Helonias thibetica (Franch.) Tanaka (1998a: 106)

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

6. Helonias thibetica (Franch.) Tanaka (1998a: 106) ( Figs. 1C, D , 3A–C , 13–14 ).

≡ Ypsilandra thibetica Franchet (1887 – 1888: 94, t. 17; 1888: 132, t. 17). Type (lectotype, designated here):― CHINA. Thibet Oriental [Sichuan]: Province de Moupin, Mars 1869, A. David s.n. (P-00282689*!). Isolectotype: K-000400237*!, P-00730550*!, P-00730551*!

= Ypsilandra cavaleriei H.Lév. & Vaniot in Léveillé (1905: 375). Type:— CHINA. Kouy-Tchéou [Guizhou]: Environs de Tou-chan , dans la montagne, 3 Juin 1899, J. Cavalerie 2638 (Holotype: P-01724137*! Isotype: E-00902989!, K-001235139!―both (in fruit) appear to be erroneously labeled as Cavalerie 1291 that is in flower).

= Ypsilandra thibetica Franch. var. angustifolia Wang & Tang (1943: 106) . Type:— CHINA. Szechuan [Chongqing]: Su-tchuen oriental, Tchen-kéou-tin [Cheng-kou Xian], Juin, R.P. Farges 788 (non ‘188’). (Holotype: PE-00035458*! Isotype: P-00730547–9*!, P-00282693*!, PE-00035457*!, K-000400238*!).

=? Ypsilandra kansuensis Zhao & Peng (1987: 57 , f. 1). Type:— CHINA. Gansu: Zhouqu Xian, Luojiayu , elev. 2060 m, 4 April 1980, Z.- J. Cao & R.-N. Zhao 550564 (Holotype LZU n.v.). Paratype: CHINA. Gansu: Zhouqu Xian, Luojiayu, elev. 2000 m, 11 June 1982, R.N. Zao 552501 (LZU n.v.).

Chinese name:―Yaruihua.

Japanese name:―Chibetto (Tibet)-shôjôbakama (nov.), Chûgoku-shôjôbakama (meaning Chinese Helonias ).

Description:―Rhizome subcylindrical, closely annulate with many scars, to 7 cm long, to 1.7 cm in diam. Roots filiform, densely pubescent, to 1.5 mm in diam., white. Leaves hysteranthous, main fresh rosette leaves usually 6– 13(–18), persistent for 1–2 years, spatulate, narrowly spatulate or oblanceolate, 7–20(–32) cm long, 0.7–2.5(–4) cm wide, tapering to linear petiole, apex acute or acuminate, apiculate, apiculus 0.3–1.3 mm long, moderately thick, soft, glossy (especially when young), entire, abaxial midvein raised. Flowering stem 6.3–21.5 cm long at anthesis, elongating to 52 cm in fruit; peduncle longitudinally ribbed, 4.5–7(–17) cm long at anthesis, to 36.3 cm long in fruit; scale-like leaves on peduncle several to many (to ca. 15; excl. basal ones), subulate, (narrowly) lanceolate, oblong or oblanceolate, to ca. 4 cm long, proximal scale-like leaves herbaceous, distal ones submembranous, apex acuminate or acute; inflorescence racemose, rachis 1.8–4.5 cm long at anthesis, becoming 5.8–15.7 cm long in fruit; pedicels 3.5–13 mm long at anthesis, 4–22 mm long in fruit, ebracteate, white, sometimes pale purplish, proximal few pedicels sometimes bracteate. Flowers 9–46, often secund, funnelform, ca. 0.8–2 cm across, strongly fragrant, protogynous. Tepals 6, ascending, spatulate, oblong-oblanceolate or narrowly elliptic, slightly canaliculate toward base, margins slightly revolute proximally, nectary at base slightly concave (or nearly flat), apex obtuse or subacute, 6.5–12 mm long, 1.6–3(–4.5) mm wide, 3–5(–7)-veined, white, sometimes pale cream or pinkish, turning yellowish to greenish at late and after anthesis; basal part slightly inflated abaxially. Stamens 6, usually exserted beyond tepals; filaments ascending, filiform, straight, (7–)11–15(–17.5) mm long, white; anthers basifixed, ovate or elliptic-ovate, bilobed at base, rounded at apex, unilocular, white with purplish tint, grayish purple, or dark (bluish) purple, (0.8–) 1.5–2.1 mm long, peltate at early stage of dehiscence; pollen (slightly purplish) white. Pistil 1, (8–) 15–21 mm long, usually exceeding stamens and tepals; ovary globose or obovoid, trigonous, trisulcate, borne on short gynophore-like floral axis (receptacle), apex emarginate, depressed in center, 1.5–3 mm long, 1.5–3 mm across, (pale) green or pale brownish green; ovules numerous, usually 8- or 9-seriate on central axile placentae; style filiform, (5–) 10–22 mm long, white or slightly purplish pink; stigma small, often only slightly broader than style, (depressed) capitate or discoid, unlobed, 0.3–1.2 mm across, white. Capsules broadly obpyramidal, broadly cordate laterally, tripartite, 3–6 mm in height, 5.5–14 mm across; lobes subconic or pyriform, obliquely or horizontally divergent, often slightly recurved distally. Seeds coated with whitish testa, narrowly fusiform, 4.5–7 mm long, 0.3–0.5 mm wide in middle, narrowly tailed at both ends; body of seed narrowly fusiform, light brown, 1–2 mm long, 0.3–0.4 mm wide.

Additional specimens examined:― CHINA. Chongqing: Nanchuan, elev. 2000 m, 9 May 1986, Jinfoshan Kaocha-dui 0127 (PE-01867400*); Su-tchuen oriental, Tchen-kéou-tin, R.P. Farges s.n. (K-000400236*). Gansu: Bai-shui-jiang Caiji-dui 4963 (PE-01823466*); Dangchang Xian, elev. 3000 m, 25 July 1951, T.P. Wang 14561 (PE-00593854*); Wen Xian, elev. 1600 m, 16 May 2007, Bai-shui-jiang caijidui 4963 (PE-01823466*); locality unclear, 8500 ft, 16 May 1916, R. Farrer & W. Purdom 68 (E-00904006*). Guangdong: Ruyuan, June 1933, fr., X.P. Gao 9117 (PE-00593853); Ruyuan, 29 April 1934, S.P. Ko 54199 (PE-00035448*). Guangxi: Chuanhsien, 14 May, 1936, Z.S. Chung 81626 (PE-00593967*); Rongshui Xian, elev. 1000 m, 2 May 1989, Beijing dui 892174 (PE-02018211*); Xing’an Xian, 24 May 1979, G.Z. Li (IBK-188673*). Guizhou: Jiangkou xian, 1100 m, 26 May 1964, Zhang et al. 400780 (PE-01318236*); (Kouy-Tcheou) Pin-fa, montagnes, lieuz humides, 10 Mars 1909, fr., J. Cavalerie 1291 (E-00115518, E-00902989*, P-00730553*); (Kweichow) June 1905, J. Esquiroll 440 (E-00902988). Hunan: Hongjiang shi, 1954, Li 1723 (PE-00593855*); Xinning Co., 1200 m, 6 April 1995, fl., L.-B. Luo 0601 (E-00071276*, TNS); Xinning Xian, 800 m, March 1985, fl., Luo 2034 (PE-00593856). Sichuan: Leibo Xian, Zhongshan, 1400–1800 m, 9 June 1959, fr., Z.T. Guan 8432 (PE-00593934); Mt. Omi, E.H. Wilson 5221 (K); Omei-hsien, Mt. Omei, 2 May 1041, fr., W.P. Fang 16781 (A); ibid, 23 Mar. 1941, fl., W.P. Fang 18232 (A); ibid., 27 February 1940, fl., W.P. Fang 13849 (A-255367); Mt. Omei, Tachengssu, 2300 m, 19 May 1939, H.C. Chow 9825 (A); Tian-quan, 14 April 1953, H.L. Tsiang 33845 (PE-00593886); Tien-chuan-hsien, 2700 m, 25 May 1936, K.L. Chun 2649 (BM-001118046); Tien-chuan-hsien, 1360 m, 9 April 1936, K.L. Chun 2301 (BM-001118044, E-00904007*); Western China. Limestone cliff, E.H. Wilson 4676 (K, TI); West Szechuen and Tibetan frontier, chiefly near Tachienlu, 9,000 –13,500 ft, fl., A.E. Pratt 766 (BM-001118045; P-00282691*). Yunnan: Ku-long-tchang, elev. 800 m, Fevrier, fl., Maire s.n. (L-1470205*, P-01724143*); Suijiang Xian, elev. 1600 m, Z.H. Hu 1351 (PE-18867402*).

Distribution:― China: Chongqing, Gansu (S), Guangdong (NW), Guangxi (N, NE), Guizhou (Central, S), Hubei (SW), Hunan (S, W), Sichuan (Central, SE), Yunnan (NE) (some data based on literature) (Fig. 15-T).

Habitat:―Shady moist slopes at elevations ca. 800–3000 m.

Conservation status:―The species is comparatively widespread, and assessed as LC according to the IUCN Red List Categories and Criteria (2001). However, local populations near human habitation may be more or less subject to disturbance and threats.

Flowering:―February–March.

Ripening:―May–June.

Remarks:― Ypsilandra cavaleriei was distinguished from Y. thibetica by Léveillé (1905), and this treatment was followed by Chen (1980) and Chen & Tamura (2000). In Helonias thibetica , specimens from the southern part of the range (Guizhou, Guangxi, Guangdong, and Hunan) tend to have somewhat smaller flowers than those from the northern part (Sichuan, including the type), and correspond to Y. cavaleriei . However, the specimens corresponding to Y. cavaleriei have no other particular difference from the typical form of H. thibetica . Chen (1980) and Chen & Tamura (2000) reported that the two species are distinguishable by the difference in the relative length of ovary lobes to the entire ovary. However, this proportion varies with the progression of flowering and fruiting, hence it seems inappropriate to apply this character for distinguishing them. Accordingly, it appears more reasonable to regard Y. cavaleriei as a local race of H. thibetica , as previously suggested in Tanaka (1998a). I have not yet seen any type

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mesostyla

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var yunnanensis

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H

:

Ym

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thibetica

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H

:

T

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parviflora

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H

:

P

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jinpingensis

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H

:

J

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alpina

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H

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A

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Asia in

Ypsilandra

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subsect of species five of ranges geographic yunnanensis .

Approximate

. var

15

.

yunnanensis material of Ypsilandra kansuensis , but a specimen collected by Farrer and Purdom (no. 68, E-00904006*!) in Gansu province, China, may represent it. Ypsilandra kansuensis differs from Helonias thibetica by its stamens equaling (vs. exceeding) the tepals. There seem to be no other particular features characterizing it. To more clearly elucidate their relationship, it is desrable to study variation of their stamen length based on more samples especially from regions near the type locality of Y. kansuensis . The occurrence of Helonias thibetica in Taiwan was recently reported by Hsu et al. (2011, under the name of Ypsilandra thibetica ). The plant is, however, not H. thibetica , but appears to be a form of H. umbellata . For further notes on this form see the remarks under H. umbellata .

Figures

FIGURE 1. Buds and their sprouting in Helonias. A. H. bullata (material Bu-1; photo on 5 April 2013). B. H. orientalis (O-3; 15 November 2014). C, D. H. thibetica (T-1; C on 19 February 2014, D on 2 March 2014). Flower buds or fresh flowering stem designated by F, leaf buds or sprouting leaves by L, rosulate leaves developed in previous years by R. In Fig. C, two of leaves developed in the preceding year marked with single white circle (dot); two of leaves developed the year before last marked with paired white circles (dots). Photo A by Jim Fowler at site of “Bu-1” in South Carolina; B–D by the author at Hachioji, Tokyo.

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 13. Helonias thibetica (material T-1). A. Young, slightly open flowers with protruding stigma (photo: 13 February 2014). B. Young flowers with anthers not yet dehiscent (19 February 2014). C. Flowers with dehiscent anthers and tepals turning creamy yellow (23 February 2014). D. Ripe fruits (5 April 2014). E. Same infructescence as D with dehiscing capsules exposing fertile seeds set by artificial selfing (3 May 2014). F. Rosette mostly of fresh leaves with young bud in center (14 August 2013). All photos taken at Hachioji, Tokyo.

FIGURE 14. Helonias thibetica (material T-1). A. Flower. B. Proximal part of flower with nectar (n) secreted from adaxial bases of tepals. Two frontal tepals removed. C. Vertical section of flower. D. Proximal interior of partial flower. E. Two anthers showing adaxial side (left) and abaxial side (right), respectively. F. Vertical section of capsule with persistent floral parts. IF; inner filament. OF; outer filament. n; nectar. Scale at C also for D.