Stachyphrynium latifolium (Blume) Schumann (1902: 49)

Niissalo, Matti A., Khew, Gillian S., Webb, Edward L. & Leong-Škorničková, Jana, 2016, Notes on Singaporean native Zingiberales II: revision of Marantaceae, with a new generic record and notes on naturalised and commonly cultivated exotic species, Phytotaxa 289 (3), pp. 201-224 : 209-214

3. Stachyphrynium latifolium (Blume) Schumann (1902: 49)

Basionym:— Phrynium latifolium Blume (1827: 37) ≡ Phyllodes laxifolia (Blume) Kuntze (1891: 695) . Note: Phyllodes Loureiro (1790: 13) is a rejected name.

Type:— INDONESIA. Java, Bantam [Banten province], without date, C. V. Blume s.n. (lectotype L! [ L 1486181], here designated).

= Hitchenia musacea Wallich ex Baker (1892: 225) ≡ Curcuma musacea Wallich (1832 : Cat. No. 6596), nom. nud. Type: SINGAPORE. Singapore, 1822, N. Wallich 6596 (lectotype K-WALL! [K001124261], designated by Suksathan & Borchsenius 2005, isolectotypes BR! [BR0000013330897]).

= Stachyphrynium cylindricum Schumann (1902: 49) ≡ Phrynium cylindricum Ridley (1899: 178) , non Roscoe (1828: table 40), nom. illeg. Type: MALAYSIA. Perak, Kwala Dipang, 1898, Ridley 9787 (lectotype K! [K000292270], designated by Turner 1998).

= Phrynium griffithii Baker (1892: 260) ≡ Stachyphrynium griffithii (Baker) Schumann (1902: 49) ≡ Phrynium spicatum Griffith (1851: 408) , non Roxburgh (1820: 5), nom. illeg. Type: MALAYSIA. Malacca, without date, W. Griffith s.n. (lectotype K! [K000292272], designated by Suksathan & Borchsenius 2005).

Rhizomatous, rosulate terrestrial herb to 1– 2 m. Leaves 2–4 per shoot; sheath 25–38 cm, pale green, light to dark brown near the margin, glabrous; petiole 30–105 cm, bright green, glabrous; pulvinus 4.5–6.0 cm; lamina oblong, 28–53 × 10–17 cm, apex acuminate, acumen 2.5 cm, base rounded to truncate, upper leaf surface dark green, glabrous, lower surface glaucous-green due to a waxy coating that is absent from one margin, where a bright green strip 1–2 cm wide is present, glabrous. Inflorescence radical, protected at the base with several narrow bracts increasing in side from 1–2 cm to 6–10 cm; peduncle 8–25 cm, glabrous, pale to yellow-green; bract at the base of the synflorescence 3–7 × 2.5–3.0 cm, with several basal thickenings (visible in dried material only); synflorescences spiciform, distichous, flattened, 8.5–21 cm; fertile bracts 8–17, each supporting one special paraclade, elliptic, obtuse with a very small acumen, 3.5–4.0 × 2.5–3.2 cm, pale green to yellow-green, glabrous; flower pairs 3–4 per special paraclade, associated prophylls, elliptic, semi-translucent light brown, glabrous 20–28 × 10–12 mm, sometimes with two keels, interphyll not always present, when present, ligulate, semi-translucent light brown, glabrous 17–22 × 2.5 mm; flower-pair shortly pedunculate (2 and 1 mm respectively). Flower ca. 3.7 cm long, with a jasmine fragrance; sepals three, free, subulate, 5 × 0.8 mm, semi-translucent white at the base, pale brown towards the apex, glabrous; floral tube 26–28 mm long, white, glabrous; petal lobes elliptic with acute apex, 9–11 × 4 mm, semi-translucent white, recurved; staminodial tube as long as the corolla tube; outer staminodes two, unequal in size, both spathulate, crumpled in bud, white, with a free part of 9 × 7 mm (larger) or 5 × 2.5 mm (smaller); hooded staminode slightly cucullate, white, yellow towards the apex, the free part 5 × 3 mm, with a minute appendix on one side, recurved, triangular, bright yellow, ca. 0.7 × 0.7 mm; fleshy staminode fused with fertile stamen for almost the entire length, curved but not very strongly tubular, white, with a free part of 5.5 × 3.6 mm, with a callose, hirsute appendix near base, and a keel-like appendix between the fleshy staminode and the fertile stamen; fertile stamen spathulate, similar in appearance to outer staminodes, 4.2–5.0 × 2.5–3.0 mm (measured from where it separates from the fleshy staminode), anther theca emerging at the point of separation of the fertile staminode and fleshy staminode, but fused to the appendix of the fertile staminode for its entire length, 1 × 0.7 mm; style with a free part 4 mm, curved to resemble a crescent, stigmatic cavity facing down at an angle ca. 1.2 mm in diameter; ovary cylindrical to weakly obovate, ca. 1.7 × 1.2 mm, cream-white, hirsute. Fruits (described from photographic material) elliptic, 30 × 18 mm, yellow-green, indumentum not seen; seeds 1–3, 25 × 9 mm, dark brown with rugose surface, arillate at base, aril white, extending into 2 curled subulate appendages, opening as the fruit dehisces, up to 13 mm.

Provisional IUCN conservation assessment:— Least Concern (LC). As currently circumscribed by Suksathan & Borchsenius (2005), this highly variable species has a broad distribution from Thailand through Peninsular Malaysia to Java and Borneo, and there are numerous recent collections across this range. Although it seems to tolerate some slight habitat disturbance, it is likely to have lost substantial parts of the habitat to forest clearance in the region. If the taxonomic status of this taxon changes, the conservation status must be re-assessed.

In Singapore S. latifolium is the most common Marantaceae , but the population size is fewer than 250 individuals and it is Endangered (EN) under criterion D.

Specimens examined:— SINGAPORE. Bukit Timah: 12 August 1889, J. S. Goodenough s.n. ( SING) ; 5 April 1890. H. N. Ridley 44 ( SING) ; 20 November 1899, H. N. Ridley 444 ( SING) ; 16 August 1983, H. Kennedy & E. P. Tay s.n. ( SING, 3 sheets) ; Fern Valley Trail, 17 October 1995, Eugene Tang & Hj. Sidek 1005 ( SING) ; Taban Loop , 27 June 1998, L. M. J. Chen LCMJ 244 ( SING) ; Taban Valley , 30 March 2005, J. Škorničková et al. SING 2005-54 ( SING, 3 sheets and spirit material) ; Tiup Tiup Path , 24 May 2011, J. Leong-Škorničková & H. D. Tran, SNG-84 ( SING) ; Tiup Tiup Path , 14 April 2015, J. Leong-Škorničková et al., SNG-335 ( SING) . MacRitchie: sector 43, 10 June 1992, S. E. Liaw et al. 1283 ( SING, SINU) ; Off Shinto Trail, 29 May 2014, J. Leong-Škorničková et al., SNG-184 ( SING), MacRitchie Nature Trail , 1 June 2016, M. A. Niissalo et al. SNG-346 ( SING) . Nee Soon: Nee Soon Freshwater Swamp Forest , in the forest on slope above stream, 26 June 2005, J. Škorničková et al., SING 2005-250 ( SING) . Pulau Seletar: 1891, J. S. Goodenough s.n. ( SING) . Sungei Buloh: 8 May 1890, J. S. Goodenough s.n. ( SING) ; 1894, H. N. Ridley s.n. ( SING) . Without precise location: 1822, N. Wallich 6596 ( BR, K) . Probably cultivated material: Botanic Gardens Jungle , 19 April 1962, Jumali bin Kafrawi 3434 ( SINU) .

Notes:— The species is morphologically uniform and the measurements are based on the most complete collections ( SING 2005-250, Eugene Tang & Hj. Sidek 1005, SING 2005-54; J. S. Goodenough s.n. 1890) as well as measurements taken from fresh material and in the field. While the species is fairly well represented in Singaporean herbaria, the only collections from Singapore in herbaria abroad are those by Wallich.

With some hesitation, we are following the very wide species delimitation of S. latifolium Suksathan & Borchsenius (2005) . The population in Singapore is identical to type material of S. griffithii , a name often applied to Singaporean material. We acknowledge that a revision of the living flowering material across the region, combined with DNA studies are needed to address if S. cylindricum and S. griffithii are indeed conspecific with S. latifolium , but due to lack of sufficient material, we are currently unable to test this. The species is fairly abundant in the small remaining primary forests in Singapore, both in freshwater swamp forests and the more humid parts of lowland dipterocarp hill forests; a few individuals are known near streams in secondary forests.

The species is easily identified even when sterile in Singapore due to a distinct glaucous surface with a darker wide margin on the underside of mature, but not old, leaves (fig. 3b, inset). While all recent voucher specimens are from Bukit Timah Nature Reserve, MacRitchie Reservoir and Nee Soon, the species occurs throughout the northern part of the Central Catchment (Upper Seletar, Upper Peirce, Ecolink), from where there are currently only silica dried collections at SING.

Typification:— Phrynium griffithii Baker : Suksathan & Borchsenius (2005) proposed a Griffith collection without a collection number from Malacca deposited at K as the lectotype and also indicated the presence of an isotype (= isolectotype) at K. There are three collections by Griffith from Malacca at Kew. From the collection style and coloration of the specimens it is likely that they are duplicates, although it is difficult to establish such a fact with certainty. Two sheets were part of Hooker’s herbarium and have a number of the late East India Company’s herbarium 5771 and 59771 respectively (the ‘9’ being likely a type error). The third sheet [ K 00292272], which was part of Bentham’s herbarium, is without a number. This sheet is therefore most likely the intended lectotype by Suksathan & Borchsenius. The other two collections present with a collection number are perhaps better considered syntypes.

Phrynium latifolium Blume : There are three Blume specimens at L. One of these matches the protologue with the vernacular name that is written on the sheet (“ pattot ”, L0041064!) and another matches the protologue locality data on the sheet (“ Bantam ”, L1486181!). The third specimen lacks annotations. As it is not clear if these three sheets are duplicates, we have selected the specimen with matching locality as the lectotype and we consider the other two sheets syntypes.

4. Stachyphrynium parvum Ridley (1910: 60) ≡ Phrynium parvum (Ridl.) Holttum (1951: 283) ≡ Stachyphrynium minus Ridley (1907: 59) , non Schumann (1902: 48), nom. illeg.

Type:— SINGAPORE. Reservoir, 1906, H.N. Ridley 12565 (lectotype SING! [SING0040953], designated by Holttum [1951], isolectotypes BM! [BM000617238], K! [K000292269]).

Rhizomatous, rosulate terrestrial herb 0.3–0.6(–0.9) m, forming large clumps. Rhizome creeping, with shoots distanced 1.5–3.5 cm. Leaves 1–4 per shoot (sometimes also with an additional laminate bract that acts as the lowest bract in the inflorescence); sheath 9–19 cm, green when young, soon drying to pale yellow-brown and later papery, light brown, tomentose; petiole 12–42 cm, medium green, glabrous; pulvinus 2–4 cm; lamina narrowly ovate to narrowly elliptic, 16–27 × 4–6 cm, apex attenuate, base shortly attenuate, upper leaf surface dark green, glabrous (including midrib), lower surface light green, glabrous, except the midrib which is puberulous. Inflorescence terminal, peduncle 10–21 cm (but sometimes the lowest bract replaced by a laminate leaf, in practice making the ‘peduncle’ a leafy shoot), erect (extending a further 1–8 cm after the lowest bract/stalked laminate leaf), glabrous, green to light brown; bract at the base of the synflorescence narrowly elliptic, 3.2–4.5 × 0.8–1.2 cm or replaced by a small leaf; synflorescence spiciform (almost capitate when heavily branched), alternate and distichous, 3.2–4.5 cm; first order branches 1–3, branches erect (sometimes spreading) distichous, 2.5–3.5 cm (measured from base to the tips of bracts), usually simple (order of branches no greater than two), fertile bracts 1–4 per branch, alternate and distichous, curled on sides (appears tubular), narrowly elliptic when opened, with acute apex, 1.7–2.5 × 5–7 mm, (green to) red-brown, hirtellous with very short glandular hairs and a few longer hairs near margins (hairs never flocculose, see notes below); flower pairs two per special paraclade, associated prophylls narrowly elliptic, glabrous, 15 × 5 mm, with two distinct keels along the entire length on the abaxial surface, with acute apex, interphyll, narrowly elliptic, glabrous, colour not recorded, 7 × 3.5 mm; flower pair sessile, but each flower with a 1.5 mm pedicel, bracteoles absent. Flower ca. 2.1 cm, fragrance unknown; sepals 3, free, subulate, 4.5–5.0 × 0.3–0.5 mm, colour not observed; floral tube 15 mm, incompletely fused for the upper 2 mm; petal lobes elliptic, obtuse, 5.5 × 2–3 mm, semi-translucent pale yellow-green, slightly darker towards the apex, reflexed; staminodial tube absent, but some of the staminodes fused to 0.5–1.0 mm beyond floral tube; outer staminodes 2, unequal, spathulate, apparently crumpled in bud (folding still visible in open staminodes), white, larger with a free part of 5.2–5.5 × 3.9–4.1 mm, smaller with a free part of ca. 4.5 × 3.5 mm; hooded staminode cucullate, entire, white, with a free part of 3.2 × 2 mm, without an obvious appendix (a simple notch ca. 0.5 mm can be seen on the side of the staminode); fleshy staminode curved towards the pistil at sides, roughly semi-circular in cross-section, more or less linear but with a very abrupt, blunt end, white, the free part 2.4 × 2 mm (but fused to the fertile staminode in the lowest 0.7 mm), with a white appendix on one side, increasingly protruding from the staminode towards the apex, ca. 1.2 × 0.3 mm; fertile stamen (including the fertile anther and appendage) white with a yellow apex, 3.1 × 1.5 mm, anther emerging at 1.2 mm, 1.5 × 0.5 mm; style with a 3.1 mm free part, enrolled like a crozier, with a trumpetlike opening facing down, collapsed on one side to give an appearance of a mouth-like opening with lower and upper ‘lip-like’ extensions. The upper ‘lip’ yellow with small hairy black calli, stigmatic cavity ca. 0.5 mm diameter; ovary 1.2 mm, hirsute. Fruits not collected.

Provisional IUCN conservation assessment:— Endangered (EN, category B1ab(iii)B2ab(iii)). EOO 1,005 km 2, AOO 24 km 2, measured using an online tool at http://geocat.kew.org. As circumscribed here, S. parvum is restricted to a small area in Singapore (two populations known) and adjacent Peninsular Malaysia. Based on revision of herbarium material, only one location is known from Johor (R.S. Holttum SFN 10294, SING!). It is only known from primary forests, which have been largely lost in Johor and Singapore.

The population in Singapore appears stable, but is very small. The species appears superficially abundant along small areas, but most individuals here seem to form clonal clumps and it is certain they represent fewer than 50 genetically distinct mature individuals. It is considered Critically Endangered (CR) under category D.

Singapore specimens examined:— SINGAPORE. MacRitchie: Reservoir Jungle, 26 October 1944, E. J. H. Corner s.n. ( SING) ; MacRitchie Reservoir, South side, 25 June 1949, J. Sinclair s.n. ( L) ; MacRitchie Reservoir, 3 November 2009, A. T. Gwee SING 2009-442 ( SING) ; MacRitchie Reservoir, MR 5, 18 February 2011, J. Leong-Škorničková et al. SNG-52 ( SING) (two sheets and spirit material) ( SING) ; Lornie Trail, 14 August 2014, J. Leong-Škorničková et al., SNG-189 ( SING) . Without precise location: Cluster 19, 11 May 1993, W. S. Chee 1802 ( SING, SINU) .

Notes:— Ridley described Stachyphrynium parvum from material collected in forest surrounding MacRitchie Reservoir, which remains the only locality for this species in Singapore. It belongs to a species complex that includes three other validly published names, Stachyphrynium sumatranum ( Miquel 1860: 616) Schumann (1902: 48) and Stachyphrynium borneense Ridley (1937: 204) and Stachyphrynium calcicola Poulsen & Clausager (2004: 162) . The described names only cover some of the diversity known from this complex and the group is in dire need of taxonomic revision. The plants in Singapore and at least one collection from Malaysia (north of Gunung Belumut [Johor], 22 May 1923, R. S. Holttum SFN 10294, SING!), are distinct in having a very long pulvinus (2–4 cm), secondary veins at an acute, ca. 30° angle at the middle of the leaf blade, and in having rather elongate, attenuate leaf-tips. The other two species described have a short pulvinus (0.9–1.5 cm), secondary veins at broader angle (> 50°) and acuminate leaf-tips (leaf tips are poorly visible due to curling in type material of S. borneense , but described as cuspidate-acuminate in the protologue). We are only aware of one collection from the main island of Sumatra (Loeboe Aloeng, not dated, 2037 HB, L!) and three more recent collections from offshore Sumatra in the Siberut Islands, not far from the type locality (Teiteibati National Reserve, Siberut Island, July 1992, J. J. Afriastini 1911, L!; Siberut, Sumatra, 26 February 2004, A.D. Poulsen et al. 2256, ANDA, BO, not seen; Siberut, Sumatra, 21 September 1924, C. Boden-Kloss 14544, SING!). We have not yet seen any duplicates of A.D. Poulsen et al. 2256, which was previously identified as S. sumatranum , but the other collections mentioned are very similar to each other. There appears to be some variability of the species in Peninsular Malaysia, with some collections appearing similar to S. sumatranum in leaf details (e.g. Sungei Semagot Kanan, 30 th mile Kota Tinggi-Mersing Road, 6 October 1963, J. Sinclair 10753, SING!) and some observed in the field probably represent new taxa ( JL-S, pers. obs.). Material of Stachyphrynium cf. sumatranum from Borneo are highly variable in terms of inflorescence length, bract shape and arrangement, and the indumentum which in some Bornean collections is clearly flocculose. We have not seen any material from Borneo that would be a good match for S. parvum in vegetative characters.

Although many specimens in this complex are now determined as S. sumatranum in various herbaria, including almost all specimens at SING, the synonymy has never been formally proposed in any publication.For the morphological reasons we have outlined above and our preliminary molecular evidence discussed below, we refrain from synonymising S. parvum with S. sumatranum (the latter having nomenclatural priority in this complex). We are of the opinion, that any synonymy in this complex should be based on comparison of living flowering material supplemented by DNA studies from across the entire distributional range of this complex as many important characters, particularly flower details, are lost in dried herbarium material. The yellow-green corolla lobes observed in Singapore material has not been reported elsewhere, but it is unclear if this is due to incomplete descriptions from those regions.