Helonias orientalis (Thunb.) Tanaka (1998a: 109)

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

11. Helonias orientalis (Thunb.) Tanaka (1998a: 109) ( Figs. 1B , 25 , 26 ).

≡ Scilla orientalis Thunberg (1794: 334) . ≡ Heloniopsis orientalis (Thunb.) Tanaka (September 1925: 2000). ≡ Sugerokia orientalis (Thunb.) Koidzumi (1930: 95) . Type:― JAPAN. E Japonia, Thunberg (Holotype: UPS-THUNB 8327*!). Note:―the type specimen was collected from Niphon [Honshû], Japan, according to the account of Scilla bifolia in Thunberg (1784: 138).

Japanese name:―Shôjôbakama (e.g. Itô 1829; Iinuma 1861, under Scilla japonica Thunb. ; Tanaka & Ono in Iinuma 1875, under Sugerokia japonica ).

Distribution:― Japan, Korea, E Russia (S Sakhalin) ( Fig. 29-O ).

Remarks:―The floral color of Helonias orientalis is usually pink or purplish, but sometimes deviates from it. The plants are expediently grouped into the following three forms depending on their flower color: (11a) the form with flowers other than white or yellow (i.e. the most common form with pink or purplish is included here), (11b) the form with white flowers, and (11c) the form with yellow or yellowish flowers. Because of sporadic occurrence in populations of the form with pink or purplish flowers (11a), the forms 11b and 11c are not treated here as independent taxa. Names formerly given to or corresponding to them are listed in synonymy under the respective forms (11a–c).

Identification key to the forms expediently grouped by their flower colors

1.

-.

2.

-. Tepals (at anthesis) pink or purplish pink or sometimes brick-red, blood-red, purple or bluish ................................................. 11a. Tepals (at anthesis) white or yellowish ............................................................................................................................................. 2 Tepals white ................................................................................................................................................................................. 11b. Tepals yellowish .......................................................................................................................................................................... 11c.

11a. The form with flowers other than white or yellow ( Figs. 1B , 25 excl. E, 26).

Heloniopsis pauciflora Gray (1859: 416) . Type:― JAPAN [N Hokkaidô]. Cape Romanzoff [northwestern extremity of Jesso―presently Cape Noshappu in Wakkanai-shi], 1853–1856, C. Wright s.n. (Holotype: GH-00029943*, p.p., excl. admixed fragments of a Luzula . vide Koidzumi (1930, 1934), Hara (1947). Isotype: US-00650433*!).

Sugerokia japonica Miquel (1866 , reprint: 24;1867a: 88―publication February 1867 at latest, for the publication date see Stafleu 1966: 84; July 1867 b: 145―publication date from Stafleu & Cowan 1981: 515); 1866–1867: 309; 1870: 92), p.p., excl. syn.? Scilla japonica Thunb. , Heloniopsis breviscapa Maxim. , and specimens from Kiusiu [Kyûshû]. ≡ Heloniopsis japonica Maximowicz (May 1867: 437). Type:― JAPAN (Japonia). Botanicus japonicus s.n. [no actual collector’s name nor detailed locality in Japan is recorded on the sheet]. The following remark is made in the protologue: A Mizutani Sugerok probabiliter in Japonia boreali detecta. (Holotype: L 908. 106-1567, barcode L-00325576*!). Note on the typification of this taxon:― Ohba et al. (2005) designated Siebold s.n. (L 908. 106-711; barcode L-0325582*!) as the lectotype and several other specimens collected by Siebold, Bürger, or Keisuke ITO as syntypes for this taxon. However, in the protologue only one collection was cited by Miquel. Accordingly, their type designations are invalid. For more details see the remarks below.

Sugerokia japonica Miq. var. racemosa Miquel (July 1867 b: 145; 1866–1867: 309). Type:― JAPAN. Botanicus japonicus s.n. . (“Legit bot. iaponicus, probabliter Sugerok [Mizutani]” in the protologue). (Holotype: L-0325576*!). This specimen is regarded as the holotype for S. japonica Miq. Sugerokia japonica var. racemosa is therefore the homotypic synonym of S. japonica . For more details see the remarks below.

Heloniopsis pauciflora A.Gray var. rubra Koidzumi (1921: 15) , cum descr. jap. Type (possible type collection needing further studies):― JAPAN. Hokkaidô. Kamifurano, 1916, H. Koidzumi (specimen collected by him in the same locality on 14 July 1917 is in TNS!). Nagano Pref.: Mt. Norikura, July 1920, H. Koidzumi (specimen collected on 10 July 1920 is in TNS!); Mt. Shirouma, August 1920, H. Koidzumi (specimen dated 20 July 1920 is in TNS!). Japanese name: Beni-shôjôbakama ( Koidzumi 1921).

Heloniopsis japonica (Thunb.) Maxim. var. purpurea Nakai (1933: 243) p.p.? ≡ Heloniopsis orientalis (Thunb.) Koidz. var. purpurea (Nakai) Nakai (1952: 146) . ≡ Hexonix japonica (Thunb.) Raf. var. purpurea (Nakai) Wang & Tang (1949: 113) . Type:―Not designated. ‘Habitatio’ and ‘nomen japonicum’ given under this name by Nakai may not be for this taxon, but those for Heloniopsis japonica as a species, probably due to the erroneous insertion of the three lines of var. purpurea into the account of the latter. For further notes see the remarks below.

Heloniopsis japonica (Thunb.) Maxim. var. sanguinea Nakai (1933 ; 243). ≡ Hexonix japonica (Thunb.) Raf. var. sanguinea (Nakai) Wang & Tang (1949: 113) . Type:― JAPAN. Hondo [Honshû]. No specimen designated. In TI, however, there is a specimen annotated with this name by Nakai, which is from the Konsei(tôge) Pass, Nikkô, 30 June 1923, fr., Y. Yamamoto s.n. Japanese name: Benibanashôjôbakama ( Nakai 1933).

Heloniopsis japonica Maxim. f. lateritia Nakai ex Honda (1938: 1679) . Type:― JAPAN. Honshû. Prov. Tôtômi [Shizuoka Pref.], Ogasagun, Mt. Ogasa, 3 April 1931, G. Hashimoto s.n. (Holotype: TI! Isotype: TNS-44182!). Japanese name:―Akabana-shôjôbakama ( Honda 1938).

– Scilla bifolia auct. non L.: Thunberg (1784: 138).

– Heloniopsis acutifolia auct. non Hayata: Honda (1938: 1678).

– Sugerokia acutifolia auct. non (Hayata) Koidz.: Honda (1938: 1678).

– Heloniopsis breviscapa auct. non Maxim.: Makino (1891: 126); Matsumura (1905: 197), p.p.

– Heloniopsis orientalis (Thunb.) Tanaka var. breviscapa auct. non (Maxim.) Ohwi: Ohba et al. (2005: 127).

Description:―Rhizome subcylindrical, to 6.5 cm long, to 2.5 cm in diam., somewhat knobby, subannulate with many close scars, whitish or pale brownish. Roots filiform, to 2.5 mm in diam., white, contractile. Leaves persistent for 1–2 years, oblanceolate, spatulate or narrowly oblong, cuneate to petiole, 4–29 cm long, 1–4.5 cm wide, petiolar part to 1 cm wide, margin entire, apex short acuminate, apiculate, apiculus ca. 0.5 mm long, sometimes with adventitious plantlet (ramet) at apex, abaxial midvein raised, adaxially glossy, chartaceous (somewhat thick in texture). Flowering stem to ca. 17 cm long at anthesis, elongating in fruit; peduncle terete, 3.5–16 cm long at anthesis, to ca. 60 cm or more in fruit; base densely covered with acute or acuminate, ovate scale-like leaves; scale-like leaves on peduncle 7–12 (excl. basal ones), narrowly oblong(-elliptic), narrowly lanceolate or ovate, acute, to ca. 2.7 cm long; inflorescence subumbellate, sometimes umbellate, becoming racemose in fruit, rachis to 1 cm long at anthesis, 1.5–9.5 cm long in fruit; pedicels 6- ribbed, 0.4–2.8 cm long (incl. those in fruit), purplish pink. Flowers 2–10(–13), funnelform, 1.2–2.5 cm across, usually slightly nodding, usually odorless, rarely slightly scented. Tepals 6, obliquely expanded, sometimes slightly recurved distally, usually pink or purplish pink (4–)5–7-veined, narrowly oblong-oblanceolate (or spatulate), 9.5–20 mm long, 2.4–5 mm wide, base affixed to receptacle (apex of pedicel) for ca. 1.2–1.5 mm in length, abaxially shortly gibbous, apex obtuse, adaxially canaliculate toward base, usually slightly revolute proximally at margin, scarcely imbricate with adjoining tepals, basal submarginal portions winged (lamellate) inward; wings extending over proximal ca. 1/4–1/3 of tepal, connate proximally to those of adjoining tepals for 0.4–0.5 mm in height, also adnate to opposite filament for (0.8–) 1.3–2 mm in height, forming a sheathing nectary. Stamens 6, usually exceeding tepals; filaments obliquely expanded, purplish pink, filiform, 8–18 mm long, proximally scarcely dilated and complanate; anthers affixed to filament at adaxial portion ca. 0.2 mm above base of connective (i.e. [sub]dorsi-basifixed), extrorse, dark purple, narrowly ovate, oblong, or narrowly elliptic-oblong, 2.5–5.1 mm long, sagittate at base, rounded at apex, bilocular with vestigial apical confluence between thecae; pollen whitish or pale purplish. Pistil 1, usually exceeding stamens and tepals; ovary sessile, globose, obtusely trigonous, trisulcate, darkly purplish pink, 2–4 mm long, 2–3.8 mm across, apex emarginate-depressed in center; style terete, straight, or slightly up-curved distally in nodding flowers, 8–24 mm long, usually purplish pink; ovules numerous, multiseriate on central axile placentae (ca. 8–12-seriate in each ovarian locule); stigma discoid, often obtusely trigonous, sometimes trilobed or tricleft, slightly concave in center, 1–2.7 mm across, (dark) purplish pink. Capsules obpyramidal, obcordate or obdeltoid in lateral aspect, triparted, 5–7 mm long, 9–12 mm across, apex depressed in center; lobes subpyriform, ascending to horizontal. Seeds numerous, coated with whitish testa, linear-fusiform, sublinear proximally, subulate distally, usually falcate, 3.5–6.6 mm long, 0.25–0.5 mm wide; body of seed narrowly oblong-fusiform, 1.1–1.6 mm long, 0.2–0.3 mm wide, (dark) brown.

Additional specimens examined (see also Tanaka 1998a):― JAPAN. Hokkaidô. Hidaka: Mt. Poroshiridake, 1600 m, 25 July 1971, fl., H. Koyama et al. 3808 (TNS-303241). Iwamizawa: Horomui, 29 June 1927, fr., K. Kondo 1630 (BM-001118050). Kamikawa: Asahikawa-shi, Kamuikotan, 27 April 1894, fl., S. Ohwaki (TNS-547312). Kamikawa-gun: Mt. Taisetsu, 18 July 1941, fl., S. Okuyama (TNS-69228). Oshima: Matsumae-gun, Mt. Daisengendake, 8 May 1907, fl., Kudo 20 (MAK-137595). Samani-gun: Hidaka, Mt. Apoi, 26 July 1941, fr., S. Okuyama (TNS-69229). Sorachi-gun: Mt. Yûbaridake, 27 July 1933, fr., H. Iwamoto (TNS-43155). Honshû. Chûbu Distr. Fukui Pref.: Takefu, Murakuniyama, 150 m, 26 March 1987, fl., T. Yamazaki (TI). Gifu Pref.: Kawai, 22 April 1960, fl., S. Okuyama 14149 (TNS-262517); Ohno-gun, Takane-mura, Dohgo-gawa, 1200 m, 7 May 1973, H. Kanai 731154 (BM-1118051). Nagano Pref.: Shimoina-gun, Etsutaro, Urugi-mura, 800 m, 16 September 2000, K. Asano & H. Asano 30185 (MAK-317546); Mt. Asama, 31 May 1937, fl., H. Ito (TI); Kita-azumi-gun, Mt. Shiroumadake, 27 August 1902, fl., Y. Yabe (TI); Kiso, Mt. Komagatake, 27 July 1925, fl., H. Koidzumi (TNS-315728). Niigata Pref.: Mt. Tairappyô, 21 June 1959, fl., H. Hara (TI); Nishikanbara-gun, Mt. Yahiko, 29 March 1987, fl., K. Inoue 4229 (TI); Isl. Sado, Nyûkawa- Tadaramine, 13 May 1954, fl., S. Okuyama 10886 (TNS-135230). Shizuoka Pref.: Abe-gun, Mt. Higashidake, 23 July 1924, fl. (TNS-315745); Hamana-gun, Kosai-chô, Bôse, April 1974, H. Kanai & H. Koyama (TNS-318330). Toyama Pref.: Yatsuo-chô, 27 May 1955, fl., A. Kirino 570 (TNS-122487). Chûgoku Distr. Okayama Pref.: Mt. Hirusen, 600 m, 4 May 1959, fl., G. Murata (TNS-186826). Tottori Pref.: Mt. Daisen, 6 June 1925, fr., Y. Nishikado (TNS-460543). Yamaguchi Pref.: Mt. Nameriyama, 14 October 1949, T. Nakai & N. Maruyama (TNS-83141). Kantô Distr. Chiba Pref.: (Sanmu-shi) Haniya, 14 April 1935, fl., T. Makino s.n. (MAK-137632). Gunma Pref.: Mt. Tanigawa, July 1956, fl., K. Okamoto (TNS-389529); Mt. Shibutsu, 20 June 1933, fl., D. Hoshi (TNS-315723). Tochigi Pref.: Mt. Takahara, 1400–1500 m, 25 May 1959, fl., H. Kanai (TNS-132582); Nikkô, Konsei-tôge, August 1883, fr., H. Sakurai (TNS-3844). Tokyo Pref.: Isl. Miyake, Mt. Oyama, 5 May 1938, fl., Hayashi (MAK-137631); Isl. Mikura, 9 August 1952, H. Tsuyama (TNS-102625); Isl. Kôzushima, 27 April 1887, fl., K. Shirai (TNS-54364). Kinki Distr. Hyôgo Pref.: Isl. Awajishima, Mt. Yuzuruhayama, 5 May 1958, fr., N. Satomi (TI). Kyoto Pref.: Takeno-gun, Yasaka-chô, 300 m, 25 April 1970, fl., G. Murata 20968 (TNS-30409); Ôi-yama, 500 m, 28 April 1968, fl., M. Togashi (TNS-210893). Mie Pref.: Mt. Fujiwaradake, 5 April 1953, fl., S. Okuyama (TNS-102335, -102338). Ôsaka Pref.: Shinjyô-Nawate, 23 April 1933, fl., Z. Yoshino (TI). Shiga Pref.: Kôka-gun, Mt. Handôjisan, 22 April 1962, fl., H. Koyama 1196 (TNS-245904). Tôhoku Distr. Akita Pref.: Akita-shi, Mt. Tegata, 2 June 1929, fr., H. Muramatsu (TI); Senpoku-gun, Mt. Komagatake, 1300 m, 5 July 1975, Y. Tateishi 2123 (TI). Aomori Pref.: Mt. Iwaki, June 1941, fl., D. Narita (TNS 132383); Mt. Hakkôdasan, 30 June 1964, fl., K. Hasegawa (TI). Fukushima Pref.: Aizu, Mt. Tashiro, 8 July 1959, fr., S. Okuyama 16611 (TNS-277605). Iwate Pref.: Hiraizumi-chô, Chûsonji Temple, 12 April 1932, fl., K. Nemoto (TNS-36033). Miyagi Pref.: Sendai, Dainohara, 5 April 1914, fl., Y. Ogura (TI); Shichikashuku-machi, Watase-fûketsu, 16 April 1978, fl., R. Suzuki (TNS-374850). Yamagata Pref. Tsuruoka-shi, Mt. Yudonosan, 22 July 1887, R. Yatabe (TI); Mt. Chôkaisan, 28 July 1887, fl., R. Yatabe (TI). Shikoku. Ehime Pref.: Between Tsutsujôsan and Mt. Ishizuchiyama, 28 May 1956, fl., G. Murata & T. Shimizu (MAK-13149). Kôchi Pref. : Takaoka-gun, Sakawa-chô, June 1893, fr., T. Makino (MAK-137668). Tokushima Pref.: Mt. Kôtsusan, 18 April 1948, fl., K. Abe 12167 (TNS-132382).

Distribution:― Japan (Hokkaido, Honshû, Shikoku, N Kyûshû*), E Russia (S Sakhalin ―e.g. Miyabe & Kudo 1932, Sugawara 1937) ( Fig. 29-O ). * The occurrence of Helonias orientalis in northern Kyûshû is recorded in Amakawa (1975), but the specimen has not yet been examined by the present author.

Habitat:―Shady moist banks along streams in forests, or sunny open mountain meadows or marshes, at elevations ca. 10–3000 m.

Conservation status:―This form is widespread, and assessed as LC according to the criteria set out in IUCN (2001). However, local populations near human habitation may be more or less subject to disturbance and threats.

Flowering:―Usually March–May. Delayed at high elevations and/or in northern regions.

Ripening:―Usually May. Delayed at high elevations and/or in northern regions.

Remarks:―On the nomenclature of two species of Helonias occurring in Japan, Hara (1947) made the following comment: “When Miquel (1866 in reprint, 1867a) first described Sugerokia japonica Miq. , he did not cite Scilla japonica Thunb. ( Thunberg 1784) as its basionym. It was in 1870 that he first cited Scilla japonica as the synonym for his species of Sugerokia . So the Miquel’s species at the time of publication is regarded as not based on the Thunberg’s species. On the other hand, Heloniopsis japonica published by Maximowicz (1867) is the combination not based on Thunberg’s Scilla japonica but on Miquel’s Sugerokia japonica ”. Judging from the protologue by Miquel (1866, 1867a), the type of Sugerokia japonica appears to be equivalent with Helonias orientalis circumscribed in the present paper. Miquel’s Sugerokia japonica is based on the specimen regarded by him as probably collected in northern Japan by [MIZUTANI] Sugerok [Sukeroku] ( Miquel 1866). As no other collections were cited in the protologue, the specimen should be the holotype. On the other hand, Miquel (1867b) published S. japonica var. racemosa on a material regarded by him as probably collected by Sugerok. According to Yamaguchi et al. (2003: 295, as Heloniopsis ), there is one specimen of Helonias orientalis collected by an anonymous Japanese (L 908. 106-1567, barcode L-0325576*!). This specimen is evidently the holotype for var. racemosa . Yamaguchi et al. (2003) also indicates that there is no specimen of Helonias (Heloniopsis) at Leiden actually accompanying the name Sugerok as the collector. Accordingly it seems possible to view the specimen collected by an anonymous Japanese botanist (L-0325576) as the type of S. japonica . Judging from the description of S. japonica in the protologue ( Miquel 1866), the specimen is at a fruiting stage. At this stage, H. orientalis usually has an elongate, racemose infructescence. This character is also consistent with the brief description of var. racemosa by Miquel (1867b). The name S. japonica var. racemosa is therefore the homotypic synonym for S. japonica . In Japan Helonias orientalis occurs mainly in snowy regions facing the Sea of Japan. Glossy, somewhat thick or firm leaves of this species seem closely related to a heavy snowfall in the habitats. Supposedly, their leaves have become durable to the heavy snowfall in the evolutionary process. The production of plantlets (ramets) by the leaves (Part I-4) might also be related to the heavy snowfall. Much snow accumulated during winter is likely to keep the leaves strongly appressed to the ground. This condition might have become a factor for the induction of such a plantlet production by leaves. If the habitats are kept moist during the growing season, the establishment of the plantlets may be favored. In contrast, in plants of H. breviscapa occurring in warmer, less snowy regions in Japan, the leaves are thinner, and adventitious plantlet formation by leaves is not seen. The authentic specimens for Heloniopsis japonica var. purpurea Nakai (1933) and H. orientalis var. purprea (Nakai) Nakai (1952) (see the nomenclatural list above) are not known at present. Nakai (1952) applied the (latter) name to the Korean plants of Helonias , so Helonias koreana with purple flowers may represent the taxon. However, plants of Helonias orientalis with purple flowers also occur in northern Japan (Tôhoku and Hokkaidô), it is therefore possible that both Korean and Japnese plants were included in his original concept of the taxon. In Helonias orientalis , floral colors other than pink or purplish pink, such as brick-red, blood-red, blue, and blackish purple-blue, have also been known. Flowers with blue tepals of this species are often found in northern cool regions like Hokkaidô ( Japan). Pink tepals often turn dirty brick-red after anthesis. Several variants in floral color were named previously as listed in the synonymy.

11b. The form with white flowers ( Fig. 25E ).

Heloniopsis grandiflora Franchet & Savatier (1877: 88 , nom. nud.; 1878: 529), ‘Helionopsis’. ≡ Heloniopsis japonica (Thunb.) Maxim. var. grandiflora Nakai (1933: 243) , p.p. Japanese name: Shirobana-shôjôbakama ( Nakai 1933). ≡ Hexonix breviscapa (Maxim.) F.T.Wang & Tang var. grandiflora (Franch. & Sav.) Wang & Tang (1949: 113) . Type (lectotype, designated here):― JAPAN. Honshû, in provincia Etchigo, circa Niigata, Vidal (Savatier 2726) (P-00730554*!). Isotype: P-00730555*!, P-00730556*!).

Japanese name:―Yuki-shôjôbakama (nov.; meaning snowy white Helonias ); Shiro-shôjôbakama ( Hiyama 1939).

Description:―Tepals, filaments, styles and stigmas white. Anthers cream or purple. Similar in other character states to the form with pink or purplish flowers (11a).

Additional specimens examined:― JAPAN. Honshû. Shiga Pref. (Ômi): Hikone-shi, Surihari-tôge , 11 April 1902, T. Makino 960 (MAK-137679). Tôkyô Pref.: Izu , Isl. Kôzu-shima , Mt. Tenjô , in herbages, elev. 400 m, 14 April 1976, flowers white, only one plant has white flowers, others purplish pink (in Japanese), T. Yamazaki s.n. (TI). Yamagata Pref.: Mt. Zaô , 20 August 1928, fr., flowers are white, C. Sto 202 (TI). Shikoku. Kôchi Pref. : Hata-gun, Ôgata-mura, Komehara, 16 March 2003, fl., flowers white, M. Tashiro et al. (MBK-131864*). The specimens listed below were recorded by collectors as having yellowish flowers, but there is a good possibility that their flowers were originally white and turned yellowish with the progression of anthesis, though the truth is unknown presently. The floral colors recorded on the herbarium sheets are varied from pale yellowish white to pale greenish yellow : JAPAN. Honshû. Prov. Iwashiro [Fukushima Pref.]: Fujita-tôge, 30 April 1929, flowers pale greenish yellow, S. Saito s.n. (TI). Nagano Pref.: Mt. Asama , Sekison-san, 9 April 1957, flowers pale yellowish white, K. Kawamoto (TNS-504635). Uzen [Yamagata Pref.]: Yonezawa, 2 May 1931, flowers pale yellow, H. Koidzumi (TNS-315689) .

Distribution:― Japan (Honshû, Shikoku). Sporadically occurs.

Flowering:―Usually March–April.

Remarks:―Franchet & Savatier (1878 in 1877–1878) described the floral color of Heloniopsis grandiflora as white. This taxon is similar in other respects to Helonias orientalis . Hiyama (1939) quoted Heloniopsis orientalis f. albidula H.Koidz. as a name for the white-flowered form. But unfortunately I have not been successful in tracing the original article with this name by Hideo Koidzumi.

11c. The form with yellow flowers.

Heloniopsis orientalis (Thunb.) Tanaka f. lutea Mochizuki (June 1978 a: 14; November 1978 b: 10). Type:― JAPAN. Honshû. Akita Pref., Nishi-senboku-machi, 24 May 1977, R. Mochizuki (Holotype: KANA-87335, n.v.).

Japanese name:―Kibana-shôjôbakama ( Mochizuki 1978a, b).

Description:―This form has yellow tepals, and is similar in other respects to the form with pink or purplish pink flowers (11a).

Specimen examined:― JAPAN. Honshû. Nagano Pref.: Sarashina-gun, Shinonoi-cho, Shinbashi-yama , 30 April 1956, flowers clear yellow, M. Minemura 681 (MAK-6543) .

Distribution:― Japan: Central and N Honshû (Chûbu and Tôhoku Districts).

Remarks:― Mochizuki (1978a, b) found an individual with yellow flowers among many individuals with purplish flowers. Minemura also collected a yellow-flowered form in Nagano Pref., central Honshû, Japan (MAK-6543!). She has left a record on the label that the flowers are clear yellow from a bud stage. One needs to be careful in recognizing a genuinely yellow-flowered form of Helonias orientalis , since white flowers of the same species consistently turn creamy yellow soon after the peak anthesis. In this connection, a few specimens recorded as having yellowish flowers are cited under the form with white flowers (11b).

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 25. Helonias orientalis (A–D, F; material O-2. G; O-1. H; O-3) and white flowered form (E; O-w-1). A. Young slightly open flowers with protruding pistil (photo: 25 March 2014). B, C, D. Sub-umbellate flowers (B on 31 March 2014; C on 29 March 2014; D on 5 April 2012). E. Compact raceme of white flowers (30 March 2012). F. Infructescence (29 April 2014). G. Dehisced capsules (14 May 2014). H. Rosette with flower bud (arrow) in center (14 August 2014). All photos at Hachioji, Tokyo.

FIGURE 26. Helonias orientalis. A–C; material O-3. D, E; O-2. A. Proximal interior of partial flower. Connate portion between adjoining tepals arrowed.Adnate portions between tepals and opposing filaments marked with asterisks (*). B. Proximal portion of flower in vertical section. Surface of nectary arrowed. IF; inner filament. C. Proximal part of flower in cross section. D. Slightly dehisced capsule with persistent floral parts. E. Three valvate, dehisced capsule. Scale at A also for C; at D also for 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.