A new species of Tillandsia (Tillandsioidae, Bromeliaceae) from the high Andean forests near Bogotá (Colombia) Author Aguirre-Santoro, Julián 0000-0002-8640-4932 Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Sede Bogotá & jaaguirresa @ unal. edu. co; https: // orcid. org / 0000 - 0002 - 8640 - 4932 jaaguirresa@unal.edu.co Author Hernández-A, Érika Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Sede Bogotá & Parques Nacionales Naturales de Colombia, Bogotá, Colombia Author Hernández-Rodríguez, David E. 0009-0005-5642-4085 Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Sede Bogotá & dehernandezr @ unal. edu. co; https: // orcid. org / 0009 - 0005 - 5642 - 4085 dehernandezr@unal.edu.co Author Betancur, Julio 0000-0002-5804-4677 Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Sede Bogotá & jcbetancurb @ unal. edu. co; https: // orcid. org / 0000 - 0002 - 5804 - 4677 jcbetancurb@unal.edu.co text Phytotaxa 2023 2023-08-14 607 5 262 272 http://dx.doi.org/10.11646/phytotaxa.607.5.1 journal article 54367 10.11646/phytotaxa.607.5.1 4639b3f3-1438-40b1-af59-4b39b7f3b7f1 1179-3163 8248846 Tillandsia chingacensis Aguirre-Santoro, E. Hern-A., & Betancur , sp. nov. Type: — COLOMBIA . Meta : municipio San Juanito , área de amortiguación del Parque Nacional Natural Chingaza , vereda El Tablón , sendero desde la finca Campo Alegre hacia la finca Brasil , áreas de bosque alto andino y potrero, 2220 m , 4°28’44.7”N , 73°38’36.6”W , 19 March 2018 , Julián Aguirre-Santoro 3592, Erika Hernández-Aldana , Paola Pulido , Lisa Anzellini , Ariel Parrales , Víctor Peña & Damián León ( holotype COL !, isotype FMB !). ( Figs 1 , 2 , 3 ) . Tillandsia chingacensis is similar to T. clavigera and T. nervisepala but differs from them by its long-triangular leaf blades ( vs. lingulate to sublingulate), inflorescence, bracts, and sepals abundantly covered with a resinous substance ( vs. without this substance), floral bracts longer ( 4–5.2 cm vs. 2.7–3.5 cm long) and strongly nerved ( vs. smooth), sepals all alike and ecarinate ( vs. abaxial sepal carinate) and longer ( 44.1–53.6 mm vs . 20–30 mm long), petal appendages present ( vs. absent), and longer capsules ( 7.4–10 cm vs. < 5.5 cm long). Plant epiphytic, solitary, acaulescent, not stoloniferous, flowering 160–200 cm tall; rosette broad, tank-forming. Leaves (47–)54–57(–61) cm long, coriaceous; sheath conspicuously differentiated from the blade, elliptic, 14.8–20.8 × 10.8–13.2 cm , white to light-brown at basal two-thirds, then dark castaneous to vinaceous distally; blade divergent to divaricate, narrowly triangular, 33–41 cm long, 6.3–8.8 cm wide at the base, 5.5–6.8 cm wide in the middle, green adaxially, grayish abaxially, sometimes dark vinaceous and maculate at the apex at adaxial side and completely vinaceous on the abaxial side, lepidote on both surfaces, more densely so on the abaxial side, apex acuminate to attenuate, pungent. Peduncle elongate, much exceeding the leaves of the rosette, erect, stout-rigid, terete, 69–151 cm long, (7.6–)20.2–20.4(–36.2) mm in diameter when dry, pink-magenta, glabrous, basal internodes 32–68 mm long, medial and distal internodes 43–85 mm long. Peduncle bracts persistent, erect, coriaceous, slightly nerved when dry; the basal ones much longer than the internodes, similar to the inner leaves of the rosette; the central ones longer to shorter than the internodes, ovate, dark-magenta, 48–83 × 25–37 mm , apex attenuate and apiculate; the distal ones equaling to shorter than the internodes, very broadly ovate to orbicular, dark-magenta, 38–57 × 23–31(–45) mm, apex attenuate and short-apiculate. Fertile part of the inflorescence copiously covered with a resinous substance (including all bracts, sepals and petals), erect, once-branched, paniculate, conical in outline, (43–) 83–103 cm long, 25.5–44 cm wide in the base, with 3 to 10 polystichously arranged branches; main axis straight, pink-magenta, (22.5–) 54–79 cm long, (4.8–)7.5–12(–21) mm in diameter. Primary bracts similar to the peduncle bracts, shorter than the stipe of the branches, not diminishing in size distally, spreading with the branches, persistent, coriaceous, strongly nerved; the basal ones broadly ovate, 4.7–6.8 × 2.7–3.8 cm , magenta; the medial and distal bracts ovate, 4.5–4.7 × 2.8–3.6 cm , magenta. Racemes oblong to elliptical in outline, 8.3–22(–36) cm long, 4–5.8 cm wide in the middle; stipe elongate, recurving, for most part exposed, 5–10(–16) cm long, 3.8–6.4 mm in diameter, terete, with 1–2(–4) carinate sterile bracts; rachis of the raceme straight, flat. Floral bracts persistent, gradually diminishing in size towards the apex of the raceme, just exceeded by the sepals, imbricate, contiguous, sub-erect, concave, coriaceous, sub-orbicular to broadly ovate, 3.9–5.2 x 1.4–4.5 mm , pink-magenta, strongly nerved when dry, ecarinate, apex acute to rounded and apiculate. Flowers 6–9(–17) in number per raceme, distichous, contiguous, 39.7–52.2 mm long, pedicellate; pedicel 5.6–10.3 mm long, 6.1–6.8 mm in diameter. Sepals all alike, free, coriaceous, narrowly elliptical, symmetrical, 44–53.6 x 8.7–17 mm , light green, strongly nerved, ecarinate, obtuse. Corolla long-tubular, apically spreading; petals free, membranaceous, oblong to lanceolate, 74 x 4–7.5 mm , dark violet, apex acute to acuminate, bearing two appendages at the base; petal appendages flanking the antepetalous stamen, free lobe 5–6.8 mm long, detaching from the petal at 12–13 mm from the base, apex fimbriate. Stamens included, nearly equaling the petals, free, all equal in length, occasionally heterandrous; filament terete, 55 cm long, dark purple at least in the distal half; anther versatile, oblong, 10 × 1.8 mm , dark purple to black, pollen yellow. Ovary conical, terete in cross section, 10.4–12.1 mm long, 3.7–4.8 mm in diameter, placentation apically axile. Style shorter to much exceeding the stamens, included or exserted, terete, to ca. 7 cm long, purple; stigma conduplicate-spiral, purple, 3.6 mm long. Ovules many, ovoid. Fruit capsular, elongate, terete, 7.4–10 cm long, 8.5–11.5 mm in diameter, brown. Seeds many, bearing a long plumose basal appendage, 5.3–6.8 cm long (including the apical and basal appendages), with each seta of the appendage bearing minuscule retrorse appendages, white. Etymology : —The specific epithet refers to Chingaza National Natural Park, a protected area where most of the specimens were collected. This area is part of the National System of Protected Areas of Colombia , and its importance is crucial for Bogotá , the capital of Colombia , as it provides water for a large part of the city’s population. FIGURE 1. Tillandsia chingacensis . A. General habit of the plant. B. Leaf blade. C. Sterile bract, floral bract, and emerging fruit. D. Flowering raceme. E. Sepals. F. Dissected corolla showing petals, petal appendages, androecium and gynoecium. G. Fruiting raceme. H. Seed showing the chalazal appendage in the lower part and the micropylar appendage in the upper part. I. Detail of the basal-most part of the micropylar appendage of the seed. J. Detail of a seed bristle. Illustration by Marcela Morales. FIGURE 2. Tillandsia chingacensis . A. Habit. B. Center of the rosette. C. Fertile part of the inflorescence. D. Part of the Peduncle. E. Branch of the inflorescence. F. Detail of the floral bracts and flowers (Photos David E. Hernández-Rodríguez and Érika HernándezAldana). FIGURE 3 . Artistic illustration of Tillandsia chingacensis showing the habit and colors of the living plant. Illustrated in the type locality by Lisa Anzellini. Distribution and habitat : —This species grows as a solitary epiphyte in robust trees of forested areas and forest edges of rugged topography crossed by deep canyons. It is endemic to the medium to high elevation forests of the eastern Cordillera of Colombia located in remote areas of Chingaza National Natural Park at 2300–3100 m elevation. Specifically, this species has been collected in three municipalities embedded in Chingaza National Park and its surrounding areas: San Juanito ( Meta ), Junín and Gachalá ( Cundinamarca ) ( Fig. 4 ). Conservation status : —The species is only known from three localities with an AOO of 20000 km 2 (AOO based on user defined cell width ( 2 km )) and a EOO of 66.174 km 2 . Two of these localities are protected by Chingaza National Natural Park and correspond to a continuous matrix of well-conserved forest. The locality in San Juanito, however, corresponds to a highly fragmented forest surrounded by intense deforestation for agricultural expansion. Based on the number of endangered localities, small AOO and EOO, we propose the categorization of this species as endangered EN B2ab(iii) following the guidelines of the IUCN ( IUCN 2022 ). Additional specimens examined:— COLOMBIA . Cundinamarca : municipio Junín , vereda Colombia , sector Carpanta , borde de bosque, quebrada Golpe de Agua , límite del Parque Nacional Natural Chingaza , 2546 m , 4º35’14.2”N , 73º40’35.2”W , 31 January 2018 , Hernández-Aldana 460 ( COL !) ; municipio Gachalá , vereda Tendidos de Río Negro , borde de bosque, quebrada La Palma , área de amortiguación del Parque Nacional Natural Chingaza , 2200 m , 4°35’50.7”N 73°36’6.9”W , 27 September 2019 , Hernández-Aldana 1041 ( COL !) . Meta : municipio San Juanito , área de amortiguación del Parque Nacional Natural Chingaza , vereda El Tablón , sendero hacia El Palmar , entre las fincas Campo Alegre y San Pio , bosque alto andino, 2180 m , 4°28’25.5”– 4°28’14.3”N 73°39’14”– 73°38’33.4”W , 13 March 2018 , Aguirre-Santoro 3513 ( COL !) ; municipio San Juanito , área de amortiguación del Parque Nacional Natural Chingaza , vereda El Tablón , sector el Palmar , bosque alto andino, 2440 m , 4°28’11.6”N – 73°38’20.9”W , 17 March 2018 , Aguirre-Santoro 3580 ( COL !) . FIGURE 4. Geographic distribution of Tillandsia chingacensis . The top-right map highlights the Chingaza Natural National Park (gray area) along with the surrounding municipalities (white areas). The larger map provides a detailed view, indicating the georeferenced collection points of Tillandsia chingacensis . Observations :— According to the latest complete taxonomic treatment of Tillandsioideae ( Smith & Downs 1977 ) , this new species should be strictly placed in the genus Vriesea because of its distichous flowers and presence of petal appendages. Nevertheless, the validity of this combination of characters to separate Vriesea from Tillandsia has been questioned by bromeliad taxonomists ( Barfuss et al. 2016 ; Costa et al. 2014 ; Grant 1993 ; Kessous & Costa 2023 ). Factors supporting this reconsideration include the plasticity of petal appendages ( Brown & Terry 1992 ; Gouda 1987 ), molecular analyses suggesting a distinct, Brazilian-centered clade for Vriesea ( Barfuss et al. 2016 ; Machado et al. 2020 ), and the resemblance of certain Andean Vriesea species to those of Tillandsia ( Barfuss et al. 2016 ; Grant 1993 ; Kessous & Costa 2023 ). In the case of new species described here, these arguments equally justify our decision to place it in Tillandsia . The new circumscription of Vriesea in the taxonomic treatment of Tillandsioideae of Barfuss et al. (2016) strongly rejects the placement of the new species within this genus because of its purple petals ( vs . yellow to brown) and Andean distribution ( vs. mainly Brazilian).According to the new taxonomic reorganizations proposed in this taxonomic treatment and the recent revision of Kessous & Costa (2023) , the Andean distribution and presence of petal appendages of Tillandsia chingacensis would rather place this species either in the genus Cipuropsis or Tillandsia . In the former case, T. chingacensis does not fit within the diagnosis of Cipuropsis because of its long-triangular leaf blades ( vs. lingulate), petals completely free ( vs. 1/4 conglutinated/connate) and purple petals ( vs. yellow or white), and free anthers ( vs. united into a tube surrounding the stigma). Additionally T. chingacensis fits with confidence within the limits of Tillandsia mainly because of its conduplicate-spiral stigma and free purple petals. Nonetheless, the precise placement of T. chingacensis within specific subgenera and species complexes of Tillandsia remains uncertain due to the absence of clear defining criteria for these infrageneric groups. While one possible option is assigning the species to T. subgenus Pseudovriesea , it lacks the distinctive xeromorphic leaves and long-triangular leaf blades characteristic of this group. Besides the phylogenetic and morphological reasons explained above, our decision to place the new species in Tillandsia is strongly based on its affinity with Tillandsia subg. Allardtia ( Dietrich (1852: 241) Baker (1888: 40) . Tillandsia chingacensis fits within this subgenus because of its included stamens with straight filaments and slender style ( Smith & Downs 1977 ). Within Tillandsia subg. Allardtia , T. chingacensis is likely related to a group of northern Andean-centered species encompassed by T. brevicapsula Gilmartin (1968: 156) , T. clavigera Mez (1896: 783) , T. fendleri Grisebach (1864: 17) , T. francisci Till & Grant (2003: 195) , and T. nervisepala ( Gilmartin 1968: 157 ) Smith (1974: 36) . These species can be recognized by their large size ( 1.5–2 m tall), inflorescences laxly compound bearing spreading to reflexed racemes or spikes, relatively long floral bracts (> 2.5 cm long) that are frequently carinate (not in T. chingacensis ), and free sepals of> 20 mm long. In Table 1 the main similarities and differences among these species are compared. TABLE 1. Comparative table of diagnostic characters for the similar species in Tillandsia subg. Allardtia that resemble T. chingacensis .

Character	T. chingacensis	T. brevicapsula	T. clavigera	T. fendleri	T. francisci	T. nervisepala
Blade shape	long-triangular	lingulate	lingulate	Lingulate, triangular	triangular	lingulate
Primary bracts vs . branch stipe	shorter	shorter	longer	longer	longer	shorter
Branch direction	reflexed	reflexed	reflexed	spreading to recurved	reflexed	reflexed
branch length	8.3–36 cm	20–22 cm	ca. 30–55 cm	10–30 cm	18-20 cm	20–22 cm
Sepal apex	acute	obtuse	acute	acute	acute	rounded
Sepal length	4.4–5.3 cm	2 cm	2.8–3.0 cm	2.5–4.5 cm	3.8–4.2 cm	2 cm
Sepal keel	ecarinate	carinate	carinate	carinate	carinate	carinate
Petal appendages	present	absent	absent	absent	absent	unknown
Capsule vs. sepals	more than two times longer	equaling	shorter	shorter to equaling	shorter	nearly two times longer

Among this group of similar species, Tillandsia chingacensis is unique because of its inflorescence (including bracts, sepals and petals) abundantly covered with a resinous substance ( vs . dry surface), ecarinate sepals ( vs . adaxial sepals carinate), presence of petal appendages ( vs . absence), and very long capsules ( 7.4–10 cm vs . < 5.5 cm long). In particular, T. chingacensis closely resembles the sympatric T. clavigera , and T. nervisepala from Ecuador because of its reflexed racemes and long stipe of the branches. However, T. chingacensis differs from T. clavigera and T. nervisepala by its long-triangular leaf blades ( vs . lingulate to sublingulate), strongly nerved ( vs . smooth) and longer floral bracts ( 4–5.2 cm vs . 2.7–3.5 cm long), and longer sepals ( 4.4–5.4 cm vs . 2–3 cm long), in addition to other differences explained above. Finally, the artificial key of Smith & Downs (1977) suggests a possible affinity of Tillandsia chingacensis with T. carnosa Smith (1963: 251) . According to the main Tillandsia key, narrow triangular leaf blades, the primary bracts shorter than the axillary branches of the inflorescence and the dense distichously flowered spikes, lead to Subkey IX. Within this subkey, T. chingacensis keys out to T. carnosa because of its ecarinate sepals and floral bracts 4–6 times longer than the internodes of the rachis.Nevertheless, T.chingacensis mainly differs from T.carnosa by its inflorescence, bracts, sepals and petals abundantly covered with a resinous substance ( vs . dry surface), erect inflorescence ( vs . nutant to pendent), once-branched inflorescence ( vs . occasionally twice-branched), floral bracts exposing the calyx ( vs . covering the calyx) and coriaceous sepals ( vs . fleshy), longer than 44 mm ( vs . up to 42 mm ), petals appendaged ( vs . naked), and stamens included in the corolla ( vs . slightly exceeding the corolla).