reddish-purple, glabrous; Pet claw narrow with suborbicular blade, 20–23 × 10 mm, margins slightly undulate, blade blue to violet or rose, claw white, corolla sometimes with a white eye; St 6–8 mm, deeply included; Anth basifixed, 1.5 mm; Ov subcylindrical, 2.5–3.5 mm, abruptly contracted into the style; Sty stout, 1.5 mm (incl. stigma); Sti disc-shaped, gemmate.

T. paraensis Mez (in Martius, Fl. Bras. 3(3): 586, t. 109, 1894). Type: Brazil, Pará (Sieber 68 [BR, GH [photo]]). — Distr: SE & S Colombia (Vaupés, Amazonas), E & S Venezuela (Bolívar, Amazonas), Guyana (Cuyuni-Mazaruni, Potaro-Siparuni, Upper Takutu-Upper Essequibo), E & S Suriname (Brokopondo, Sipaliwini), French Guiana (Saül, Régina, Maripasoula), Brazil (Amazonas, Mato Grosso, Rio Grande do Norte, Rondônia, Roraima, Pará, Rio de Janeiro, Bahia, Pernambuco), NE Ecuador, Peru, NE Bolivia; epiphytic in forest, 20–1120 m.

Incl. Vriesea sanctae-crucis S. Moore (1895) ≡ Tillandsia sanctae-crucis (S. Moore) Mez (1896); incl. Tillandsia juruana Ule (1907).

[5] Plant stemless, flowering 12–40 cm tall; L 10–25, forming a narrowly utriculate rosette, often exceeding the inflorescence or at least the peduncle, 10–35 cm, the outer greatly reduced and sheath-like, very densely subappressed-lepidote throughout; L sheaths forming an elongate pseudobulb, 3–7 × 1.7–3 cm, ovate, inflated-convex, stiff-coriaceous, with membranous margins, slightly distinct from and merging into the concolorous lamina; L lamina very narrowly triangular, succulent (fleshy) and coriaceous, 6–25 (−33) × 1.4–2 cm, suberect or curved, often finely keeled on both faces, often the margins involute towards the apex when dry, apex attenuate and abruptly pungent; Inf simple (or rarely compound of 2 spikes), 10–30 (−40) cm; peduncle completely covered by its bracts or partly visible, often curved, short and concealed by the leaves or elongate, 8–19 cm, 2–3.5 mm ∅, sparsely lepidote or subglabrous; lowest peduncular Bra leaf-like, erect, densely imbricate to remote, ovate, chartaceous, (densely) appressed-lepidote or floccose at least at the apex, the upper ones more apiculate; spikes mostly curved, 6–15 (−20) cm, 1–1.5 cm wide, linear (-lanceolate), complanate, subdensely distichously 4- to 15-flowered, acute, rarely with >1 sterile bract at the apex, rachis sharply angled when dry, ± flexuous, sparsely lepidote or glabrate, only partly hidden at anthesis but often wholly exposed when dry; floral Bra scarcely imbricate (if at all), equalling to much exceeding the sepals, 23–39 mm, (broadly) elliptic, clasping the flower, scarcely incurved, subchartaceous, even or veined, ecarinate, so narrow as to expose the rachis in part, <2–4× as long as the internodes, densely lepidote or subglabrous, often yellow-green or tinged with red or wholly reddish, obtuse or the lower ones minutely apiculate; Fl subsessile, erect or suberect; Sep slightly incurved at the apex, 15–23 mm, ovate-oblong, thin-coriaceous, veined only when dry, free, sparsely or minutely lepidote, rounded, then obtuse or subacute; Pet erect with slightly divergent apex and forming a tube, linear-lanceolate, 33–65 mm, rose to pink, attenuate and obtuse; St unequal in length, 40–45 mm, exserted or ± equalling the petals, shorter then or equalling the pistil; Fil flat or subterete towards the apex, straight; Anth dorsifixed just below the middle; Ov slenderly ovoid, gradually contracted into the slender style.

T. paucifolia Baker (Gard. Chron., ser. nov. 10: 748, 1878). Type: K, GH [photo]. — Distr: SE USA (Florida), SE Mexico, N & E Guatemala, C & S Honduras, E El Salvador, Nicaragua, NW & W Costa Rica, Bahamas, Cuba, Jamaica, Haiti, Dominican Republic, N Colombia, C & NE Venezuela; epiphytic in low forest and scrub.

Incl. Tillandsia yucatana Baker (1887).

[5] Plant stemless, flowering 10–45 cm tall, with few offsets; outer L reduced, without lamina, other leaves densely appressedly cinereous-lepidote; L sheaths forming an elongate pseudobulb 5–15 cm long, large, broadly ovate, merging gradually into the lamina; L lamina thick, to 20 cm but often much shorter even in large plants, 3–7 mm wide at the base, curved, involute-subulate, the lower often contorted, apex attenuate and abruptly pungent; Inf simple, digitate (or slightly spaced), composed of (1–) 3–6 spikes; peduncle erect, very short and concealed by the leaves; peduncular Bra leaf-like with spreading lamina, erect, imbricate; primary Bra like the peduncular bracts, leaf-like, at least the lower ones much exceeding the spikes with their lamina, rose; spikes suberect or divergent and often curved, to 12 cm but normally much smaller, linear-lanceolate, complanate, 2- to 10-flowered, acute; floral Bra erect, imbricate, exceeding or slightly shorter than the sepals, 20–21 (−30) × 11 mm, elliptic or ovate, subchartaceous, margins hyaline, veined when dry, the apex scarcely or not at all carinate, concealing the rachis (alive), 3–4× as long as the internodes, appressed-lepidote, acute or obtuse; Fl sessile, distichous; Sep 18–20 × 5 mm, lanceolate-oblong or narrowly obovate, subchartaceous, finely veined when dry, unequally short-connate (adaxial ones more) for 2–4 mm, adaxial ones carinate in the lower 2/3, glabrous or abaxially sparsely lepidote at the base centre, rounded, then obtuse (or subacute); Pet forming an erect narrow tube clasping the filaments at the apex, linearly tongue-shaped, to 43 × 7 mm, basally narrowed to 3 mm, violet, blade tip margins recurving, rounded; St in 2 whorls of unequal length, ±45 mm, exserted; Fil exposed part slightly inflated and (sub-) terete, violet in the upper part; Anth dorsifixed near the middle, 2.5 × 1 mm, elliptic, apex obtuse, dark brown; Ov 6 × 2.5 mm, narrowly ovoid, contracted into the style; Sty 54 mm, exceeding the stamens; Sti dilated, spreading, white.

T. pedicellata (Mez) A. Castellanos (Lilloa 11: 144, 1945). Type: Argentina, Córdoba (Hieronymus 795 p.p. [B, BM, CORD, GH]). — Distr: Cuba (Camagüey), S Brazil (Paraná), Bolivia (Chuquisaca, Cochabamba, La Paz, Potosí, Santa Cruz, Tarija), Argentina (Buenos Aires, Catamarca, Córdoba, Jujuy, Mendoza, Salta, San Juan, San Luis, La Pampa, Rio Negro, Tucumán); epiphytic, 60–3660 m. – Fig. 25.

≡ Tillandsia coarctata var. pedicellata Mez (1896) (nom. inval., ICN Art. 35.1); incl. Tillandsia coarctata Gillies ex Baker (1878) (nom. inval., ICN Art. 32.1c).

[4] Plant small, flowering to ±6 cm tall, forming dense clusters; L rosulate, densely imbricate, to 8 × ±1 mm; L sheaths suborbicular, with broad veinless margins, 3- to 4-veined, glabrous; L lamina lanceolate in outline, densely lepidote with green-centred trichomes, fleshy, subulate, apex apiculate or obtuse; Inf peduncle at flowering time very short (0.3 cm) or almost missing to elongate at fruiting time, glabrous; floral Bra (5.5–) 6.5–7 × (2.6–) 3–3.3 mm, ovate to broadly triangular, 1- to 3-veined, glabrous or sparsely lepidote towards the apex with large trichomes, apiculate; Fl erect, slighly zygomorphic with 2 petals spreading, sometimes barely exceeding the leaves, weakly fragrant; Sep 5.8–7 × 1.3–2 mm, lanceolate, 3- (to 5-) veined, outer vein much shorter than the midvein, equally subfree or adaxial ones connate for ±½ their length or all connate for up to 3 mm, glabrous, apiculate or obtuse; Pet tongue-shaped, often widest at ± the middle, 8–9 × 1.3–1.8 mm, (4- to) 5- to 6-veined, pale yellow to dark violet or brown, broadly rounded; St all of equal length, 3.5 mm, deeply included; Fil 2.7–3 × 0.15 mm, thread-like, 1-veined, whitish; Anth basifixed, 1–1.5 × 0.25–0.3 mm; Ov 1.5 mm, cylindrical or subglobose, abruptly contracted into the style; Sty ±1/3 as wide as the ovary, 1/3 as long as the ovary; Sti disc-shaped, gemmate.

T. peiranoi A. Castellanos (Lilloa 2: 14, t. 2, 1938). Type: Argentina, Salta (Peirano s.n. [LIL, BA, GH [photo]]). — Distr: NE Argentina (Salta); saxicolous, 1200 m.

[3] Plant caulescent, flowering 20 cm tall but normally much smaller, few-branched; stem to 10 cm; L subdensely spirally arranged, 2–5 cm, cinereously-lepidote with subappressed scales; L sheaths large but obscure, broadly ovate, with broad thin margins, many-veined, merging into the lamina; L lamina narrowly triangular or triangular, rigid and fleshy, 0.5 cm wide, suberect to recurved, veined, with subulate apex, strongly canaliculate except at the attenuate and pungent apex; Inf simple or 1× branched with 1–2 small suberect branches, composed of 1–3 (−5) spikes; peduncle completely covered by its bracts, erect, very slender, glabrous; peduncular Bra laxly imbricate, lanceolate, thin, veined, lepidote, acuminate; primary Bra like the floral bracts, much shorter than the spikes; spikes to 6 cm, linear (at least the terminal one), ±6-flowered, rachis slender, flexuous or suberect, glabrous; floral Bra erect, slightly shorter than the sepals, 12 mm, elliptic or ovate, closely enveloping the flowers, thin, veined, ecarinate, so narrow as to expose the rachis in part and sometimes part of the sepals, slightly <2× as long as the internodes, glabrous, green or purple-brown or soon dry and stramineous; Fl distichous, suberect or divergent caused by the flexuous rachis, always partly contiguous with the rachis; Sep 10–12 × 3 mm, elliptic, thin, centrally (3– to 5–) veined with broad veinless margins, free, adaxial ones carinate, tinged reddish-purple, glabrous, sharply acute; Pet claw narrow with suborbicular blade, 16 × to >7 mm, pale blue to nearly white, blade recurved. St deeply included, exceeding the style; Anth 2.5 mm, linear.

T. porongoensis L. Hromadnik & P. Schneider (Haussknechtia 4: 39–41, 1988). Type: Argentina, La Rioja (Hromadnik & Hromadnik s.n. [WU]). — Distr: NW Argentina (La Rioja); epiphytic, ±850 m.

[4] Plant caulescent, stem arched, hardly branched; L few, forming a spreading open rosette, laxly rosulate, to 15 cm, stiff and succulent, densely cinereous-lepidote, silver-white; L sheaths 1.2–1.5 × 1.5–2 cm, triangular to ovate, clasping the stem and concealing it, adaxially smooth, glabrous towards the base to densely lepidote in the upper ½, cherry-red up to ½ the length, abaxial face paler with spots, distinct from the lamina; L lamina very narrowly triangular, near the sheath with extending trichomes at the margins, to 12 × 0.8–1 cm wide at the base, spreading to recurved into a semicircle, margins strongly involute, canaliculate and subulate from ± the middle to the apex, apex acute and pungent with a sharp point; Inf simple; peduncle completely covered by its bracts, 4.5–5 cm, 1.5 mm ∅, glabrous; peduncular Bra ± equalling the internodes, 3.2 × 0.9 cm, the upper ones completely enveloping the peduncle, with only the tip flaring slightly, distinctly veined, silvery cinereous-lepidote, stramineous, the lower ones with leaf-like blade and more apiculate; spike 4–4.5 cm, 7–9 mm wide, lanceolate, complanate, 4- to 8-flowered, rachis 4-angled, geniculate, glabrous, hidden or partly exposed, internodes 4 mm; floral Bra ± imbricate, exceeding or much shorter than the sepals, upper ones very short, (10–) 16–20 × 10 mm, coriaceous, veined, ecarinate, concealing the rachis or so narrow as to expose the rachis in part, adaxially glabrous, the lower ones densely lepidote or the upper ones lepidote at the margins with large trichomes, tinted reddish-brown, apiculate; Fl distichous, erect; Sep 12–14 × 4 mm, narrowly ovate, coriaceous, soon straw-like, veined, evenly shortly connate, adaxial ones slightly carinate, reddish-brown, adaxially glabrous, abaxially glabrous in the lower ½ and densely grey-scaled in the upper ½, apiculate; Pet 20 × 5 mm, 2–3 mm wide at the base, yellowish-brown, blade ovate, recurved, rounded; St deeply included; Fil 10 × 0.3 mm, ribbon like, straight, white; Anth 1.5 mm; Ov 4 × 2.5 mm, slenderly ovoid; Sty stout, somewhat thinner than the ovary, 8 mm including the stigma.

T. praschekii Ehlers & K. Willinger (Bromelie 1989(2): 36, ill., 1989). Type: Cuba, Pinar del Río (Prášek 32/83 [WU]). — Distr: Cuba (Pinar del Río); on rocks, 200–300 m. – Fig. 26.

[5] Plant stemless, flowering 8–10 cm tall, growing in large groups; L to 15, forming an almost bulbous rosette, succulent; L sheaths 1.5 × 1.5 cm, suborbicular to elliptic, inflated, with small asymmetrical trichomes on the margins, densely furfuraceous-lepidote, outer face dark green and brownish at the base, inner face brown, abruptly contracted into the lamina; L lamina narrowly triangular, to 7 cm, 6 mm wide near the base, cinereous-green, densely and furfuraceously lepidote with small spreading scales, divergent to strongly recurved, the edges grooved, canaliculate, apex attenuate and then abruptly acute; Inf 1× branched, appearing simple but composed of 10 polystichously arranged hidden spikes, erect or almost upright, dense, capitate, to 4 × 3 cm,the uppermost 2–5 spikes at the apex of the inflorescence remaining undeveloped; peduncle completely covered by its bracts, short and concealed by the leaves, ±2 cm, 5 mm ∅; peduncular Bra leaf-like; primary Bra spreading to recurving, like the peduncular bracts, blade narrowly triangular, the lower ones 4 cm long, upper ones shorter, much exceeding and wholly hiding the spikes, subsucculent, densely furfuraceous-lepidote, leaf-like and almost forming an involucrum, green to pink; spikes shortly stipitate, with sterile bracts at the base, erect and appressed to the axis, excluding the petals 2 cm, 8–10 mm wide, complanate, 1- to 3-flowered (upper spikes 1-flowerd), with a reduced flower at the apex; floral Bra 3–4 mm shorter than the sepals, 7–9 × 7 mm at the base, broadly triangular, rigid and coriaceous, margins hyaline, strongly veined, alate-carinate, inside sparsely lepidote, outside furfuraceously lepidote, green with red hooked tip; Fl with a short pedicel-like receptacle; Sep 12 × 5 mm, elliptic, rigid, with hyaline margins, evenly shortly connate for 1 mm, adaxial ones carinate, at the base green, otherwise wine-red, sparsely lepidote with large trichomes, acuminate; Pet erect and forming a tube, tongue-shaped, 36 × 6 mm (base 3 mm wide), violet-blue with white base, blade tip margins divergent; St in 2 whorls of unequal length, to 45 mm, exserted for 10 mm; Fil slender and flattened at the base and dilated and becoming terete at the distal end, violet with white base; Anth dorsifixed near the middle or at 1/3 from the base, 2 × 0.75 mm, elliptic, brown; Ov 5 × 3 mm, narrowly ovoid, angled; Sty slender, exceeding the anthers; Sti 2 × 2 mm, erect or lobes spreading, papillose, white.

T. pruinosa Swartz (Fl. Ind. Occid. 1: 594, 1797). Type: Jamaica (Swartz s.n. [S]). — Distr: SE USA (Florida), SE Mexico, C & S Belize, N Guatemala, Honduras, Nicaragua, Costa Rica, Cuba, Jamaica, Dominican Republic, Puerto Rico, Colombia, Venezuela, Guyana, Suriname, Brazil, Ecuador; epiphytic, 0–1400 m.

≡ Platystachys pruinosa (Swartz) Beer (1856); incl. Tillandsia breviscapa A. Richard (1850).

[5] Plant stemless, flowering 8–12 (−23) cm tall; L ±10–20, 7–25 cm, white-tomentose-lepidote (often ferrugineous when dry); L sheaths forming an ovoid pseudobulb, 2–4.8 × 2–4.3 cm, suborbicular or elliptic, much inflated-convex, papyraceous, with broad membranous purple-red margins, densely ferruginous-lepidote to somewhat ferrugineously tomentose-lepidote at the base, inner face pale brown to deep purple, abruptly contracted into the lamina; L lamina fleshy-coriaceous and flexible, 5–21 cm, 2–5 mm wide, secund or suberect, often slightly flexuous, very densely white- to somewhat ferrugineous-lepidote to tomentose-lepidote, involute-subulate, apex filiform-attenuate; Inf simple or rarely digitately compound of 2–4 spikes, 5 (−15) cm, densely tomentose-lepidote; peduncle densely covered by its bracts and concealed within the leafy rosette, erect, 5–8 cm, 2–3 mm ∅; peduncular Bra leaf-like, suberect, with densely imbricate sheaths, the upper ones more oblong, acute; axes of the inflorescence hidden, very short; primary Bra like the upper peduncular bracts, subspreading with the spikes, exceeding the spikes with the leaf-like lamina, shorter than the axillary branches; spikes shortly stipitate (<1 cm), with a sterile bract (prophyllate) or with few sterile bracts at the base and apex, suberect or divergent when compound, dense, 4–6.5 cm, 1.8–4.5 cm wide, (broadly) elliptic, slighly complanate, 3- to 6- (to 15-) flowered, acute, rachis angled, nearly straight, densely tomentose-lepidote, hidden; floral Bra suberect, densely imbricate, nearly 2× as long as the sepals, 20–29 mm, elliptic, thin-coriaceous, even, obscurely carinate towards the apex, 4–5× as long as the internodes, pale silvery green tinged with or wholly dull roseate-red, acutish or obtuse and obscurely apiculate; Fl sessile, distichous, contiguous with each other and/or the rachis; Sep 13–19 mm, elliptic, very thin-coriaceous, with broad membranous margins, even, abaxial one subfree, adaxial ones about ½ connate for ±6 mm, adaxial ones bluntly carinate at the base, sparsely appressed-lepidote on both faces or glabrescent, rounded, then obscurely acuminate; Pet erect and forming a tube, tongue-shaped, 30 mm, violet, blade tip margins recurved, obtuse or acute; St in 2 whorls of unequal length, long-exserted, the shorter equalling and the longer exceeding the style; Fil slender and flattened at the base and dilated and becoming terete at the distal end, violet; Anth dorsifixed at 1/3 from the base, >3 mm; Ov ovoid (?), contracted into the style; Sty slender; Sti conduplicate-spiralized, linear.

T. pseudobaileyi C. S. Gardner (Selbyana 7(2–4): 363, ill., 1984). Type: Mexico, Chiapas (Gardner 1118 [SEL]). — Distr: S Mexico, C & SE Guatemala, SW & S Honduras, El Salvador, W & C Nicaragua; epiphytic on trees of open seasonally dry forests, 100–1500 m.

[5] Plant stemless, flowering to 30 cm tall, with few offsets; L 10–12, the outer greatly reduced, the others 20–35 cm; L sheaths forming an ovoid pseudobulb, 4–5 × 5–6 cm, orbicular, much inflated, stiff-coriaceous, with reddish margins, veined esp. in the upper ½, adaxially red-brown-lepidote and abaxially appressed-lepidote, brownish towards the base; L lamina narrowly triangular or linear, succulent and rigid, 1–1.5 cm wide, minutely appressed-lepidote, with striking longitudinal brown-red stripes (veins) in bright light, curved or divergent, distinctly veined (fresh and dry), the margins involute and often contorted; Inf compound, composed of 2–5 spikes; peduncle erect, short and concealed by the leaves or slighly elongate, to 15 cm, blood-red; peduncular Bra leaf-like, laxly imbricate, involute, the sheath blood-red, the lamina green or reddish; primary Bra lanceolate, the lowest ±½ as long as the spikes, acuminate and pungent, blood-red; spikes with a sterile bract, bicarinate at the base, spreading at 40–45°, linear-lanceolate, complanate, 4- to 6-flowered; floral Bra slightly exceeding the sepals, 17–20 × 9–11 mm, ovate, convex, rigid, scarcely or not at all carinate at the apex, the adaxial face glabrous with a few trichomes near the apex, abaxially appressed-lepidote, cinereous, tinged red or with reddish margins; Fl suberect, contiguous with the rachis and each other; Sep 14–16 × 5–6 mm, elliptic, thin or rigid, evenly shortly connate or free, adaxial ones slightly to moderately carinate, green with red edges, adaxially glabrous and abaxially appressed-lepidote; Pet erect and forming a tube, spatulate, 29–32 × 6–8 mm, margins slightly undulate, lavender, blade tip margins recurving; St in 2 whorls of unequal length, exserted, the longest just exceeding the style; Fil 36–43 mm, broadened towards the apex, upper part lavender; Anth dorsifixed at 1/3 from the base or near the middle, 2–3 mm, black; Ov not described; Sty 29–34 mm; Sti lobes erect or spreading, dilated, slightly papillose, green.

Material separated as this species was treated as part of T. baileyi by Smith & Downs (1977).

T. pueblensis L. B. Smith (Contr. Gray Herb. 104: 81, t. 3, figs. 1–2, 1934). Type: Mexico, Puebla (Purpus 5856 [GH, BM, F, MO, NY, UC, US]). — Distr: E & S Mexico (Puebla, Oaxaca); saxicolous, ±1680 m. – Fig. 27.

[5] Plant stemless or short-stemmed or with a rhizomatous stem, flowering 16–24 cm tall, stem sometimes few-branched; L ±15, densely rosulate, erect or curved in the upper part, the outer ones reduced to pointed sheaths, the inner to 16 cm, densely tomentose-lepidote, silver-grey; L sheaths scarcely distinct from the lamina; L lamina narrowly triangular, brittle and fleshy, ±1 cm wide at the base, involute-subulate or stongly convex, apex acute or subpungent; Inf simple; peduncle completely covered by its bracts, erect, short and concealed by to ± equalling the leaves; lowest peduncular Bra somewhat leaf-like and long-laminate, the upper ones lanceolate, membranous, cinereous-lepidote, grey-green, acute; spikes erect, densely flowered, 8–9 × 1 cm, linear-lanceolate in outline, slightly complanate, typically 4- (to 7-) flowered; floral Bra erect, imbricate, much exceeding the sepals, 25–33 × 10 mm, oblong-lanceolate, convex, membranous, margins hyaline, even or finely veined when dry, ecarinate, concealing the rachis (alive), ±3× as long as the internodes, the base glabrous and subdensely lepidote towards the apex and margins, bright roseate or more often wine-red with green margins, narrowly rounded or obtuse; Fl subsessile, distichous; Sep 20–21 × 5.5 mm, narrowly obovate, chartaceous, with broad hyaline margins, veined at least when dry, adaxial ones connate or subfree, connate for up to (if at all) 6 mm, adaxial ones carinate, broadly obtuse or narrowly rounded; Pet forming an erect narrow tube clasping the filaments at the apex, tongue-shaped with long-cuneate base, 40–50 × 8.5 mm (base 3 mm), violet, blade tip margins recurved, rounded and then obtuse; St in 2 whorls of unequal length, ±60 mm, exserted; Fil slender and flattened at the base and dilated, becoming terete at the distal end; Anth dorsifixed near the middle, 2.5 × 1 mm, apex obtuse; Ov 5.5 × 2.5 mm, ovoid, tapering into the style; Sty slender, 62 mm, exceeding the stamens; Sti spreading, dilated.

T. rectangula Baker (J. Bot. 16: 238, 1878). Type: Argentina, Córdoba (Lorentz 127 p.p. [K, G, WU]). — Distr: C & S Bolivia (Cochabamba, Tarija), Argentina (Catamarca, Córdoba, La Rioja, San Luis, Santiago del Estero); epiphytic, 500–2720 m.

Incl. Tillandsia propinqua var. rectangula Grisebach (1879).

[4] Plant small, caulescent, forming a globose mass, stems many from a single point, richly branched at the base; L losely polystichous, rarely >2 cm; L sheaths densely imbricate making the stem appear stout, ±0.6 cm, suborbicular, scarious, with broad veinless margins, several-veined, glabrous except for the extreme apex, distinct from the lamina; L lamina narrowly triangular, 0.2 cm wide, densely cinereous-lepidote, divergent or spreading from the stem at ±90°, with a central keel, with small nearly symmetrical subappressed trichomes, subulate and strongly angled, with a furrow not reaching the apex, apex attenuate and pungent; Inf 1-flowered; peduncle naked for most of its length, erect to ascending, apparently always terminal, almost none or elongate to 2 (−4) cm, slender, strongly angled, glabrous; peduncular Bra usually 2, one at the extreme base of the peduncle, mostly hidden by the leaves, the other just below the flower, lanceolate, subcoriaceous, several-veined, even, glabrous or occasionally pale-appressed-lepidote, acute or obtuse; floral Bra much shorter than the sepals, 8 × 4 mm, like the upper peduncular bract but smaller, hyaline, glabrous, pale green, obtuse; Fl subsessile; Sep 9–10 × 4 mm, narrowly elliptic, membranous, with broad hyaline margins, even, evenly short-connate or slightly longer connate posteriorly for up to 2 mm, pale green, glabrous or sparsely lepidote, acute, obtuse; Pet tongue-shaped, 13 × 3 mm, yellow to ochre-brown, blade spreading or recurved at anthesis, obtuse; St 6 mm, deeply included, exceeding the style with most of the anthers; Anth dorsifixed near the base, >1 mm, linear, apex obtuse; Ov subprismatic, ±2.5 mm, contracted into the style; Sty slender, shorter than the ovary; Sti capitellate.

T. reducta L. B. Smith (Phytologia 5: 399, t. 1, figs. 11–12, 1956). Type: Peru, Cajamarca (Ferreyra 3216 [US, USM]). — Distr: NE Peru (Cajamarca); only on rocks in higher regions, 2500–2600 m; only known from the type.

[1] Plant caulescent, forming dense clusters, stem >8 cm, 0.5 cm ∅ but appearing stout (0.8 cm ∅) because of the densely imbricate leaf sheaths; L persistent, densely tomentose-lepidote with fine linear spreading trichomes; L sheath 1.2–2 × 1.2–1.8 cm, broadly ovate, thin at the base and margins; L lamina very narrowly triangular, fleshy, spreading or slightly recurved, 4–5 (−8) cm, subulate but adaxially flat or with a weak furrow; Inf depauperately compound, subdigitate, composed of 3–5 spikes, dense, ellipsoid, fertile part 2–3 × 1 cm; peduncle completely covered by its bracts, erect, 2–10 cm, 2 mm ∅, slender, lepidote at least near the nodes and in the upper part; lowest peduncular Bra almost leaflike with short lamina, the upper like the primary bracts, erect and with tightly inrolled margins, imbricate, sheath rose, axes of the inflorescence hidden and very short; primary Bra dense, suberect, ovate, to 2.2 cm, nearly as long as the spikes and covering them, thin, strongly veined and carinate, acuminate, abaxially lepidote; spikes with 1 (−2) sterile bracts at the base, (1- to) 2- to 3-flowered (top-most spike to 5-flowered), with a reduced flower at the apex, rachis stout and short, densely lepidote, internodes 2 mm; floral Bra suberect, ± equalling the sepals, 11–13 × 7–8 mm, ovate or triangular-ovate, incurved, closely enveloping the flowers, chartaceous, veined, scarcely or not at all carinate, so narrow as to expose the rachis in part, sparsely lepidote with few coarse white subappressed trichomes, roseate, acute or apiculate; Fl subsessile, distichous; Sep 9–11 × 5 mm, oblong, thin, somewhat veined, evenly shortly connate or subfree, adaxial ones carinate (faintly when fresh), rose in the upper part, glabrous, sometimes with a few trichomes at the base, broadly acute; Pet barely exceeding the stamens, tongue-shaped, 15–21 × 3–4 mm (base 1.5 mm), centre violet, apical part white (often turning purple after anthesis), blade slightly divergent and then recurved, rounded-emarginate; St 15–17 mm; Fil narrowly ribbon-like, straight; Anth dorsifixed near the base, 3–3.5 × 0.7 mm, lanceolate, grey-brown; Ov 3–4 × 1.5 mm, narrowly ovoid or subglobose, contracted into the style; Sty slender, 8–9 mm, slightly longer or shorter than the stamens; Sti spreading, shortly lobed.

T. reichenbachii Baker (Handb. Bromel., 166, 1889). Type: K [icono, Reichenbach drawing]. — Distr: SE Brazil (Rio de Janeiro), Bolivia (Santa Cruz, Tarija, Cochabamba, La Paz, Chuquisaca), N Argentina, Paraguay; epiphytic in woods, 100–2500 m.

≡ Tillandsia duratii ssp. reichenbachii (Baker) Halda (2005); incl. Tillandsia tucumanensis Mez (1896); incl. Tillandsia herzogii Wittmack (1916); incl. Tillandsia euosma Spegazzini (1917).

[3] Plant caulescent, flowering to >20 cm tall, stem 2–5 cm, stout, curved; L laxly spirally arranged, to 14 cm; L sheaths suborbicular or very broadly elliptic, lustrous, glabrous except for the lepidote apex, very distinct from the lamina; L lamina very narrowly triangular, thick, to 8 cm, 0.8 cm wide, cinereous-lepidote, spreading to reflexed and often recurved at the apex, strongly canaliculate (involute-subulate when dry), slightly flexuous; Inf subdigitate or sometimes simple, composed of 2–8 spikes, to 7 × 3–4.5 cm; peduncle completely covered by its bracts, erect, ±15 cm, 2.5 mm ∅, slender; lowest peduncular Bra somewhat leaf-like but soon with much reduced lamina, imbricate, sublanceolate, enveloping the peduncle, veined, lepidote, subacute; primary Bra like the upper peduncular bracts, much shorter than the stipe of the axillary branches, obtuse; spikes long-stipitate, stipe to 3 cm and densely bracteate, first often erect and then suddenly divergent, densely flowered, 3 cm, 0.9–1 cm wide, linear, complanate, 5- to 7-flowered; floral Bra much shorter than the sepals, 10–11 × 5 mm, elliptic, clasping the flower, subchartaceous, veined when dry, ecarinate, exposing most of the rachis, 3× as long as the internodes, subglabrous or sparsely lepidote, green tinged with reddish-brown, obtuse or acutish; Fl subsessile, slightly divergent or contiguous with each other and/or the rachis; Sep 12–12.5 × 6 mm, elliptic or obovate, thin-coriaceous, with broad hyaline margins, even, adaxial ones or all evenly shortly connate for 1–2 mm, ecarinate, green tinged with reddish-brown, glabrous, rounded or obtuse; Pet with narrow claw and suborbicular blade, 23 × 13 mm (base 2.5 mm wide), white or the blade often pale blue, margins crenate-serrate, blade spreading at right angle; St ± of equal length, 8 mm, deeply included; Fil ribbon-like, straight; Anth basifixed, 1.5 mm, greenish; Ov ±2.5 mm; Sty stout, ± as long as the ovary, shorter than the stamens; Sti disc-shaped, gemmate.

T. retorta Grisebach ex Baker (J. Bot. 16: 238, 1878). Type: Argentina, Córdoba (Lorentz 125 [BM, B, GOET, M, P]). — Distr: N and C Argentina (widespread to Mendoza and La Pampa in the S); epiphytic, 600–1460 m.

Incl. Tillandsia caespitosa Gillies ex Baker (1878); incl. Tillandsia nappii var. darwinii Lorentz & Niederlein (1881) (incorrect name, ICN Art. 11.4).

[4] Plant flowering usually 7.5 or rarely to 15 cm tall, forming a globose mass, stem 2–8 cm, many from a single point, usually much branched; L distichous, 3–5 (−7) cm, densely cinereous-lepidote; L sheaths imbricate, suborbicular, making the stem appear 3–5 mm thick, with broad veinless margins, veined, normally glabrous with a ciliate margin of elongate scales or occasionally in the upper part lepidote like the lamina; L lamina barely over 2 mm wide, recurved, then often towards the apex spreading again, veined when dry, subulate, apex acuminate and pungent; Inf 1- or 2-flowered, densely cinereous-lepidote; peduncle bractless, from almost none to much surpassing the leaves but usually quite short, slender, densely lepidote, reduced rachis extended from the base of the terminal flower; floral Bra erect, equalling or exceeding the sepals, 10–12 (−15) × 7 mm, triangular-ovate, veined, slightly >2× as long as the internodes, densely lepidote, acute, acuminate; Fl subsessile, erect; Sep 10–10.5 × 3 mm, lanceolate-oblong or narrowly obovate, thin, with broad hyaline margins, finely veined, adaxial ones connate for 3 mm, abaxial one almost free, adaxial ones carinate, sparsely lepidote, acute or obtuse; Pet erect with slightly divergent apex, tongue-shaped, 12.5 × 3.3 mm (base 1.3 mm wide), yellow, blade distinct, narrowly elliptic, rounded-emarginate, subtruncate; St 7 mm, deeply included, exceeding the style by ½ of the anthers; Fil ribbon-like and tapering from base to apex, straight; Anth basifixed, 2 mm, linear; Ov 3.5 × 5 mm, ovoid, abruptly contracted into the style; Sty stout, 2.5 mm; Sti erect, shortly lobed.

T. schatzlii Rauh (Trop. subtrop. Pfl.-welt 27: 8, 1979). Type: Mexico, Oaxaca (Rutschmann s.n. in BG Heidelberg 46304 [HEID, US]). — Distr: S Mexico (Oaxaca); steep rock walls; altitude not recorded.

[5] Plant stemless or sometimes with a rhizomatous stem; flowering Ros 25 × 12 cm; L ±10, forming a small funnel-shaped rosette, densely appressed-lepidote with trichomes with a green centre, outer leaves reduced and deltoid; L sheaths 4 × 3 cm, broadly ovate, wrinkled, light brownish to violet inside, inconspicuous from and merging into the lamina; L lamina narrowly triangular, slightly succulent, to 20 cm, 2 mm wide at the base, divergent and often the upper ½ spreading, apex attenuate and subpungent; Inf simple, often subpendulous or erect; peduncle completely covered by its bracts, erect or curved, shorter than the leaves, 5 mm ∅; lower peduncular Bra with short leaf-like blade but soon bladeless, raspberry-red or rose but colour masked by the trichomes; spikes to 16 × 2 cm, lanceolate, slightly complanate, 8- to 10-flowered; floral Bra imbricate, much exceeding the sepals, 47–60 × 20 mm, (narrowly) ovate, margins with extended scales, finely veined when dry, ecarinate, densely lepidote, raspberry-red or rosy yellow-green towards the base; Fl to 80 mm (incl. stamens and style); Sep 21.5–27 (−29) × 6–7 mm, narrowly obovate, finely veined when dry, adaxial ones connate for 3–4 mm, adaxial ones bluntly carinate at the base, green, sparsely lepidote with large trichomes, obtuse or acute; Pet forming an erect narrow tube clasping the filaments at the apex, narrowly tongue-shaped with long cuneate base, 65–80 × 9 mm (base 2.5 mm wide), dark violet fading to the white base, blade tip margins recurving; St in 2 whorls of unequal length, 70–85 mm, exserted, shorter than the style; Fil slender and flattened at the base and dilated becoming terete at the distal end, violet in the upper part; Anth dorsifixed, 3 mm, apex obtuse; Ov 5 × 2.5 mm, ovoid, tapering or gradually contracted into the style; Sty slender, exceeding the petal tube.

T. schiedeana Steudel (Nomencl. Bot. ed. 2, 2: 688, 1841). Type: Mexico, Veracruz (Schiede & Deppe 1004 [B, BM, HAL]). — Distr: USA (Missouri), Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Cuba, Jamaica, Haiti, Dominican Republic, N & C Colombia, Venezuela, NW Peru; epiphytic, 0–2250 m.

≡ Tillandsia vestita Schlechtendal & Chamisso (1831) (nom. illeg., ICN Art. 53.1); incl. Tillandsia flavescens Martens & Galeotti (1843); incl. Tillandsia caerulea Grisebach (1865) (nom. illeg., ICN Art. 53.1); incl. Tillandsia grisebachii Baker (1887); incl. Tillandsia eggersii Baker (1889).

[5] Plant caulescent, flowering usually ±20 (−40) cm tall, forming a globose mass, stem 5–20 cm, simple or few-branched; L polystichously few-ranked, varying greatly in density, to 25 cm, densely cinereous-lepidote or sometimes ferruginous-lepidote; L sheaths large, suborbicular, densely imbricate; L lamina very narrowly triangular, divergent to spreading at a right angle from the stem, adaxially often with 1 or few grooves, subulate (involute when dry) and angled, shallowly canaliculate, slightly flexuous, apex filiform-attenuate; Inf always simple; peduncle completely covered by its bracts, erect, shorter than the leaves and ± as long as the spike; lowest peduncular Bra leaf-like, imbricate, nearly all with leaf-like lamina; spikes ±7 cm, 0.7 cm wide, lanceolate, distichously arranged, rachis slender, glabrous; floral Bra densely imbricate, much exceeding the sepals, 27 × 9 mm, chartaceous and with coriaceous base, veined near the apex, sublustrous, ecarinate, concealing the rachis, 3–4× as long as the internodes, glabrous or the lower ones sparsely lepidote, red or sometimes yellow, obtuse; Fl sessile, distichous, 45–55 mm; Sep 14 (−20) × 3 mm, lanceolate, thin-coriaceous, even, evenly short-connate for 1 mm, adaxial ones bluntly carinate, yellow, glabrous, rounded or obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, 44 × 8 mm, lemon-yellow, blade tip margins recurved; St in 2 whorls of unequal length, exserted; Fil slender and flattened at the base and dilated becoming terete at the distal end, green; Anth dorsifixed near the middle, apex apiculate, brownish; Ov ellipsoid; Sty shorter than or equalling the shorter stamens; Sti spreading, green.

T. seleriana Mez (Bull. Herb. Boissier, sér. 2, 3: 84, 1903). Type: Mexico, sine loco (Seler 3439 [B]). — Distr: SE Mexico (Jalisco, Michoacán, Chiapas, Nayarit, Oaxaca, Veracruz), Guatemala, C & SW Honduras (Francisco Morazán, Lempira), NW El Salvador (Santa Ana), N & W Nicaragua (Estelí, Nueva Segovia); epiphytic in pine and oak woods, 270–2100 m. – Fig. 28.

[5] Plant stemless, flowering 20–25 cm tall; L much shorter than to exceeding the inflorescence, outer ones reduced and sheath-like, densely cinereous-lepidote or ferruginous-lepidote; L sheaths forming an ovoid pseudobulb 7–12 cm, broadly ovate or suborbicular, whitish-cinereous-green, merging into the lamina; L lamina linear to triangular, fleshy, 1 cm wide at the base, green (masked by the trichomes but less so than on the sheaths), erect or curved or secund, involute-subulate, apex attenuate; Inf 1× branched, subdigitate or nearly so, composed of 3–6 spikes, dense, 6–10 cm; peduncle often partly visible, erect, short and concealed by the leaves or rarely elongate; peduncular Bra leaf-like, lamina shorter than or exceeding the inflorescence, erect, densely imbricate except the upper ones, with rose-coloured sheath; primary Bra broadly ovate or elliptic, the lower ones slightly shorter and the upper ones much shorter than the axillary branches, shortly laminate or apiculate, densely lepidote, rose-red; spikes sessile, 4 cm, broadly elliptic, strongly complanate, 4- to 8-flowered, rachis straight, densely lepidote, exposed; floral Bra exceeding the sepals except in the top flower and so narrow as to expose part of the sepals, 20–28 × 9 mm, narrowly elliptic, incurved at the apex, coriaceous, margins with extended scales at the margins, slightly if at all carinate, not concealing the rachis, 4× as long as the internodes, densely cinereous-lepidote with coarse spreading scales, acute; Fl sessile, densely distichous, contiguous with each other; Sep 15.5–17 × 4 mm, narrowly elliptic, prominently veined, adaxial ones connate for 4–6 mm, adaxial ones distinctly carinate, green or red, glabrous, acute, obtuse; Pet forming an erect narrow tube clasping the filaments at the apex, constricted above the ovary, linear-oblong, 35–37 × 6 mm (3 mm at the base), violet for most of the length, blade tip margins recurved, narrowly rounded; St in 2 whorls of unequal length, 44–46 mm, exserted; Fil slender and flattened at the base and dilated becoming terete at the distal end; Anth dorsifixed near the middle, black; Ov ovoid, 6 mm; Sty slender, 44 mm, exceeding the stamens; Sti spreading, dilated, white.

T. subulifera Mez (Repert. Spec. Nov. Regni Veg. 16: 74, 1919). Type: Trinidad (Broadway 4200 [B, B [photo]]). — Distr: NE Nicaragua (Atlántico Norte), W Costa Rica (Puntarenas), C Panama (Panamá), Trinidad & Tobago (Couva-Tabaquite-Talparo (Chaguanas)), NW Colombia (Antioquia), Venezuela (Miranda, Sucre, Táchira); epiphytic in forests and plantations, 5–980 m.

[5] Plant stemless, flowering 15–19 cm tall; L few, erect, the outer greatly reduced, the inner to 18 cm, appressed-lepidote, with distinct or faint white cross-bands; L sheaths forming a cylindrical pseudobulb, inflated, about ½ as long as the lamina, concolorous with the lamina; L lamina linear, fleshy, coriaceous, 5 mm wide at the base, strongly angled, subulate and shallowly canaliculate, apex obtuse or abruptly acute; Inf simple; peduncle erect, shorter than and largely concealed by the leaves, slender; peduncular Bra erect, imbricate, many-veined, densely lepidote; spikes erect or curved downwards, 5–12 cm, ±1.5 cm wide, oblong-lanceolate or linear in outline, complanate, 4- to 12-flowered, without sterile bracts at the apex or base, rachis slender, flexuous to geniculate (when dry), appressed-lepidote, exposed, mostly not or only partly covered by the floral bracts; floral Bra erect, much shorter than the sepals, 20–25 × 15 mm, elliptic, incurved, margins hyaline, prominently veined, ± carinate towards the apex, appressed-lepidote, (yellowish-) green or roseate, broadly acute; Fl with a short pedicel-like receptacle, distichous, suberect; Sep slightly incurved at the apex excluding the apiculus, 22–25 × 8 mm, narrowly elliptic or narrowly obovate, finely veined (prominently when dry), free, ecarinate, rose, cinerous-appressed-lepidote, apiculate or narrowly obtuse; Pet forming a tube slightly widening towards the apex, linear-oblong, 32–34 × 7 mm (at the base 5 mm), rose, upper part yellow, often with rose margins and apex, narrowly rounded; St ± 32 mm, included and visible in the throat; Fil terete, slightly coiled halfway, rose towards the distal end; Anth dorsifixed near the middle, 3 × 1.8 mm, oblong, base bilobed, apex obtuse and apiculate; Ov 5.5 × 3 mm, ovoid, tapering form near the base, slightly contracted into the style; Sty slender, to 27 mm; Sti weakly conduplicate, spreading.

T. tectorum E. Morren (Belgique Hort. 27: 328, t. 18, 1877). Type: Peru (Wallis s.n. [BR]). — Lit: Hromadnik (2005: synopsis T. tectorum-complex). Distr: N & S Ecuador (Azuay, El Oro, Loja, Pichincha), Peru (Amazonas, Cajamarca, Huánuco, La Libertad, Ancash, Lima); saxicolous, and cultivated on roofs, 680–3400 m. – Fig. 29.

Incl. Tillandsia argentea K. Koch (1867) (nom. illeg., ICN Art. 53.1); incl. Pourretia nivosa hort. ex E. Morren (1877) (nom. inval., ICN Art. 36.1c); incl. Tillandsia saxicola Mez (1906).

[1] Plant short-stemmed to long caulescent, flowering 30–100 cm tall; L densely rosulate, 12–25 cm, white-tomentose-lepidote; L sheaths 1.2–3 cm wide at the base, triangular-ovate, clasping the stem, fleshy in the upper part, densely lepidote except the inner base, distinct from the lamina; L lamina very narrowly triangular, fleshy and 0.8–1.5 cm wide at the base, silvery-white, divergent to spreading, apex filiform-attenuate, trichomes eccentric, with elongated wings; Inf compound, densely digitate with ±4–6 spikes, 5–10 × (2.5–) 5–8 cm; peduncle erect, much exceeding the leaves, 10–40 cm × 5 mm ∅, slender, densely lepidote; lowest peduncular Bra somewhat leaf-like with setaceous lamina, sheaths imbricate, 1.5–2.5 × 0.8–1.2 cm, broadly ovate, membranous, veined, adaxially scatteredly lepidote and abaxially densely lepidote; primary Bra like the upper peduncular bracts, much shorter than the spikes; spikes sessile, erect or somewhat spreading, 3–4.5 × 1–1.2 cm, lanceolate, strongly complanate, (3- to) 5- to 10-flowered; floral Bra densely imbricate, ± equalling to exceeding the sepals, 10–17 × 7–11 mm, ovate, thin, veined, carinate at least at the apex, lepidote to subglabrous, red, green or roseate, acutish or apiculate; Fl subsessile, 20 mm; Sep 9–15 × 3–4 mm, ovate-lanceolate, thin, with broad hyaline margins, free, adaxial ones carinate, green and flushed red in the upper ½, abaxially sparsely lepidote along the midvein or glabrous; Pet erect and forming a tube, tongue-shaped, 15–25 × 3–4 mm, white with a blue band in the middle section, paler blue to whitish towards the apex, becoming pink after anthesis, blade divergent, obtuse; St all of equal length, 12–17 mm, included but visible in the throat; Fil narrowly ribbon-like, flaccid, straight or plicate, white; Anth subbasifixed or dorsifixed at ¼ from the base, 2–3 mm, linear, black or brown; Ov 2.5–3 × 2 mm, ovoid; Sty slender, 5–15 mm, longer than the stamens; Sti with short somewhat spreading lobes.

T. tenebra L. Hromadnik & W. Till (Bromelie 1991(2): 32, ill., 1991). Type: Argentina, La Rioja (Hromadnik & Hromadnik 7275 [WU, WU]). — Distr: Argentina (Catamarca, La Rioja, Buenos Aires, San Luis); saxicolous, 800–1900 m. – Fig. 30.

Incl. Tillandsia myosura var. saxicola Hieronymus ex A. Castellanos (1945) (nom. inval., ICN Art. 36.1c).

[4] Plant short-stemmed, flowering 4–6 cm tall, few-branched or forming clumps; L densely distichous, appearing to be equitant; L sheaths densely imbricate, 1–1.2 × 1.4 cm, upper part densely cinereous-lepidote like the lamina; L lamina succulent, rigid, 4–6 × 0.4–0.5 cm, recurved or spreading, adaxially grooved in the lower ½, subulate and laterally compressed, apex subulately acuminate and pungent, densely cinereous-lepidote, with extending trichomes at the margins, trichomes suborbicular; Inf simple; peduncle bractless or with 1–2 bracts at the base, erect, very short to elongate, lustrous, glabrous or lepidote near the first flower, often dark brown; peduncular Bra 2–4.5 cm, tightly enveloping the peduncle, fleshy, short-caudate; spike 1- to 2- (to 3-) flowered, rachis straight, lepidote, barely visible, internodes 0.25–0.4 (−0.65) cm; floral Bra much shorter than the sepals, (7–) 10–12 (−15) × 6–7 (−8) mm, broadly ovate or rarely triangular, margins hyaline, distinctly 14- to 16-veined, the lower one densely appressed-lepidote and the upper sparsely lepidote esp. at the base, cinereous or brown-red, apiculate; Fl subsessile, distichous, strongly fragrant; Sep (8–) 9.5–11 × (3–) 3.5–4.5 mm, elliptic, thin-coriaceous at the base, 8- to 10-veined (distinctly so at the apex), unequally short-connate for 1–3 mm, adaxial ones with a strong midvein and slightly carinate near the apex, brown to red in the upper part, glabrescent or sparsely appressed-lepidote at the apex, rounded or apiculate; Pet spatulate, 12–14 (−15) × 3.5–4.2 mm (base 1.5 mm wide), fleshy, with a thick midvein or with ridges, dark brown-violet, blade broadest near the apex, tapering into the cuneate claw, recurved, rounded; St all of equal length, 7 mm, deeply included; Fil 4–5.5 mm, ribbon-like and tapering from near the base to the apex; Anth basifixed, 1.5 × 0.5 mm, yellow; Ov 3.4–4 (−6) × 2–2.5 (−3) mm, angular-obconical, abruptly contracted into the style; Sty stout, 1.1–1.5 mm, 0.5–0.8 mm ∅, almost conical; Sti capitate.

T. tortilis Klotzsch ex Baker (J. Bot. 25: 237, 1887). Type [lecto]: Mexico, San Luis Potosí (Parry & Palmer 872 [BM, E, GH, K, MO, P, US]). — Lit: Till (1986). Distr: C & E Mexico, Jamaica; epiphytic or saxicolous, 1830–2440 m.

≡ Viridantha tortilis (Klotzsch ex Baker) Espejo (2002); incl. Platystachys ehrenbergii Beer (1856) (nom. inval., ICN Art. 36.1); incl. Platystachys tortilis Beer (1856) (nom. inval., ICN Art. 36.1); incl. Tillandsia ehrenbergii Klotzsch ex Beer (1856) (nom. inval., ICN Art. 38.1); incl. Tillandsia tortilis Klotzsch ex Beer (1856) (nom. inval., ICN Art. 38.1); incl. Platystachys ehrenbergii K. Koch (1874) (nom. inval., Art.?) ≡ Tillandsia ehrenbergii (K. Koch) Klotzsch ex Mez (1896); incl. Tillandsia ehrenbergiana Baker (1889).

[1] Plant short stemmed to distinctly caulescent, flowering 10–20 cm tall, forming dense clusters; stem 3–5 cm, simple or branched; L ±15, densely rosulate, to 17 cm, densely tomentose-lepidote or villous, with fine silver-grey scales; L sheaths 2 × 1.2 cm, elliptic, the lower part membranous, strongly veined, densely tomentose except at the base, distinct from the lamina; L lamina 6–15 × 0.5–1 cm, mostly spreading to reflexed, with a distinct midvein adaxially, with involute margins, subulate towards the long-attenuate apex, trichomes with elongated wings; Inf always simple; peduncle erect or ascending, much shorter than to just exceeding the leaves, 4–8 cm, 1–2 mm ∅, strongly sulcate, glabrous; lowest peduncular Bra leaf-like, imbricate, much exceeding the internodes, lanceolate, involute, thin, strongly veined, lepidote, the upper roseate, acuminate; spike 3–4 × 1–1.6 cm, elliptic, strongly complanate, 3- to 10-flowered, rachis slender, nearly straight; floral Bra densely imbricate, much exceeding the sepals, 20–26 × 8–12 mm, lanceolate, membranous, veined at least when dry, upper ½ distinctly carinate, concealing the rachis, 3–4× as long as the internodes, abaxially cinereously lepidote, roseate, acute or acuminate; Fl subsessile, distichous, suberect, dense; Sep (16–) 17–20 × 3–5 mm, lanceolate, membranous, strongly veined, subfree, adaxial ones carinate, rose but green in the lower ½, lepidote, acuminate; Pet erect with slightly divergent apex, linearly tongue-shaped, 25–38 (−40) × 4 mm, green, lower ½ white, rounded; St 24–31 mm, included (reported as exserted by Mez); Fil white; Anth basifixed, 4 mm; Ov ±4.5 mm, ellipsoid; Sty slender, 11–20 mm, shorter or ± equalling the stamens; Sti erect to spreading at maturity, green.

T. violaceiflora L. Hromadnik (Bromelie 2012(2): 49, ills. (pp. 48, 50), 2012). Type: Argentina, Salta (Hromadnik 7176 [MCNS, WU]). — Distr: NE Argentina (Salta: Quebrada de las Conchas); rocky cliffs, ±1500 m.

[5] Plant saxicolous, nearly stemless forming a short, ±3 mm thick rhizome, flowering Ros only 10–12 × 6–10 cm, growing in groups; L 4.5–9 cm, stiff; L sheaths 0.6–1 × 1.2–2 cm, trapezoid, amplexicaul, adaxially veined, glabrous, abaxially the upper ½ lepidote like the lamina, pale brownish, slightly distinct from the lamina; L lamina narrowly triangular, fleshy, rigid, 0.7–1 cm wide at the base, divergent and the longer ones upwardly secund, finely veined, strongly canaliculate and subulate at the somewhat pungent apex, cinereous-lepidote; Inf simple, erect or somewhat decurved, 6–8 cm, equalling or exceeding the leaves; peduncle completely covered by its bracts, short and concealed by the leaves, 3–4 cm, 2 mm ∅, glabrous, white; peduncular Bra 2–3 × 0.7 cm, oblong, membranous, strongly veined, glabrous or sparsely pale-lepidote on the lamina, the basal ones pale green, greenish-white or whitish, acuminate; spike 4 × 1.2 cm, lanceolate, strongly complanate, densely 3- to 5-flowered, rachis 4-angled, slender, glabrous, partly exposed, green, internodes 5 mm; floral Bra exceeding the sepals, 30 × 12 mm, lanceolate, subcoriaceous, smooth but rapidly veined after anthesis, glabrous, greenish to ruby-red, apiculate; Fl sessile, distichous, erect and densely appressed to the rachis, 40 mm, not fragrant; Sep 15–18 × 4 mm, lanceolate, the adaxial pair nearly free, adaxial ones carinate, keels red; Pet forming a narrow tube exceeding the floral bracts, then spreading, spatulate, 40 × to 5 mm (base 2 mm wide), violet, the lower 1/3 white, obtuse, abaxially weakly carinate; St 25 mm, included; Fil ribbon-like, straight, white; Anth subbasifixed, 3 mm; Ov 5 × 2 mm, ellipsoid; Sty slender, 25 mm, exceeding the stamens; Sti reaching the throat of the corolla, the narrow lobes spreading, white.

T. virescens Ruiz & Pavón (Fl. Peruv. 3: 43, t. 270: fig. b, 1802). Type: Peru, Huánuco (Ruiz & Pavón s.n. [MA?/BM?, F]). — Lit: Till (1989); Maruska (2012); Castello & Galetto (2013). Distr: SE Brazil (Rio de Janeiro), Peru, Bolivia, N & C Chile, N to NC Argentina, C Paraguay; saxicolous or epiphytic in dry habitats, 3–4000 m.

≡ Diaphoranthema virescens (Ruiz & Pavón) Beer (1856) ≡ Tillandsia capillaris fa. virescens (Ruiz & Pavón) L. B. Smith (1935); incl. Tillandsia propinqua Gay (1853); incl. Tillandsia recurvata Grisebach (1874) (nom. illeg., ICN Art. 53.1); incl. Tillandsia cordobensis Hieronymus (1885) ≡ Tillandsia capillaris fa. cordobensis (Hieronymus) L. B. Smith (1935); incl. Tillandsia stolpii Philippi (1895); incl. Tillandsia dependens Hieronymus ex Mez (1896); incl. Tillandsia dependens var. percordobensis Mez (1896) ≡ Tillandsia dependens fa. percordobensis (Mez) A. Castellanos (1933); incl. Tillandsia dependens var. perusneoides Mez (1896) (nom. inval., ICN Art. 26.1?) ≡ Tillandsia dependens fa. perusneoides (Mez) A. Castellanos (1933) (nom. inval., ICN Art. 26.1?); incl. Tillandsia williamsii Rusby (1910); incl. Tillandsia dependens var. sanzinii Hicken (1914) ≡ Tillandsia virescens var. sanzinii (Hicken) A. Castellanos (1945); incl. Tillandsia tomasii Halda (2005).

[4] Plant small but very variable in both size and form, flowering 2–16 cm tall, densely clump-forming; stems many from a single point, simple or branched; L distichous, mostly 1–4 (−9) cm (rarely <1 cm), cinereous- or ferruginous-lepidote; L sheaths usually elliptic, thin, several-veined, densely lepidote except where overlapping; L lamina linear or very narrowly triangular, ± succulent, to 2 mm wide, erect to spreading or recurved; Inf normally 1- or rarely 2-flowered; peduncle always naked, almost none or elongate (mostly after anthesis) to 8 cm, mostly slender after anthesis, glabrous or slightly lepidote near the flower; floral Bra usually equalling or exceeding the sepals, 5–7 × 3.5–4.5 mm, deltoid to oblong, closely enveloping the flower, thin, strongly 3- to 6-veined, ecarinate, densely lepidote at least at the very tip to glabrous, bluntly apiculate; Fl subsessile; Sep 6.5 × 1.3–1.5 mm, narrowly oblong, thin, with broad hyaline margins and apex, 3- to 5-veined with weakly developed vein branches, unequally connate for 1 or 5 mm, adaxial ones connate for ¾ or more, ecarinate but sometimes with pronounced midvein, green but often purple-speckled above the middle, obtuse or rounded; Pet tongue-shaped or linear or spatulate, 9–11.5 × 1 (−2) mm, white or yellow or brown, rounded or obtuse; St 3–4.5 mm, deeply included, exceeding the style; Fil thread-like, straight; Anth basifixed, 1–1.5 mm, linear, apex obtuse, green or cream-coloured; Ov 2 × 1.5 mm, subglobose; Sty stout, 1 mm; Sti disc-shaped, gemmate.

Difficult to distinguish from T. capillaris, which is also highly variable in habit and succulence. The main difference are the highly connate adaxial sepals (vs. < ½ connate in T. capillaris), but intermediate specimens are occasionally found.

T. weberi L. Hromadnik & P. Schneider (J. Bromeliad Soc. 37: 105, ills., 1987). Type: Mexico, Jalisco (Hromadnik & Schneider 12117 [JE, HAL, WU]). — Distr: S Mexico (Jalisco); epiphyte, ±800 m. – Fig. 31.

[5] Plant stemless, flowering to 30 cm, solitary or in small clusters; L few, to 10 cm, succulent, stiff, velvety white-lepidote with distinct fine longitudinal veining; L sheaths forming a pseudobulb almost closed at the top, conspicuous, 3 × 2.7 cm, ovate, inflated, abaxially densely cinereous-lepidote like the lamina except the lowest 3 mm of the base which are glabrous, inside brownish and with a ± distinct violet band about 2 mm wide, merging into the lamina; L lamina narrowly triangular, to 8 × 1.8–2 cm wide at the base, cinereous-lepidote, usually secund or erect, finely veined, the margins involute forming a seemingly subulate tip, apex stiffly pungent; Inf simple or rarely with 2 spikes; peduncle completely covered by its bracts, curved to ascending, to 15 cm, stout, glabrous, bright salmon-red; lowest peduncular Bra somewhat leaf-like, ±2.7 × 1 cm, enveloping the peduncle, membranous, veined, densely lepidote, green but the upper ones dark pink, acuminate and carinate at the tip; spikes to 14 × 0.6–0.8 cm, linear-lanceolate, complanate, (4- to) 10- (to 15-) flowered, rachis straight, completely covered by the bracts, internodes 9 mm; floral Bra exceeding the sepals, 25–30 × 9 mm, narrowly ovate, coriaceous, margins membranous and somewhat wavy and smooth, veined, ecarinate, abaxially lepidote or becoming subglabrous, pink, acute; Fl sessile, to 50 mm; Sep 20 × 5 mm, lanceolate, membranous, faintly veined, adaxial ones connate for 3–4 mm, thickened at the base, adaxial ones carinate, whitish-green with reddish tips, glabrous, acute; Pet forming an erect narrow tube clasping the filaments at the apex, 40 × 7 mm (3 mm at the base), upper ½ blue-violet, white below, blade tip margins recurving, apiculate; St exserted; Fil to 45 mm, slender and flattened at the base and dilated becoming terete at the distal end, spirally twisted around the pistil in the lower ½, thin, upper part blue-violet, otherwise white; Anth 3 × 1.5 mm; Ov 6 × 2 mm; Sty slender, 42 mm, exceeding the stamens; Sti shortly conduplicate-spiralized, spreading, dilated, white.

T. xiphioides Ker Gawler (Bot. Reg. 2: t. 105 + text, 1816). Type: [icono]: l.c. t. 105. — Lit: Gouda (2015). Distr: S Brazil (Rio Grande do Sul), S Uruguay, S Paraguay, Bolivia (Chuquisaca, La Paz, Potosí, Santa Cruz), Argentina (Jujuy, Salta, La Rioja, Córdoba, San Juan, Santiago del Estero, Mendoza); epiphytic and saxicolous in dry habitats, 1000–2700 m. I: Till & Barfuss (2014: 101, as var. minor). – Fig. 32.

≡ Anoplophytum xiphioides (Ker Gawler) Beer (1856) ≡ Phytarrhiza xiphioides (Ker Gawler) E. Morren (1879); incl. Tillandsia suaveolens Lemaire (1843); incl. Tillandsia macrocnemis Grisebach (1879); incl. Tillandsia sericea Hort. Ruch ex E. Morren (1879) (nom. inval., Art. 32.1c); incl. Tillandsia unca Hicken (1912) (nom. illeg., ICN Art. 53.1); incl. Tillandsia xiphioides var. minor L. Hromadnik (1989).

[2] Plant flowering 15–30 cm tall or taller, not or few-branched, stem shorter than the leaves; L laxly polystichous or sometimes almost distichous, few to many, erect to spreading or ± curved, densely cinereous- or ferruginous-lepidote, scales mostly with broad lobes; L sheaths large, densely imbricate making the stem appear 1–2 cm thick, passing almost imperceptibly into the lamina; L lamina narrowly triangular, to 25 × 2 cm but often much shorter, flat at the base but mostly with involute margins upwards, apex attenuate-subulate; Inf always simple; peduncle hidden by the inner leaves and bracts, erect, almost none to 12 cm; peduncular Bra densely imbricate and hiding the peduncle, elliptic-oblong, thin, lepidote or quite glabrous or sparsely pale-lepidote, caudate (lower ones) to apiculate (upper ones); spike with several sterile bracts at the base, to 12 cm (excl. petals), lanceolate or oblong, 2- to 10-flowered, acute, rachis narrowly alate, flexuous, to 3 mm ∅; floral Bra densely imbricate, much exceeding the sepals, 40–56 (−70) × 11–13 mm, lanceolate-oblong, somewhat incurved at the apex, submembranous, margins broadly hyaline, veinless but finely to strongly veined when dry, ecarinate, usually many times longer than the internodes, glabrous or sometimes the lower ones sparsely lepidote, mostly greenish or stramineous or suffused with red or violet, attenuate or acutish; Fl sessile or subsessile, distichous, to 100 mm, fragrant; Sep (25–) 35 (−42) × (6–) 7–9 mm, linear-lanceolate, submembranous, prominently veined, free or subfree, adaxial ones finely carinate from base to apex, pale green, glabrous, acuminate or acutish; Pet forming a narrow tube exceeding the floral bracts, then spreading, claw narrow with suborbicular blade, 70–85 × 10–15 (−20) mm, margins conspicuously crenate-serrate, white, blade subrhombic or obovate, recurved to weakly helicoid, obtuse; St 43–53 mm, included or visible in the throat; Fil flaccid, becoming filiform upwards, straight; Anth basifixed, 8 mm, linear; Ov slenderly prismatic-ovoid, 6–10 mm, tapering into the style; Sty slender, including stigma 55–68 mm, exserted from the corolla throat at late anthesis; Sti recurved, linear.

A very variable species as to size, succulence and indumentum, with many microspecies. The leaves vary from long and leathery to short and fleshy.

T. yuncharaensis W. Till (J. Bromeliad Soc. 40: 214, ills., 1990). Type: Bolivia, Tarija (Haugg & al. WT55 [WU]). — Distr: S Bolivia (Tarija); saxicolous, ±3000 m. – Fig. 33.

[2] Plant long-caulescent, stem to 30 cm; L laxly polystichous; L sheaths 0.7 × 1.8 cm, broadly triangular, amplexicaul, glabrous in most parts, brownish, merging into the lamina; L lamina with subciliate scales at the margins, to 5 × 1 cm, spreading to recurved, not secund, canaliculate, veined when dry, ± appressed-lepidote, apex attenuate and acute; Inf simple; peduncle completely covered by its bracts, only 2–3 cm, 2 mm ∅, angled when dry, glabrous; peduncular Bra few (3–4), the lowest somewhat leaf-like, the upper ones like the floral bracts, much exceeding the internodes, veined, lepidote at least towards the apex, reddish, acuminate or broadly acute; spike 5.5–6.5 × 0.8–1.2 cm, linear-lanceolate, slightly to strongly complanate, (1- to) 2- to 4- (to 6–) flowered, acute, rachis glabrous; floral Bra exceeding the sepals, 35–42 × 11–13 mm, margins hyaline, even, ecarinate, concealing the rachis, glabrous in part with very few scale-hairs near the apex, carmine-red with greenish margins, acute; Fl distichous, 37–40 mm, strongly fragrant; Sep 23–25 × 6–7 mm, ovate-lanceolate, smooth, evenly short-connate for 1 mm, adaxial ones very bluntly carinate, subalate only at the apex, green tinged with purple, glabrous, subobtuse; Pet forming a narrow tube exceeding the floral bracts, then spreading, 45–47 × 7 mm, margins slightly undulate, violet or white (sometimes with blue margins), claw 30 × 3–3.5 mm, blade 17 mm, subrhombic, curved-spreading; St in 2 whorls of unequal length, 24–30 mm, visible in the throat; Fil 28–29 × 1 mm, ribbon-like or filamentous upwards, not plicate, white; Anth subbasifixed, 2.5 (5 when fresh) × 0.5 mm, linear; Ov 5 × 2–2.5 mm, conical-cylindrical or ovoid, contracted into the style; Sty slender, 33 mm (incl. stigma), slightly exceeding the petal tube and the stamens; Sti dilated-conduplicate, spreading.

T. zecheri W. Till (Bromelie 1983(1–2): 6–7, ill., 1983). Type: Argentina, Salta (Hromadnik & Hromadnik 7162 [WU]). — Distr: NE Argentina (Salta); saxicolous on river banks, 1900–2500 m. – Fig. 34.

[1] Plant short-stemmed, branched or solitary; L suberect, densely cinereous or densely tomentose-lepidote in some forms; L sheaths veined, lepidote, indistinctly merging into the lamina; L lamina narrowly triangular, 13–17 cm, rigid or succulent in some forms, slightly secund to curved, densely cinereous-lepidote, often veined with the trichomes in longitudinal lines, trichomes with elongated wings esp. at the leaf base, lamina with involute margins forming a channel, apex obtuse or pungent; Inf simple; peduncle completely covered by its bracts, slightly curved, short and concealed by to equalling the leaves, 2–7 cm, straight; lowest peduncular Bra with short leaf-like lamina, erect, imbricate, 3–4 cm, the upper ones lanceolate, enveloping the peduncle, greyish, acute; spike with a sterile bract at the base, recurved, 14 × to 3 cm, lanceolate, complanate, densely (3- to) 4- to 6- (to 8-) flowered, acute, rachis laterally compressed, slightly flexuous, glabrous, normally hardly visible at anthesis, green; floral Bra ± densely imbricate but partially exposing the rachis after anthesis, exceeding the sepals, (18–) 27–35 (−48) × 9–11 mm, often greenish, margins finely veined or even and sometimes waxy, ecarinate, glabrous, reddish, obtuse; Fl subsessile, not fragrant; Sep (18–) 20–26 × 4.5–5 mm, oblong-lanceolate, equally free or evenly short-connate for 1–2 mm, adaxial ones carinate, green, (obscurely) acuminate; Pet tongue-shaped, 37–40 × ±5 mm, dark purple-violet or bluish, blade spreading to recurved, obtuse; St unequal in length, ± equalling the petal claw at anthesis; Fil slightly (if at all) plicate in the middle; Anth dorsifixed near the base, linear, apex apiculate; Ov ovoid, 5 mm; Sty exceeding the stamens and exserted from the throat; Sti weakly conduplicate, erect or slightly divergent.

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Curator University Botanic Gardens, Utrecht, The Netherlands
E. J. Gouda

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Gouda, E.J. (2019). Tillandsia BROMELIACEAE. In: Eggli, U., Nyffeler, R. (eds) Monocotyledons. Illustrated Handbook of Succulent Plants. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-662-56324-3_96-1

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DOI: https://doi.org/10.1007/978-3-662-56324-3_96-1
Received: 05 January 2019
Accepted: 05 January 2019
Published: 16 May 2019
Publisher Name: Springer, Berlin, Heidelberg
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Online ISBN: 978-3-662-56324-3
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