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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Billbergia Thunberg (Pl. Bras. 3: 30, 1821). Type: Billbergia speciosa Thunberg. — Bromelioideae — Lit: Smith & Downs (1979: 1975–2036, Fl. Neotropica); Barros & Costa (2008: monograph Rio de Janeiro State); Gaiotto F. & al. (2010: monograph Paraná State). Distr: S Mexico to Bolivia and N Argentina, with a centre of diversity in Brazil. Etym: For Gustaf Johan Billberg (1772–1844), Swedish botanist and zoologist.
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Incl. Anacyclia Hoffmannsegg (1833). Type: Anacyclia farinosa Hoffmannsegg [nom. illeg., ≡ Bromelia zebrina Herbert].
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Incl. Eucallias Rafinesque (1838) (nom. illeg., Art. 52.1). Type: Eucallias versicolor Rafinesque [nom. illeg., ≡ Bromelia zebrina Herbert].
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Incl. Jonghea Lemaire (1852). Type: Jonghea splendida Lemaire.
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Incl. Cremobotrys Beer (1854) (nom. illeg., Art. 52.1). Type: Bromelia zebrina Herbert.
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Incl. Helicodea Lemaire (1864). Type: Helicodea baraquinia Lemaire [typification according to L. B. Smith & Downs, Fl. Neotrop. 14(3): 1977, 1979].
Perennial terrestrial or epiphytic rosette plants, usually acaulescent, offsetting and with usually short stolons; Ros broadly funnel-shaped to tubular; L sheaths inconspicuous to distinct, L lamina narrowly to broadly linear-oblong, often with conspicuous pale cross-bands, tip often rounded tongue-shaped with a small mucro, margins coarsely to finely serrate or rarely entire; Inf simple or compound, peduncle erect or curved, floriferous part often arching to nutant; peduncular Bra usually large, thin, often bright red to magenta, overlapping; Fl large and showy, actinomorphic or slightly zygomorphic, sepals and petals forming a short to long epigynous tube; Sep lobes free, erect, glabrous to densely farinose or floccose-lanate; Pet free, at the basis with 2 scales, with a long claw and a somewhat broader, long but still narrow tip, ascending to recurved or tightly recoiled at anthesis; St usually exserted; Fil all free or those of the inner whorl adnate to the petal base to the height of the scales; Ov completely inferior; Sty longer than the stamens; Fr green to violet fleshy berries, globose, with persistent calyx.
A genus of some 60 species. Billbergia forms part of the higher core Bromelioideae in all recent molecular phylogenies. It is with the exception of 2 species strongly supported as monophyletic, and is closely related to Quesnelia and a small group of Aechmea species (Sass & Specht 2010). Evans & al. (2015) also found 3 species outside a well-supported core Billbergia clade. The genus is traditionally divided into 2 subgenera, Subgen. Billbergia (inflorescences simple or branched, peduncle often glabrous; petals recurved or slightly coiled) and Subgen. Helicodea (Lemaire) Baker 1889 (inflorescences simple, peduncle densely white-lanate; petals coiled), but the molecular phylogeny of Sass & Specht (2010) found no support for this division.
Succulence: Succulence is absent or ill-defined in most species of the genus, although many have distinctly coriaceous leaves and are native to semi-arid places. In cultivation, the degree of succulence of several species is strongly influenced by the watering and light regime, with more pronounced succulence under dry and bright conditions. Below, a very small selection of succulent-leaved species (all from subgen. Billbergia) is presented by way of example.
Horticulture: Most species have relatively short-lived flowers, making them unattractive for horticultural purposes. The only species more commonly seen in the trade is B. nutans, and several cultivars and selections are encountered. Formally named intergeneric hybrids are known with Aechmea (= ×Billmea), Cryptanthus (= ×Biltanthus, see separate entry in this handbook), Neoregelia (= ×Neobergia, see separate entry in this handbook), and Quesnelia (= ×Billnelia).
The following name is of unresolved application but is referred to this genus: Tillandsia osyana hort. ex K. Koch (1862) (nom. inval., Art. 32.1c).
B. brasiliensis L. B. Smith (Arq. Bot. Estado São Paulo ser. 2, 1: 105, 1943). Type: Brazil, Santa Catarina? (De Vos s.n. in Hort. Verschaffelt s.n. [[icono]: Rev. Hort. 41: 87, Fig. 21, 1869]). — Lit: Barros & Costa (2008: 1175–1176). Distr: Brazil (Rio de Janeiro, Santa Catarina?); ecology not described. – Fig. 1.
Incl. Helicodea leopoldii Hort. Verschaffelt ex Lemaire (1864) ≡ Billbergia leopoldii (Hort. Verschaffelt ex Lemaire) Linden (1869) (nom. illeg., Art. 53.1); incl. Billbergia ianthina hort. ex E. Morren (1871) (nom. inval., Art. 32.1c); incl. Billbergia nuptialis Hort. Makoy ex E. Morren (1871) (nom. inval., Art. 32.1c); incl. Billbergia kuhlmannii L. B. Smith (1950).
Acaulescent, to 80 cm tall, with short ascending rhizomes; Ros rather narrowly tubular, with 8–10 leaves; L sheath broadly elliptic, rigid, margins spiny, L lamina to 80 × 6–7 cm, basal parts with broad pale cross-bands, thick and fleshy (E. Gouda, pers. comm.), somewhat lepidote, esp. on the lower face, tip broadly acute to roundish, apiculate, margins with laxly arranged Sp, these brown, antrorse, 1.5–3 mm; Inf decurved, simple, floriferous part usually nodding, somewhat densely 28- to 30-flowered, densely white-farinose, peduncle 35–41 cm, white-farinose; peduncular Bra congested below the floriferous part, lanceolate, acute, rose-red; floral Bra minute, almost covered by the indumentum; Fl sessile, suberect, 65 mm; Sep very slightly asymmetrical, oblong, 10−17 × 6 mm, rose-red, white-farinose; Pet linear, acute, ± 50 mm, greenish-rose with dark blue-purple apical part, with 2 fimbriate scales, contorted or somewhat spirally recurved for a short distance only; St included; Fr not described.
Originally described from Santa Catarina, but without locality data. Barros & Costa (2008) report the taxon from Rio de Janeiro, and its occurrence in Santa Catarina is unlikely. Suspected as probable intersubgeneric hybrid by Smith & Downs (1979: 2007) and Reitz (1983: 485–486), but compared with the similar B. pyramidalis by Barros & Costa (2008).
B. euphemiae E. Morren (Belgique Hort. 22: 11, tt. 1–2, 1872). Type: Cult. BG Liège (Anonymus s.n. [LG, GH [photo]]). — Distr: Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro); epiphytic or lithophytic in forest. I: Smith & Downs (1979: 2013); Barros & Costa (2008: 1178); Machado & al. (2016: 138).
Incl. Billbergia euphemiae var. nudiflora L. B. Smith (1955); incl. Billbergia euphemiae var. saundersioides L. B. Smith (1955); incl. Billbergia euphemiae var. purpurea M. B. Foster (1957).
Acaulescent, to 60–70 cm tall, with short, stout, ascending rhizomes; Ros broadly tubular, with few leaves; L sheath large, narrowly elliptic, entire, L lamina 30 − 64 × (2.4−) 3–6 cm, thickly coriaceous, long-oblong, with white-lepidote transverse bands, tip rounded to broadly acute, apiculate, margins with laxly arranged small teeth; Inf decurved, 22–35 cm, floriferous part nodding to pendent, laxly to densely 6- to 20-flowered, simple, peduncle red, densely white-farinose; peduncular Bra suberect, large, lanceolate-elliptic, acute, whitish-red, upper Bra congested below the floriferous part, lax to somewhat dense; Fl spreading, 50–60 mm, very shortly pedicellate; Sep asymmetrical, 12–18 mm, narrowly elliptic, rounded, minutely apiculate; Pet linear, ± 30 mm, linear, obtuse, green, purple or dark blue towards the tip, with 2 fimbriate scales, never coiled; St included; Fr not described.
Divided into 4 ill-defined varieties by Smith & Downs (1979: 2012), but the differences appear to be minor, and var. nudiflora was synonymized already by Barros & Costa (2008: 1176). The leaves are quite succulent at least in some clones, and Pereira & al. (2011) record up to ± 50% water storage volume.
B. nutans H. Wendland ex Regel (Gartenflora 18: 162, t. 617, 1869). Type: BG Herrenhausen (Anonymus s.n. [LE?; [icono]: l.c. t. 617]). — Lit: Fagundes & Mariath (2010: fruit anatomy). Distr: Brazil (Paraná, Santa Catarina, Rio Grande do Sul), Uruguay, Paraguay, Argentina (Jujuy, Salta, Misiones, Córdoba); epiphytic or rarely lithophytic. I: Rauh (1990: 119: Fig. 88); Gaiotto F. & al. (2010: 89); Klein V. & Klein (2013: 76–77); Roguenant & al. (2016: 527). – Fig. 2.
Incl. Billbergia schimperiana Wittmack ex Baker (1889) ≡ Billbergia nutans var. schimperiana (Wittmack ex Baker) Mez (1896); incl. Billbergia nutans fa. rupestris Hassler (1919); incl. Billbergia nutans var. striata Reitz (1965).
Acaulescent, 40 (−50) cm tall, with numerous short stolons and forming dense somewhat untidy-looking clumps; Ros narrowly tubular, with 12–15 erect to somewhat arching leaves; L sheath oblong, 4 − 5 × 2 − 2.5 cm, subglabrous, L lamina linear to very narrowly triangular, to 60 (−70 or 100 cm under shady conditions) × (0.6−) 1−1.7 (−2) cm, somewhat canaliculate, leathery-coriaceous to distinctly fleshy, sparsely appressed-lepidote, margins near the base spinose-serrate, Sp 1 mm; Inf decurved, to 40 cm, appearing simple but composed of usually 1-flowered branches, peduncle very slender, glabrous, flowering part arching to drooping, few-flowered; peduncular Bra similar to the leaves, densely imbricate, basal Bra long-attenuate, green, upper Bra shorter, red, all grey-lepidote, longer than the internodes; floral Bra small, inconspicuous; Fl somewhat distichously arranged, sessile to shortly pedicellate; Sep erect, narrowly elliptic to ovate-lanceolate, acute, 15–20 (−27?) mm, rose with dark blue margins, towards the tip green with bluish blotch; Pet 35–46 mm, linear, obtuse, pale green with blue margins, with 2 coarsely serrate scales, tips pure green, spreading to recurved; St included; Fil of the inner series shortly connate with the petals; Fr ellipsoid, 35−45 × 12–14 mm, green, glabrous.
A naturally occurring form from Rio Grande do Sul with yellow longitudinal lines on the leaves was separated as var. striata. According to Gaiotto F. & al. (2010), B. schimperiana falls within the variability of the species, and recognition as separate taxon at any rank seems unwarranted. Guillot Ortiz & al. (2016) report the taxon as local neophyte in Spain.
References
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Eggli, U. (2020). Billbergia BROMELIACEAE. In: Eggli, U., Nyffeler, R. (eds) Monocotyledons. Illustrated Handbook of Succulent Plants. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-662-56486-8_76
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