Fosterella penduliflora (C.H. Wright) L.B. Smith, Phytologia

Leme, Elton M. C., Forzza, Rafaela C., Halbritter, Heidemarie & Ribeiro, Otávio B. C., 2019, Contribution to the study of the genus Fosterella (Bromeliaceae: Pitcairnioideae) in Brazil, Phytotaxa 395 (3), pp. 137-167 : 157-164

6. Fosterella penduliflora (C.H. Wright) L.B. Smith, Phytologia 7: 172. 1960. ( figs. 12 , 13 )

Basionym:— Catopsis penduliflora C.H. Wright , Bulletin of Miscellaneous Information Kew 1910: 197. 1910. Type:— PERU. Without further locality, M. Forget s.n., described from a living specimen sent by Messrs. F. Sander & Sons, March 1910 [lectotype K!, designated by Peters et al. (2008)].

≡ Lindmania penduliflora (C.H. Wright) Stapf, Bot. Mag. 150: pl. 9029. 1924.

= Fosterella chiquitana Ibisch, R. Vásquez & E. Gross, Rev. Soc. Boliviana Bot. 2: 118. 1999. Type:— BOLIVIA. Depto. Santa Cruz: Prov. Ñuflo de Chávez, about 10 km from Concepción, on the road to San Javier, on granitic outcrops, 16° 10’ S, 62° 05’ W, 500 m elev., 16 May 1997, P. Ibisch 98.0125, flowered in the garden of Pierre L. Ibisch, Santa Cruz de la Sierra, from 21 August 1998 onwards (holotype LPB, isotypes FR, WU!).

= Fosterella latifolia Ibisch, R. Vásquez & E. Gross, Rev. Soc. Boliviana Bot. 2: 123. 1999. Type:— BOLIVIA. Depto. Santa Cruz: Prov. Florida, Refugio Volcanes, in understory of semihumid forests, 18° 05’ S, 63° 40’ W, 1000 m elev., 16 May 1998, P. Ibisch 98.0098, (holotype LPB, isotype SEL!).

Description:— Plants acaulescent, flowering 40–60 cm tall. Leaves ca. 10 in number, subspreading; sheath 20–50 mm wide, entire, whitish, glabrous; blade narrowly lanceolate, apex acuminate-caudate, distinctly narrowed towards the base but not petiolate, thick towards the base, thin in texture toward the apex and margins, 15–45 × 3–8 cm, laxly to subdensely white lepidote abaxially, with peltate, dentate trichomes (type III), adaxially glabrous, green or sometimes reddish, margins entire, undulate, sometimes the basal margins densely white lepidote. Peduncle erect, 20–36 cm long, 2–5 mm in diameter, green to reddish, glabrous; peduncle bracts the basal ones subfoliaceous, the upper ones narrowly lanceolate, long caudate, erect, longer to shorter than the internodes, membranaceous, green, nerved, entire, subdensely white lepidote abaxially. Inflorescence compound, racemose to paniculate, with second to third order branches, erect, 24–40 cm long, 10–25 cm in diameter; main axis ca. 1.5 mm in diameter, glabrous; primary bracts resembling the upper peduncle bracts, entire, 5–15 × 2–3 mm, distinctly shorter than the stipe, white lepidote abaxially; primary branches 10–15 in number, suberect, laxly arranged, (6-) 11–22 cm long, 12–35-flowered, bearing 1–2 third order branchs; stipe slender, 7–30 × 1.5–2 mm, naked or bearing 1–2 sterile bracts or with aborted flowers or dormant secondary branches; rachis ca. 1 mm in diameter, straight to slightly flexuous, green, glabrous; secondary bracts resembling the upper primary bracts; secondary branches 3–5 cm long, 5–10-flowered; floral bracts ovate-lanceolate, acuminate, 2–4 × 1–1.5 mm, entire, membranaceous, green, nerved, inconspicuously and sparsely white lepidote with filamentous trichomes, slightly shorter than to equaling the pedicel. Flowers 9–10 mm long (with extended petals), 3–7 mm apart, doward secund; pedicel 2–6 mm long, ca. 0.5 mm in diameter, curved downward; sepals ovate, acute, 2–3 × 1 mm, green, glabrous; petals narrowly oblong-ovate, obtuse, 7–10 × 2.5 mm, whitish, distinctly recurved at anthesis but not recoiled, somewhat straight afterward; stamens distinctly shorter than the petals, but completely exposed at anthesis; anther narrowly oblong, 1.5–2 mm long, dorsifixed near the base, distinctly recurved-recoiled at anthesis; stigma apparently conduplicate-spiral; ovary ellipsoid. Capsules ovoid, ca. 4 × 2 mm; seeds filiform, bicaudate, 2–2.4 mm long.

Additional specimens examined:— BRAZIL. Rondônia: Ariquemes, Mineração Mibrasa, Setor Alto Candeias, km 128, 10° 35’ S, 63° 35’ W, Sudoeste de Ariquemes, Mata sobre serra, beira de cachoeira, abundante sobre as rochas, 17 May 1982, L.O.A. Teixeira et al. 549 (K!, MO!, RB!).

Distribution and habitat: — According to Peters (2009), F. penduliflora has the broadest geographical distribution in the genus, growing in Peru (the type collection only), Bolivia, and Argentina, with some registered Bolivian populations thriving about 210 km, in straight line, from Brazil’s border. There is only one collection now reported (and examined) from Brazil, in Rondônia state, about 550 km apart, in straight line, from the known Bolivian populations. In Brazil, the cited specimens were found growing on a hill 280–300 m elevation, saxicolous on rocks inside the forest, along a waterfall. It was reported as an abundant species when it was collected 36 years ago. However, an expedition organized few months ago with the specific purpose to locate it did not succeed, probably because the area is now severely affected by cattle breeding and periodical fires.

Etymology:— The name of this species is an explicit reference to its pendulous flowers.

Observations:— The absence of living specimens for deeper observation of floral characteristics and the presence of leaf trichomes of the peltate, dentate type III, documented by SEM images ( figs. 3 M, N, O ) encourage us to identify the specimens from Rondônia, at least provisionally, as F. penduliflora on the basis of the identification key provided by Peters (2009). However, the specimens from Rondônia (e.g. fig. 12 ), when compared to the Bolivian specimens and to the lectotype ( fig. 13 ), in the morphological circumscription of Peters (2009), present some discordant features, like leaf blades subdensely lepidote (vs. laxly lepidote) with margins densely lepidote along the basal portion (vs. glabrous), upper peduncle bracts shorter than the internodes (vs. equalling to longer than the internodes), and inflorescence with third order branches (vs. with second order branches only).

In comparison to the other Brazilian species, F. penduliflora is closer related to Amazonian F. batistana , with which it shares the same peltate, dentate trichomes (type III), despite distinctly sparsely arranged in F. batistana . Other differences of F. penduliflora are its larger size when in flower (40–60 cm vs. 15–30 cm high), longer peduncle (20–36 cm vs. 5–15 cm), longer inflorescence (24–40 cm 6–18 cm), longer primary branches [(6-) 11–22 cm vs. 2–8.5 cm], and longer pedicel (2–6 mm vs. 1–3 mm).

7. Fosterella windischii L.B. Smith & R.W. Read , Bradea 6: 137. 1992. Type:— BRAZIL. Mato Grosso, Serra Ricardo Franco, em fendas na encosta, 450–500 m elev., 15º S, 60º W, 23 September 1978, P. C. Windisch 2044 (holotype US!, isotypes HB!, HRCB, MBM!). ( figs. 14 A to G)

Description:— Plants acaulescent, flowering 40–70 cm tall, propagating by basal shoots. Leaves ca. 12 in number, fasciculate, subspreading, equaling to exceeding the peduncle, the basal ones slightly reduced; sheath inconspicuous, subreniform, ca. 1 × 2.5 cm, slightly thickened, corrugated, greenish, densely white lepidote distally with interwoven trichomes somewhat forming a membrane; blade linear-lanceolate, apex slenderly caudate, slightly narrowed at the base, thin in texture toward the margins and apex, thickened on the central channel and toward the base, 17–50 × 1.5–2.8 cm, green, finely nerved, abaxially completely covered by a thick and coarse layer of white, interwoven peltate, shortly fimbriate trichomes (type I), adaxially and near the base densely white lepidote with large and coarse adpressed trichomes and glabrous toward the apex, abaxial and adaxial faces strongly contrasting in color, margins inconspicuously and subdensely spinulose at the base and entire toward the apex, undulate, lepidote near the base to glabrous. Peduncle erect, 15–20 cm long, 2–3 mm in diameter, green, glabrous; peduncle bracts 1.5–7 × 0.3–0.5 cm, narrowly triangular-lanceolate, slenderly caudate, erect, thin in texture, nerved, entire, densely white lepidote with coarse adpressed trichomes, longer or the upper ones slightly shorter than the internodes. Inflorescence shortly paniculate, with third order branches, erect, 14–40 cm long, 8–20 cm in diameter; main axis 1–2 mm in diameter, green, glabrous; primary bracts resembling the upper peduncle bracts, filiform-caudate, entire, 3–20 × 1–2 mm, densely and coarsely white lepidote, longer than (basal ones) to equaling the middle of the stipe (upper ones); primary branches 5–19 in number, suberect-ascending, subdensely to laxly arranged, 3–18 cm long, 6–26-flowered, the basal to median ones bearing 1–3 secondary branches; stipe slender, 10–25 × 1 mm, naked; secondary bracts resembling the floral bracts, many times shorter than the stipe; secondary branches 1.5–8 cm long, 3–19-flowered; rachis ca. 1 mm in diameter, nearly straight, green, glabrous; floral bracts ovate to suborbicular, acute but appearing filiform due to the enrolling margins, 1.5–3 × 1–2 mm, entire, membranaceous, finely nerved, castaneous, glabrous, equaling to shorter than the pedicel. Flowers 7–10 mm long (with extended petals and including the stamens), 3–8 mm apart, nutant-secund; pedicel 1.5–4 mm long, ca. 0.5 mm in diameter, strongly curved downward, glabrous; sepals broadly ovate, acute, 2–3 × 1.5 mm, membranaceous, finely nerved, greenish, glabrous; petals lanceolate, narrowly obtuse to acute, 5–6 × 1.5–2 mm, white, membranaceous, forming an apically acute flower bud before anthesis, strongly recoiled-recurved at anthesis and afterward, forming a corolla ca. 4 mm diameter; stamens exceeding the petals; anther narrowly oblong, 1.7–2 mm long, basifixed, base bilobed, apex acuminate, slightly to distinctly recurved at anthesis; pollen oblong-elliptic, sulcate, sulcus broad, without exine elements, sulcus margins sharply cut and bearing a distinct entire marginal area without microreticulate exine elements, exine microreticulate, muri very narrow; ovary ovoid, ca. 2.5 × 1.5 mm, green; ovules subcylindrical, long caudate; style longer than the ovary and exceeding the petals, ca. 4 mm long, white; stigma simple-erect, white, lobes ca. 0.7 mm long, apical margins crenulate. Capsules a subglobose basket-like structure hanging with the opening downward; seeds narrowly fusiform, bicaudate, ca. 1.3 mm long.

Additional specimens examined:— BRAZIL. Mato Grosso: Vila Bela da Santíssima Trindade, Serra de Ricardo Franco , Cachoeira dos Namorados, in rocks along the river on vertical rocky wall, 639 m elev., 14°55’03.43” S, 60°04’26.26” W, 13 July 2007, W.M. Kranz 212, cult. E. Leme 7144 (HB, RB 670708, RB 756772) ; Cachoeira Seca , on vertical rocky walls, 520 m elev., 14°54’55.39” S, 60°04’13.36” W, 13 July 2007, W.M. Kranz 213, cult. E. Leme 7145 (RB, HB) ; limite superior da mata de encosta, em paredão de rochas, 550 m elev., 15° S, 60° W, 22 March 1978, P.C. Windisch 1796 (RB); close to the border with Bolivia , July 1975, P.C. Windisch 1326 (HB) .

Distribution and habitat:— Fosterella windischii is growing saxicolous on sandstone rock outcrops and rock walls along rivers and near waterfalls ( figs. 14 A, B ), within the humid forest, in the region of the Parque Estadual Serra de Ricardo Franco, in the municipality of Vila Bela da Santíssima Trindade, in the west region of Mato Grosso state, close to the border with Bolivia, at 500–640 m elevation. It is also reported by Ibisch et al. (2008) and Peters (2009) from Bolivia, where it grows in the Parque Nacional Noel Kempff, not distant from Brazil’s border, at lower altitudes.

The region of Serra de Ricardo Franco is known by its rich biodiversity due to the confluence of different ecosystems, like the Amazonian, Cerrado, and Pantanal.

Etymology:—The name of this species honors its collector, the Brazilian botanist, Paulo Gunter Windisch, specialist in ferns and lycophytes.

Observations:— Fosterella windischii is an easy-to-identify species within Brazilian species, due to its very characteristic combination of morphological features: leaf blades abaxially completely covered by a thick and coarse layer of white, interwoven peltate, shortly fimbriate trichomes (type I), abaxial and adaxial faces strongly contrasting in color, margins inconspicuously and subdensely spinulose at the base, inflorescence with distinct third order branches, basal primary bracts longer than the stipe, which are densely and coarsely white lepidote, sepals greenish, petals strongly recoiled-recurved at anthesis and afterwards, white with acute apex, pollen with sulcus margins sharply cut and bearing a distinct entire marginal area without microreticulate exine elements.

Despite the 1600 km separating, in straight line, the Brazilian population of F. windischii , situated in the west region of Mato Grosso state, from the population of its close relatives, F. atlantica , in Minas Gerais state, their morphological proximity is evidenced by the trichome type on the abaxial surface of the leaf blades and petal curvature at anthesis and afterwards. However, F. windischii differs from it by leaf blades inconspicuously and subdensely spinulose at the base (vs. entire), inflorescence with distinct third order branches (vs. usually with second order branches and rarely with inconspicuous third order branches.), the basal primary bracts longer than the stipe (vs. many times shorter), being densely and coarsely white lepidote (vs. inconspicuously white lepidote to glabrescent), flowers longer (7–10 mm vs. 7–8 mm long), with longer pedicel (1.5–4 mm vs. 1.5–2.5 mm long), sepals larger (2–3 × 1.5 mm vs. 1.5–2 × 1–1.5 mm), greenish (vs. vinaceous toward the apex), petals white (vs. vinaceous at the central-apical portion mainly abaxially), apex acute (vs. obtuse), and forming a flower bud with acute apex before anthesis (vs. flower bud apically rounded), and pollen with sulcus margins sharply cut and bearing a distinct entire marginal area without microreticulate exine elements (vs. sulcus margins well defined but not sharply cut, bearing microreticulate exine elements decreasing in size, but without any entire marginal area).

According to Peters (2009), considering the non Brazilian species, F. windischii is similar to F. kroemeri Ibisch, R. Vásquez & J. Peters (2008: 189) , endemic to Bolivia, differing in its shorter and narrower leaf blades with more pronounced indumentum on the abaxial leaf blade surface, shorter peduncle and the less branched inflorescence.

8. Fosterella yuvinkae Ibisch, R. Vásquez, E. Gross & S. Reichle , Selbyana 23: 216. 2002. Type:— BOLÍVIA. Santa Cruz, Chiquitos, Santiago de Chiquitos, 18°20’10.8” S, 59°34’30.1” W, August 2001, S. Reichle P-SR1 (holotype LPB, photo!) ( figs. 15 A to D)

Description:— Plants acaulescent, flowering 44–55 cm tall, propagating by basal shoots. Leaves ca. 10 in number, all alike, fasciculate, subspreading, exceeding the peduncle; sheath inconspicuous, subreniform, ca. 20 × 35 mm, thick, green, abaxially and distally densely white tomentose with long multifilamentous-stellate trichomes, adaxially glabrous, margins bearing coarse white filamentous trichomes; blade narrowly linear-lanceolate, inconspicuously narrowed at the base, thin in texture throughout, slightly channeled, without or bearing a slightly thicker channel toward the base, apex long acuminate-caudate, 30–40 × 2.3–3.3 cm, green, finely nerved, abaxially densely white tomentose with long multifilamentous-stellate trichomes (type IV), adaxially glabrous including the inner ones, margins entire, slightly to distinctly undulate, glabrous. Peduncle erect, 15–18 cm long, 1.5–2.5 mm in diameter, green to reddish, glabrous; peduncle bracts 15–40 × 2–5 mm, narrowly subtriangular-attenuate, then caudate, erect, membranaceous, paleaceous, nerved, bearing filamentous trichomes, margins entire, equaling to longer than the internodes. Inflorescence shortly paniculate, with third order branches, erect, 15–35 cm long, to 10–18 cm in diameter, main axis slender, 1–2 mm in diameter, glabrous, green to reddish; primary bracts resembling the upper peduncle bracts, entire, 5–20 × 1–2 mm, equaling to distinctly shorter than the stipe; primary branches 8 to 14 in number, suberect-ascending, laxly arranged, sometimes 2 developing from the same node, 8–16 cm long, (13–)20–35-flowered; stipe slender, 10–20 × 1 mm, naked; rachis 0.5–1 mm in diameter, slightly flexuous, greenish to reddish, glabrous, the terminal branch not at all distinct from the lateral ones; secondary bracts resembling the floral bracts; secondary branches 1 to 2 in number, 3–8 cm long, bearing 9 to 18 flowers; floral bracts ovate, acuminate, 2–3 × 1.5 mm, entire, membranaceous, greenish to reddish, glabrous, equaling to exceeding the pedicel. Flowers 11–12 mm long (with extended petals), 3–10 mm apart, downward secund at anthesis; pedicel ca. 2 mm long, ca. 0.5 mm in diameter, strongly curved downward, green, glabrous; sepals ovate, acuminate, 2.5–3.5 × 2 mm, greenish except for the reddish apex, glabrous, membranaceus; petals linear-lanceolate, apex acute, 9–10 × 2 mm, strongly recurved at anthesis but not recoiled, erect afterwards, white, membranaceous, naked; stamens shorter than the petals, but exserted at anthesis; anther narrowly oblong, ca. 2 mm long, basifixed, base bilobed, apex acute, distinctly recurved to recoiled at anthesis; pollen broadly ellipsoid, sulcate, yellow, sulcus without exine elements, margins sharply cut, with a narrow to broad entire marginal area without microreticulate exine elements, exine microreticulate; ovary ovoid, green; ovules short caudate; style shorter than the petals; stigma conduplicate-erect, white, distinctly exceeding the anthers, lobes short, erect, margins inconspicuously crenulate. Capsules unknown.

Additional specimens examined:— BRAZIL. Mato Grosso do Sul: Rio Verde do Mato Grosso, road MS 427, to Alto Pantanal , ca. km 50, ca. 241 m elev., 19°57’45.3” S, 55°03’38.2” W, 17 September 2011, E. Leme et al. 8586 (RB 756858, RB 756728) ; Rio Verde de Mato Grosso, Serra da Pimenteira na Serra de Maracaju , 262 m elev., -19.129083, -55.060639 WGS84, 17 September 2011, A. Pott 16242 ( CGMS) .

Distribution and habitat:— Fosterella yuvinkae was found growing terrestrial or saxicolous on shaded sandstone walls (fig, 15 A), at about 250 m elevation, in the region of Rio Verde de Mato Grosso, Mato Grosso do Sul state. This is the first record of the species for Brazil, distant ca. 300 km from the Bolivian border, and ca. 200 km from the type locality of its close relatives, F. hatschbachii (in straight line). In Bolivia, it is reported from the Dpto. Santa Cruz, in the Prov. Chiquitos and Prov. Cordillera, where it grows in dry, deciduous lowland forests, along small ravines and riverbanks in higher elevated sites, 480–1200 m ( Peters 2009).

Etymology:—The epithet honors the Bolivian zoologist Yuvinka Gareca, for her ecological research in the area of Santiago de Chiquitos.

Observations:— Fosterella yuvinkae is closely related to F. hatschbachii and can be easily confused with it. However, it differs from it by leaf blades abaxially densely white tomentose with long multifilamentous-stellate trichomes (type IV) [vs. subpeltate, long filamentous trichomes (type V), which are more densely arranged], primary branches with a higher number of flowers (20–35 vs. 10–27 in number), sepals acuminate (vs. acute), and by the longer petals (9.5–10 mm vs. 7–9 mm long), white (vs. whitish or whitish with a pale reddish-wine colored central-apical portion), and acute apex (vs. obtuse and inconspicuously cucullate-emarginate).

When compared to the non Brazilian species, Peters (2009) indicates it is close to F. penduliflora , distinguished from it by the densely tomentose abaxial leaf surface, with trichomes type IV [vs. scattered lepidote with peltate, dentate trichomes (type III)], narrower petals, and the distinct habitat preference in Precambrian rocky outcrops within Chiquitano-Dry-Forest of Bolivia (vs. widely distributed in montane Andean forests in Bolivia, as well as in Peru and Argentina).

Figures

FIGURE 3. Photographs and SEM micrographs of trichomes on the abaxial surface of leaf blades.A–C. Fosterella atlantica (Leme 9223), interwoven peltate, shortly fimbriate trichomes type I, (bars: A = 10 mm; B = 200 μm; C = 100 μm). D–F. Fosterella windischii (Leme 7144), interwoven peltate, shortly fimbriate trichomes type I (bars: D = 10 mm; E = 400 μm; F = 100 μm). G–I. Fosterella bodoquenensis (Leme 8786), peltate, long fimbriate trichomes type II (bars: G = 5 mm; H = 200 μm; I = 100 μm). J–L. Fosterella batistana (Leme 7696), peltate, dentate trichomes type III (bars: J = 5 mm; K = 200 μm; L = 100 μm). M–O. Fosterela penduliflora (Teixeira 549), peltate, dentate trichomes type III (bars: M = 500 μm; N = 500 μm; O = 100 μm).

FIGURE 12. Fosterella penduliflora: Specimen (Teixeira 549) from Rondônia state, Brazil, deposited in the herbarium of the Missouri Botanical Garden (MO). Eletronic image obtained from herbarium’s website.

FIGURE 13. Fosterella penduliflora: Lectotype (Forget s. n.) from Peru, without further locality, deposited in the herbarium of the Royal Botanical Gardens, Kew (K). Eletronic image obtained from herbarium’s website.

FIGURE 14. Fosterella windischii from Vila Bela da Santíssima Trindade, Serra de Ricardo Franco, Mato Grosso: A. General view of the habitat at Cachoeira dos Namorados. B. Population at Cachoeira Seca. C–D. Specimens flowering in cultivation (Leme 7144). C. Habit. D. Flower details. E–G. Specimens flowering in cultivation (Leme 7145). E. Inflorescence details. F. Flower details. G. Habit. (A–B, photos by W.M. Kranz)

FIGURE 15. Fosterella yuvinkae from Rio Verde do Mato Grosso, Mato Grosso do Sul (Leme 8586): A. Population during dry season. B. Habit. C. Details of the branches. D. Detail of a raceme with open flowers.