Tanaka, N., Hannon, D.P. & Vislobokov, N.A. Kew Bull (2018) 73: 32. https://doi.org/10.1007/s12225-018-9754-5
- Possibly at risk from future limestone quarrying. Otherwise safe due to inaccessible habitat and probably low collection pressure as an ornamental or economic target. Data on the occurrence of this species in other areas is lacking, hence further investigations in wider areas are needed. Currently categorised as Data Deficient (DD) according to the IUCN Red List Categories and Criteria (IUCN 2012).
- Endemic to central Laos.
- On shaded dry-appearing limestone cliff; alt. 142 m.
- Morphology General Habit
- Lithophytic, glabrous, rhizomatous, evergreen, perennial herb
- Morphology Leaves
- Leaves produced before development of floriferous stem, usually 2 – 3 per year, basal, tufted, spreading-ascending, arched and distally recurved-pendent, persistent for 2 or more years, petiolate; petioles laterally compressed (slightly deeper than wide), C-shaped in cross-section, adaxially sharply and narrowly canaliculate, moderately flexible, to c. 18 cm long, to 5 mm wide; blades narrowly oblanceolate, proximally narrowly cuneate or long-attenuate, margins entire, broadly and weakly wavy, distally acute or acuminate, to c. 75 cm long, to 5.5 cm wide, subcoriaceous, midvein paler and acutely keeled abaxially, sunken adaxially, lateral parallel veins many, faintly raised on both sides (becoming prominent when dry), both surfaces weakly glossy
- Morphology Reproductive morphology Flowers
- Flowers many (c. 35 per spike), nearly synchronously blooming, sessile, ascending or horizontal, proximal ones often slightly descending, faintly odorous
- Morphology Reproductive morphology Flowers Androecium Stamens
- Stamens 6 on apical portion of perianth tube or on basal portion of perianth segments; anthers hippocrepiform, biloculate, thecae on low rounded (papillary) projection from perianth, (sub)lunate, distally contiguous, latrorse, 1 – 1.5 mm long, 1.1 – 1.5 mm wide
- Morphology Reproductive morphology Flowers Gynoecium Pistil
- Pistil unicarpellous, erect, ellipsoid-obovoid or obpyriform, usually shortly clawed towards base, 2.3 – 2.7 mm long; ovary and style externally undifferentiated, fleshy, 2 – 2.5 mm long, 1.4 – 1.7 mm in diam. (at subdistal broadest portion); ovary superior, occupying basal small portion of ovary-style column, uniloculate, c. 0.6 – 0.8 mm long and across, bearing 1 ovule on ventral lateral wall; ovule ellipsoid or pyriform, hemitropous with micropyle orientated downwards, 0.5 – 0.6 mm long; stylar portion broader than ovarian portion, 1.5 – 2 mm long, with a shallow longitudinal furrow on distal portion (probably corresponding to ventral suture of carpel); stigma reniform or depressed reniform, with lobation extending down (as furrow) to distal stylar portion, centrally depressed, 0.8 – 1 mm broad, densely papillulate
- Morphology Reproductive morphology Flowers Perianth
- Perianth clavate-tubular with truncate apex, 9 – 12 mm long, 3.5 – 5 mm in diam. at distal broader portion, proximally white and distally pale cream or (pale) fulvous (dull pale yellowish brown or beige), proximally gamotepalous and tubular, distally 6-lobed; tube 7.5 – 9.5 mm long, 3 – 3.2 mm in diam. in middle part, longitudinally few-angled externally, portions below stamens longitudinally ribbed adaxially; segments (narrowly) triangular or ovate-triangular, moderately or strongly incurved (upper 3 or 4 segments more abruptly inflexed), distally connivent, largely covering orifice of flower (each flower has 3 – 4 narrow slits mainly between lower segments), apex often inrolled, acuminate, acute or subacute, often with a few minute incisions, 2 – 2.5 mm long, 1.5 – 2 mm wide at base, adaxial medial portion ribbed longitudinally
- Morphology Reproductive morphology Fruits
- Fruits ellipsoid, somewhat clawed proximally, often slightly narrowed near apex, forming papillary apex, 1.9 – 2.1 cm long, 1 – 1.1 cm diam., pericarp fleshy, surface nearly smooth, green when young, orange at maturity, semi-glossy, 1-seeded
- Morphology Reproductive morphology Inflorescences
- Inflorescence a terminal dense spike, straight, cylindric, 3 – 5 cm long (rachis 2.5 – 4.5 cm long), 1.7 – 2 cm in diam. Floral bracts 2 (outer and inner) per flower, borne on rachis; outer bract larger than inner one (bracteole), clasping floral base from below, cucullate, broadly ovate or flabellate (transversely broadly ovate or triangular when opened up), light green, 2 – 3 mm long, distally often antrorse, apex (sub)acute, obtuse, rounded or subtruncate, distal margins membranous and often minutely irregularly denticulate; bracteole lying lateral to flower, ovate or broadly ovate, apex antrorse, dorsally midribbed, 1.3 – 1.8 mm long, 1 – 1.8 mm wide, proximally imbricate with bract
- Morphology Reproductive morphology Inflorescences Peduncles
- Peduncle (excluding inflorescence) (sub)terete, usually straight, spreading-ascending in same plane as attendant leaves, 6 – 20 cm long, ebracteate or with one small sterile bract (similar to fertile floral bracts) near apex, green
- Morphology Reproductive morphology Seeds
- Seeds obovoid-ellipsoid, 1.2 – 1.4 cm long, 0.8 – 0.9 mm diam., translucent and dingy ivory-coloured, glossy.
- Morphology Roots
- Roots cord-like, to 5 mm in diam. Sheath leaves (cataphylls) 6 – 9 at apical portion of stem, equitant, ovate to linear-lanceolate, apex acuminate, to 19 cm long, to c. 1 cm wide, withering earlier than fresh foliage leaves but persistent
- Morphology Stem
- Stem rhizomatous, cylindric, decumbent (distally pendant and free from cliff face in habitat), to 20 cm or more long (in habitat), 2 cm in diam., sometimes branched, externally medium brown, slow in growth (less than 1 cm in length in 7 years under cultivation), nodes a few mm apart Floriferous stem (including inflorescence) erect or inclined (sometimes declined), to c. 24 cm long, basally sheathed with 2 or 3, linear, acute sheath leaves 2 – 14 cm long
The perianth segments of Tupistra are usually excurved (Hooker 1902; Tanaka 2003a, c, 2010a, d; Vislobokov et al. 2014a; Averyanov et al. 2016), whereas those of two recently described species, T. breviscapa Aver. & N. Tanaka from central Laos and T. theana Aver. & N. Tanaka from central Vietnam (Averyanov & Tanaka 2012), are somewhat incurved or only slightly excurved. The two species also have small pistils and are lithophytic, and their habitats are near to the habitat of T. siphonantha. In addition, T. breviscapa has a single ovule in the locule, as aforementioned. Therefore, it appears that both T. breviscapa and T. theana are phyletically comparatively near to T. siphonantha. Since the floras of Laos and adjacent regions are still insufficiently explored (Newman et al. 2007, for Laos), there is a high possibility that new entities phyletically linking T. siphonantha and other congeners will be discovered from there in future.
Fruits of Tupistra are usually muricate and brown, purplish-brown (Wan 1984) or nearly black, while those of Rohdea are invariably smooth and red or orange (Tanaka 2010b). In the present study it became evident that T. siphonantha also has a smooth orange fruit similar to that of Rohdea. It is known that there is a strong relationship between reddish fruit and frugivorous birds as regards seed dispersal (Karasawa1978; Wheelwright & Janson 1985; Kobayashi et al. 2017). Hence it is likely that the orange or red berries of T. siphonantha and Rohdea evolved as a result of (probably parallel) adaptation to such avian frugivores for effective seed dispersal. In Rohdea, the fruits become ripe 7 – 10 months after anthesis. In T. siphonantha, they require approximately 9 – 10 months for full ripening, similar toRohdea. In a few other species of Tupistra , they fully ripen about 15 months after anthesis (Tanaka 2010a).
Tupistra siphonantha is also unique in having an inflorescence with flowers blooming almost simultaneously (Fig. 1). In contrast, in other species of Tupistra , inflorescences are indeterminate and flowering proceeds from proximal flowers to distal ones (Hooker 1902; Tanaka 2010a; Averyanov & Tanaka 2012; Vislobokov et al. 2014a; Averyanov et al. 2016). Having several features, such as synchronously blooming flowers, latrorse thecae prominently separated by a roundish protrusion of the perianth, and simple pistils with a single ovule, T. siphonantha is deemed as a highly specialised lineage among the species of Tupistra .
Tupistra is known to have a tricarpellary compound pistil of which the ovary is trilocular, bearing two ovules in each locule (Tanaka 2003a, 2010a; Vislobokov et al. 2014a; Averyanov et al. 2016). Our observations revealed that the pistil of T. siphonantha markedly deviates in organisation from that of its congeners. In T. siphonantha, the stigma and the distal part of the ovary-style column has a longitudinal lobation (furrow) that appears to correspond to the ventral suture of a carpel, and the ovary is unilocular, bearing a single ovule on the ventral lateral wall (Fig. 2R). No other carpellar structures, including rudimentary ones, were found. The fruits consistently contain only one seed (Fig. 1). These observations indicate that the pistil of T. siphonantha is unicarpellous (simple). Presumably, this simple pistil developed secondarily through reduction in the number of carpels and ovules from the tricarpellary pistil possessed by all other congeners. In connection with this, it is noteworthy that the ovary of T. breviscapa Aver. & N. Tanaka is trilocular but contains only one ovule in each locule (Averyanov et al. 2015).
Having markedly inflexed perianth segments, small pistils, and smooth orange fruits, Tupistra siphonantha resembles species of Rohdea (Tanaka 2003b, 2010b, Averyanov et al. 2014). However, it agrees more with Tupistra in having slender petioles, creamy white, beige or fulvous perianths (Fig. 1) and sessile anthers (Fig. 2E & F). In Tupistra , compared with Rohdea, the petiole is consistently slender and elongate relative to the blade (vs relatively short and widely decurrent), perianth colour is purplish, whitish, beige, or fulvous (vs green or greenish at early and middle stages of flowering) and anthers are mostly sessile or subsessile (vs mostly distinctly stipitate). Further, T. siphonantha and (other species of) Tupistra share an ovary strongly confined to the basal small portion of the pistil and a style as broad as or broader than the ovary portion (Fig. 2). This indicates that T. siphonantha is closer to Tupistra than to Rohdea of which the ovary occupies a relatively large part of the pistil and the style is consistently narrower than the ovary (Averyanov et al. 2014; Hooker 1902; Tanaka 2003a, 2010a, b; Hu et al. 2013; Vislobokov et al. 2014a).
The specific epithet refers to the tubular flowers.Differs from all other members of Tupistra by the synchronously blooming flowers, clavate-tubular, cream-white or fulvous perianths with strongly incurved segments, latrorse anthers with thecae prominently separated by a round protrusion from the perianth, and very small, simple pistil bearing a single ovule in the locule.
In Aspidistra and Rohdea, entomophilous flowers are reported (Migliorato1910; Vislobokov et al. 2014b). The flowers of Tupistra siphonantha may also be zoophilous (probably entomophilous), judging from their perianth colour, fragrance, and floral structure. The perianth segments of T. siphonantha are strongly incurved and distally connivent, largely covering the orifice of the flower during anthesis. The flowers have 3 – 4 narrow slits between the lower, adjoining perianth segments (Figs. 1, 2) and it is highly probable that small pollen-vectors enter the floral interior through the slits and pollinate the stigma while moving around inside the perianth.
Flowering: (late Oct. –) Nov. – Jan. (– Feb.) (under cultivation). Fruiting: late Sept. onwards (under cultivation).
- Type: Laos, Khammouane Province, D. P. Hannons.n. (holotype HNT!; isotype K!).
First published in Kew Bull. 73(3)-32: 2 (2018)
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Kew Backbone Distributions
The International Plant Names Index and World Checklist of Selected Plant Families 2022. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0
Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2022. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0