Helonias umbellata (Baker) Tanaka (1998a: 108)

Tanaka, Noriyuki, 2019, Taxonomy, evolution and phylogeography of the genus Helonias (Melanthiaceae) revisited, Phytotaxa 390 (1), pp. 448-450 : 448-450

9. Helonias umbellata (Baker) Tanaka (1998a: 108) ( Figs. 3D, E , 21 , 22 ).

≡ Heloniopsis umbellata Baker (1874: 278 , ‘ Heleniopsis ’; 1879: 460). ≡ Sugerokia umbellata (Baker) Koidzumi (1930: 95) . ≡ Hexonix umbellata (Baker) Wang & Tang (1949: 114) . Type:― TAIWAN (Formosa). R. Swinhoe s.n. (Holotype: K-001045008*!).

= Heloniopsis acutifolia Hayata (1920: 144 . f. 53). ≡ Sugerokia acutifolia (Hayata) Koidzumi (1930: 95) . Japanese name: Taiwanshôjôbakama ( Koidzumi 1930). ≡ Hexonix acutifolia (Hayata) Wang & Tang (1949: 113 , ‘ acutiflora ’). Type:― TAIWAN. [Ilan Hsien] Bonbonzan, 10 May 1917, B. Hayata & S. Sasaki (Holotype: TI!).

= Heloniopsis arisanensis Hayata ex Honda (1938: 1679) . ≡ Sugerokia arisanensis (Hayata ex Honda) Koidzumi (1939: 53) . ≡ Hexonix arisanensis (Hayata ex Honda) Wang & Tang (1949: 114) , ‘(Hayata)’. Typ e:― TAIWAN. [Chiayi Hsien] Arisan [Alishan], 2500 m, Junio 1914, U. Faurie 942 (Holotype: TI!. Isotype: BM-1118049!; P-01762777*!).

= Heloniopsis taiwaniana Ying (1980: 222 , f. 108 on p.67). Type:― TAIWAN. Taipei Hsien, Urai , Mt. Tatungshan , 20 April 1976, S.S. Ying 3615 (Holotype: NTUF-F00008243*!).

– Heloniopsis orientalis (Thunb.) Tanaka var. yakusimensis auct. non (Masam.) Ohwi: Masamune (1957: 103), p.p.

– Ypsilandra thibetica auct. non Franch.: Hsu et al. (2011: 99–104).

Chinese name:―(Taiwan) humahua.

Japanese name:―Hime-shôjôbakama ( Hayata 1917), Arisan-shôjôbakama ( Honda 1938), Shima-shôjôbakama ( Kawakami 1910).

Description:―Rhizome subcylindrical, to 1.7 cm long, 1 cm in diam. Roots filiform, ca. 1 mm in diam. Leaves persistent for 1–2 year, narrowly spatulate or oblanceolate, 7–16 cm long, 1–2.2 cm wide, tapering to linear petiole to ca. 6 mm wide, margin entire (not undulate), apex short acuminate or obtuse, apiculus (0.5–) 1–2 mm long, somewhat thick in texture, green, glossy, midvein raised abaxially. Flowering stem erect, fistulose, ca. 2–17 cm long at anthesis, elongating to ca. 29 cm long in fruit; peduncle 2–20(–28) cm long incl. in fruit, 1.5–5.5 mm in diam. in middle; scale-like leaves on peduncle 4–10 (excl. basal ones), narrowly elliptic or oblong, acuminate, to 23 mm long, submembranous; inflorescence umbellate or sub-umbellate, rarely racemose, rachis usually to 1.1 cm long (excl. raceme); pedicels ribbed, (3–)6–14(–18) mm long, whitish, sometimes tinged pink, ebracteate, 1 or a few proximal pedicels sometimes bracteate. Flowers 1–21, funnelform, often secund, strongly fragrant, protogynous. Tepals 6, oblong-oblanceolate or spatulate, 8–12 mm long, 2–3.7 mm wide, apex obtuse to acute, 3- or 5-veined, white or pale pink, not imbricate with adjoining tepals proximally, adaxially canaliculate toward base; proximal margins usually slightly revolute; proximal submarginal portions raised (ridged), sometimes minutely connate basally to adjoining tepals (connate portion to 0.3 mm in height); nectary at base concave adaxially; base of tepal affixed to receptacle (apex of pedicel) for ca. 1 mm in length. Stamens 6, equaling or slightly longer than tepals, obliquely divergent; filament arising from (adaxial) rim of nectary of opposing tepal, filiform, proximally scarcely dilated, (4–) 6–14 mm long, white or pale (purplish) pink, not adnate to opposing tepals; anthers basifixed adaxially, slightly extrorse, white or pale (bluish) purple, narrowly ovoid, usually 1.3–3.1 mm long, base sagittate, apex obtuse, (sub)bilocular with vestigial apical confluence between thecae; pollen white or creamy. Pistil 1, exceeding stamens and tepals; ovary sessile or short stipitate, whitish, pale green or purplish pink, globose-ellipsoid or obovoid, obtusely trigonous, slightly trisulcate, 1.5–2.8 mm long, 1.8–3 mm across, apex emarginate-concave; ovules numerous, multiseriate on central axile placentae; style terete, often slightly broadened above, 5–19 mm long, white or purplish pink; stigma thick discoid, often slightly trigonous, 1.3–2.4 mm across, 0.5 mm thick, white or pale purple. Capsules broadly obpyramidal (obcordate or obdeltoid in lateral aspect), apex depressed in center, tripartite, 4–7 mm long, 6.5–10 mm across; lobes mitriform or ovoid-pyriform, divergently protruding. Seeds numerous, linear-fusiform, testa whitish, 3.4–5 mm long, 0.2–0.4 mm wide; body of seed narrowly fusiform, 1.2–1.8 mm long, 0.2–0.3 mm wide, brown.

Additional specimens examined (see also Tanaka 1998a):― TAIWAN. Hsinchu Hsien: Chienshih Hsiang , ca. 2000 m, 20 February 2002, fl. Y.-Y. Huang 947 et al. (MAK-386385) . Ilan Hsien: Mt. Taiping , 4 March 1967, fl., M.T. Kao 6796 (TI); ibid., 1750m, 22 May 1949, fr., H. Keng 1217 (TAI-029039, PH-66762) . Miaoli Hsien: Nanchuang Hsiang, Fengmeihsi , ca. 1500–1700 m, 7 January 1994, fl., T.Y. Liu et al. 358 (HAST-40520*, BM-001118048) . Nantou Hsien: Ching-shui-kou, 18 February 1959, Huang, Kou & Kao 938 (TAI-029016*); Hsinyi Hsiang , ca. 1500 m, 12 February 2003, fl., C.- I. Peng 19224 (HAST-93318*) . Taipei Hsien: Shichiseizan ( Mt. Chihsing ), 3000 ft, 25 January 1932, fl., N. Fukuyama 3015 (TNS-59639); Shihting Hsiang, ca. 200–500 m, 17 February 2008, fl., P.-F. Lu 15304 (HAST-123134*); in monte Taitun, ad 600 m, 7 mai 1903, U. Faurie 537 (MAK-137680, P-01762775); Yangminshan to Chihsing, 600–1100 m, 26 February 1961, fl., T. Shimizu 11117 (A, TI); Daitonzan ( Mt. Tatun ), 20 April 1932, fr., S. Sasaki (TNS-321209) . Taichung Hsien: Leeshan , 20 April 1962, fr., C.S. Feung & M.T. Kao 4554 (TI); 710 logging tract, 1900–2000 m, 1 April 1993, fl., J.C. Wang et al. 8094 (HAST-27016) . Taitung Hsien: Taito (Taitung), Sinsuiei ( Chinshuiying ), 10 February 1925, fl., S. Sasaki (TNS 321217) . Taoyuan Hsien: Fuhsing Hsiang , ca. 1840 m, 11 Apr. 2002, fr., W.-C. Leong 2881 et al. (MAK-386590) . Pingtung Hsien: Kosyun (Oluanpi), Garanbi ( Garambi ), 29 March 1932, S. Sasaki 40061 (TAI-250118*) .

Distribution:― Taiwan ( Fig. 29-U ).

Habitat:―Shady and semishady moist rocks and slopes at elevations usually 200–2500 m.

Conservation status:―The species is comparatively widespread, and may be assessed as LC according to the IUCN Red List Categories and Criteria (2001).

Flowering:―Usually late January–March (to June at high elevations).

Ripening:―April–May(–June).

Remarks:― Hsu et al. (2011) reported Ypsilandra thibetica as being in Taiwan, but the plant on which the report is based appears to be a form of Helonias umbellata (see also the remarks under H. thibetica ). Having a raceme, their plant may look like H. thibetica , but other characters such as anthers (e.g. Fig. 2E, H in their paper, appearing subbilocular) and stigmas (e.g. Fig. 2E, G in their paper, appearing slightly lobed or trigonal and slightly larger) agree with H. umbellata . Further, the karyotype in their paper ( Fig. 4 ) does not agree with H. thibetica but rather with H. umbellata (N. Tanaka unpublished data). In Helonias , racemes are considered to be more primitive than umbels (Part III-2-2), so forms with racemes of H. umbellata may represent relict ancestors. It is also possible that racemose inflorescences were acquired secondarily from umbels. Further studies are needed to resolve this question. Helonias umbellata has tepals sometimes minutely connate to adjoining ones basally ( Fig. 22D ; Tanaka 1997b), suggesting a close relationship between H. umbellata and ser. Heloniopsis . Helonias umbellata occurs from hillsides to high peaks on Taiwan, and the habitats are not necessarily close to streams, indicating that the plant can tolerate diverse environmental conditions. Helonias kawanoi and H. leucantha , in contrast, are mostly confined to moist places along or near streams at low elevations on small, subtropical islands.

Figures

FIGURE 2. Diagram of phenology of rosette leaves and flowering stems (scapes) in Helonias. Phenological transition of rosette leaves (L) and flowering stem (F) in their lifetime (from bud formation to death) is schematized.Approximate magnitude of growth rate of two organs is expressed by line or shape of ellipse or lance (expanded portion). State of organs showing static, increasing or decreasing growth rate is represented respectively by line, or by expanding or narrowing (closing) portion of an ellipse or lance. Broken line represents period during which older leaves die, although their death is usually concentrated in growing season (April to July or August). In H. bullata, for example, leaf bud (see L-row) is initially formed in summer of year 0, it stays dormant until coming spring, and then develops rapidly in spring to summer (expanded open portion) of year 1. Mature leaves wither in growing season of following year (broken line in year 2). Flower bud of H. bullata (see F-row) is initially formed in fresh rosette (in L-row) in summer of year 1; like leaf bud it stays dormant until next spring, and then develops rapidly in spring (expanded shaded portion) of year 2. Flowering stem dies around summer of same year (closing shaded portion). Leaf bud (see L-row) of H. kawanoi, H. leucantha or H. umbellata after formed soon starts to sprout in late summer onwards (of year 0), although its growth appears more or less suppressed in subsequent cooler seasons (*). Phenological progression of flowering stem of H. kawanoi (F-row) tends to be lagged (**) when cultivated under lower temperature (in Tokyo) than in natural habitats.

FIGURE 3. Young slightly open flowers of Helonias thibetica (A–C; material T-1) and H. umbellata (D, E; U-1), showing protogyny. A. Flowers with precociously protruding pistil. B, D. Flowers similar in stage to A with frontal tepals removed to show indehiscent anthers. C, E. Closeup of densely papillulate mature stigma from flower in B and D, respectively.

FIGURE 4. Helonias bullata (material Bu-1; photos A–E on 28 April 2014, F on 13 June 2002). A. Habit and boggy habitat. B. Young opening flowers with indehiscent anthers. C, D. Young flowers visited by a Bombus species sucking nectar.E. Flowers visited by a halictid bee. F. Fully dehisced, ascending capsules. All photos taken by Jim Fowler.

FIGURE 21. Helonias umbellata. A. Young slightly open flowers with protruding pistils (material U-1; photo on 3 March 1989). B. Flowering plant (U-2; 14 March 1989). C. Umbel (U-1; 30 March 1992). D. Plant with fresh leaves (light green) and leaves developed in preceding year (darker green) grown in greenhouse during winter (U-3; 14 March 2015). All photos at Hachioji, Tokyo.

FIGURE 22. Helonias umbellata. A–E: material U-1. F: U-4 (in spirit). A. Flower with two frontal tepals and stamens removed. B. Closeup of proximal interior of flower. C. Vertical section of flower with ovary shortly stipitate at base. D. Proximal interior of partial flower. Minutely connate portions between adjoining tepals arrowed. E. Proximal part of flower in cross section. F. Distal part of flower with anthers showing apical confluence between thecae (arrowed). Scale at B also for C–E.

FIGURE29.Distributionofsixspeciesbelongingtotwoseriesofsubsect.HeloniopsisinEAsia.B:Heloniasbreviscapa.Ka:H.kawanoi.Ko: H.koreana.L:H.leucantha.O:H.orientalis.U:H.umbellata.Ser.UmbellataecomprisesKa,LandU.Ser.HeloniopsisconsistsofB,KoandO.