植物研究雑誌 J. J. Jpn. Bo t. 73: 73: 102-115 (1 998)

Phylogenetic Phylogenetic and Taxonomic Studies onHelonias ，Ypsilandra andHeloniopsis 111. 111. Taxonomic Revision

Noriyuki Noriyuki TANAKA

Department Department of Education ，School of Liberal Arts ，Teikyo University 359 359 Otsuka ，Hachioji-shi ，Tokyo ，192-0395 JAPAN (Received (Received on April 2 ，1997)

The three genera ，Helonias ，浄silandra and Heloniopsis have not sufficient differ- ences ences to be distinguished from each other in generic rank. Accordingly the three genera were were unified into one genus. Heloniopsis and Ypsilandra were reduced to Helonias. The following following taxonomic reductions were also made in this paper; i.e. ， both Y. cav αleriei and Y. Y. kansuensis were reduced to Helonias thibetic α; both Heloniopsis arisanensis and eloniopsis H eloniopsis taiwanensis to Helonias umbellata; Y. yunnanensis var. himalaica to Helonias yunnanensis; yunnanensis; Sugerokia nipponica ，Heloniopsis japonica v紅 .flavida ， H. japonica v紅. albiflora albiflora and H. japonica var. tessellata to Helonias breviscapa. Fl oral colour variants were were not distinguished as distinct taxa. As a result ， a total of nine species were recognized in in the emended Heloni ω. Eight new names were produced as a consequence of the transference transference of both ゆsilandra and Heloniopsis to Helonias. (Continued (Continued from 1. Jpn. Bo t. 72: 329-336 ， 1997)

According to my previous studies (Tanaka H. kawanoi ， the basal part of the inner fila- 1997a ，1997c-e) ，Helonias and Ypsilandra ments is located very close to that of the ovary ， differ differ mainly in the following characters; i.e. ， and this state does not differ so markedly from themo ゅhology of seeds ， the number of ovules that in Ypsilandra in which the two floral per per loculus of an ov 紅 y， the number of flowers organs are adnate to each other only very per per scape ，and the presence or absence of a shortly at their bases. Accordingly ， the differ- longitudinal longitudinal furrow between thecae on the ence conceming the adnation of these two ventral ventral side of anthers. As for the third charac- floral organs between the two genera seems ter ， there is some ove r1 ap in the range of not so great virtually and may not be worthy of variation variation between the two genera (Tanaka generic delimitation. Meanwhile ， both the in- 1997d). 1997d). All these differences seem not to be ner and outer filaments of Ypsilandra are not sufficient sufficient for the generic demarcation of these adnate to ， while those of the two species . plants. of Heloniopsis (H. orientalis and H. Meanwhile ， as for Ypsilandra and bre ν iscapa) are adnate basally to their co 町 e- Heloniopsis ， the following differences are spondingtepals(Tanaka 1997b ，1997c ，1997e). known to exist between the two genera. That Franchet (1887) earlier noted that there is this is ， in 浄silandra the inner filaments are adnate difference between the two genera. However ， basally basally to the ovary ， while inHeloniopsis they it was later disclosed that there are also three are are free from the ovary (Tanaka 1997 c，1997 e). other species in Heloniopsis whose filaments However ， in some species of Heloniopsis like are not adnate to tepals (Tanaka 1997b ，1997 c，

一一 102 一一 April1998 Journal of Japanese Botany Vo l. 73 No. 2 103

1997e). 1997e). Therefore ， the presence or absence of A11 the differences so far known to exist the the adnation between these two flora1 organs between the three genera seem not to deserve (f i1 aments and tepa1s) cannot be regarded as an a generic de1imitation. They possess many iritergeneric iritergeneric difference in this case. As regards characters in common and c1 ose1y resemb1e the the anthers ，均 silandra has perfectly un i1 ocu- each other in basic habits. It seems more ap- 1ar 1ar anthers ， whi1e Heloniopsis has virtua11y propriate to treat them in one genus ， as Kawano bi10cu1ar bi10cu1ar anthers. It is a noteworthy that fact (in Kawano and Masuda 1980) ear1ier sug-

Heloniopsis Heloniopsis sti11 retains 位aces of a theca1 co 任 gested. In this paper 1 attempted to revise the fluence fluence at the apex of the anthers. Especia11y in of this p1ant group ，in c1 uding the some species 1ike H. leucantha ，the 住aces unification of the three genera into one genus. appear appear to be re1ative1y we11 retained. As far as the the anthers ofthese species of Heloniopsis 紅 e Genus Helonias 3 compar 吋 with those of Ypsilandra ， the gap Helonias L. )， Sp. P1.， ed. 1 ，1: 342 (1 753) ， between the two genera seems not so deep ed. 2 ，1: 485 (1762) ，p.p. ，exc l. P1ukenet; Gen. substantia11y substantia11y (Tanaka 1997c). Pl.， ed. 5 ，1: 159 (1 754) ， ed. 6 ，1: 180 (1 764); Gray (1859) once pointed out that there is a Amoen. Acad. 3: 12 ，t. 1 ，f.1 (1756); Mu 町 ay ， difference difference in the sty1ar structure between Sys t. Vege t.， ed. 14 ，349 (1784); Wi11d. ， Sp. Pl. Helonias Helonias and Heloniopsis. But ，1ater it was 2: (1): 273 (1 799) ，p.p.; Redoute ，Li1iac.l ，t. 13 proven that the sty1ar structure varies (1 802) ，exc l. P1ukenet; Michx. ，Fl. Bo r. -Amer. transitiona11y transitiona11y from that of Helonias through 1: 211 (1803) ，p.p.; Curtis' s Bo t. Mag. 20: 吻silandra to that of Heloniopsis (Tanaka t. 747 (1 804); Pers. ， Syn. P1. 1: 398 (1 805) ， 1997a ，1997c ， 1997e). Therefore ，what Gray p.p.; Aiton ， Hort. Kew. ， ed. 2， 2: 330 (1 811) ， observed observed was the two ex 佐emes of an a1most p.p.; Spreng. ，Sys t. Veg. ，ed.16 ，2: 144 (1 825) ， continuous continuous variation in this structure. Gray p.p.; Schu1 t. & Schu1 t.， f. Sys t. Veg. 7: 1561 (1859) (1859) a1so pointed out that Helonias and (1830) ，p.p.; End l.， Gen. Pl.， p.135 ， no. 1066 Heloniopsis Heloniopsis differ in seed character ， but 1ater (1 836-1840) ，p.p.; Kunth ，Enum. Pl. 4: 174 it it tumed out that the seeds he examined as (1843) ，exc l.“ H. ? japonica"; Gray ，Man. those those of Heloniopsis were ， in fact ， not those of Bo t.， ed. rev. ，478 (1 859); Baker in J. Linn. Heloniopsis ， but those of a species of Luzula Soc. 17: 467 (1 879); Benth. ， Gen.. P1. 3 (2): (Koidzumi 1930 ，1934 ，Hara 1947). There- 827 (1 883); Eng l.， Na t. Pfl.-fam. 11.5: 22 fore ，Gray' s description of the seeds of (1888); Britton & A. B r.， I11. Fl. N. States & Heloniopsis Heloniopsis must have been nu11ified. With- Canada ， ed. 2. 1: 488 (1 913); Kr ause in Eng l. out out knowing this fact ，Mique1 (1867) 1) set up a & Prantl ，N a t. Pfl. -fam. ，ed. 2， 15a: 258 (1930); new genus Sugerokia ， because his materia1 Hutch. ，Fam. F10w. Pl. 2: 85 (1934); Fema1d ， differed differed especia11y in the seed character from Gray's Man. Bo t.， ed. 5，424 (1950); G1eason ， Gray's Heloniopsis. Both Sugerokia and I11. Fl. NE. U.S. 1: 406 (1952). 4 Heloniopsis Heloniopsis were based on specimens from Abalum Adans. )，Fam. Pl. 2: 47 ，511 Japan ，and it is evident that they are the same “OAbalon" (1763) ，p.p. ，exc l. P1ukene t. genus ， as many taxonomists have ea r1 ier Hexonix Rafin. (nom. rejic. vs. Heloniopsis pointed pointed out (Maximowicz 1867 ，Baker 1874 ， A. Gray 1859) ， F l. Te11ur. 2: 13 (1 837)5). 2 1879 ， Franchet and Savatier 1877 )，Bentham Kozola Rafin. (nom. rejic. vs. Heloniopsis 1883 ， Eng1er 1888 ，Koidzumi 1930 ，Kr ause A. Gray 1859) ， F l. Te11ur. 2: 25 (1837)5). 1930 ，Nakai 1933 ，Honda 1938 ，Hara 1947 ， Heloniopsis A. Gray (nom. cons.) in Mem. Ohwi 1953 ，1965 ，Chen 1980). Amer. Acad. ser. 2 ，6: 416(1859) ，exc l. charac t. 104 104 植物研究雑誌第73 巻第2号 平成10 年4月 seminum (vide H. Hara 1947 ， infra ci t.); about as long as or exceeding tepals; anthers Maxim. in Bull. Acad. Sci. S t. -Petersb. 11: ovate ， oblong or hippocrepiform ，extrorse to 436 (1 867); Miq. in Ann. Mus. Bo t. Lugd.- latrorse ，unilocular or virtually bilocular; fila- Batav. Batav. 3: 146 (1 867) ，“ Helionopsis"; Baker in ments filiform ，persistent ， outer filaments free J. J. Bo t. 12: 278 (1 874) ，“ Heleniopsis"; in J. from ov 訂 y， while inner ones free from or

Linn. Linn. Soc. 17: 459 (1 879); Franch. & Sav. ヲ adnate basally to ovary. Both whorls of fila-

Enum. Pl. Jap. 2: 87 (1 877)2) ，529 (1878)2) ラ ments adnate basally to or free from their Helionopsis"; “Helionopsis"; Benth. ，Gen. Pl. 3 (2): 827 corresponding tepals. Gynoecium 1， (1 883); Eng l.， Na t. Pfl.-fam. 11.5: 22 (1 888); tricarpellate. Style either simple with a capi- Krause in Eng l. & Prantl ，Na t. Pfl.-fam. ， ed. 2， tate to depressed capitate stigma ， or divided to 15a: 15a: 259 (1 930); Hutch. ，Fam. Flow. Pl. 2: 86 various degrees in 3， of which ventral side is (1 934); H. Hara in J. Jpn. Bo t. 21: 148 (1 947); stigmatic. Ovary superior ，globose ，ellipsoid Ohwi ， Fl. Jap. ，ed. rev. ，336 (1 975); S.C. Chen ， or obovate ，trilobed to trigonous ，stipitate or l. Fl. Reip. Pop. Sin. 14: 15 (1 980); Satake in estipitate ，trilocular ， with ovules on the central Satake Satake et al.， Wild Flow. Jap. 1: 26 (1 981). marginal placenta. Capsules tr i1 obed ， Sugerokia Sugerokia Miq. in Ann. Mus. Bo t. Lugd.- loculicidally dehiscen t. Seeds coated with a Batav. Batav. 3: 144 (1867) 1); Koidz. ， F l. Symb. Or.- whitish to brownish soft testa ， fusiform to Asia t. 94 (1 930). linear ，tailed at both ends. Flowering generally Ypsilandra Ypsilandra Franch. in Nouv. Arch. Mus. in spring ， except H. kαwanoi which flowers in His t. Na t. Paris ser. 2， 10: 93 ，t. 17 (1887); Pl. summer to autumn. One species (H. bullata) David.2: David.2: 132 ，t. 17 (1 888); Kr ause in Eng l. & distributed in eastern N. America ， while the Prantl ，Natur l. Pfl.-fam. ，ed. 2， 15a: 260 (1 930); other eight species in Asia. Chromosome Hutch. ，Fam. Flow. Pl. 2: 85 (1 934); S.C. number 2n = 34. Chen ， F l. Reip. Pop. Sin. 14: 15 (1 980); G.H. Type species: Helonias bullata L. Yang ， F l. . 7: 11 (1 991); Noltie ， F l. 3(1): 90 (1 994). Key to the species

Perennial Perennial glabrous herb. stout ， 1. Style trilobed or trisected ..，・ H ・H ・H ・.....・ H ・.2 conic a1 to tuberous ，bearing many fibrous roots. 1. Style not distinctly lobeq ， with a capitate to

Leaves many in a basal rosette ，evergreen ， depressed capitate stigma .…....・ H ・.....・ H ・..4 oblanceolate ， narrowly oblanceolate or spatu- 2. Style shortly trilobed ， with the columnar late-oblanceolate ， acute to acuminate at apex part being c.6 mm long. Flowers 2-3 per with with a short pointed tip ，attenuate to base. scape. Tepals linear to linear-oblanceolate Scape erect or ascending ，simple ，terete ， with … 3) H. alpina several several short scaly leaves ，1-45 cm in height at 2. Style (廿 ifid 一) triparted to trisected ， with the

anthesis. anthesis. racemose to umbel 由 undivided (columnar) p紅 t being less than late ， bearing many to few flowers ， usually c.l.5 mm long. Flowers usually more than ebracteate ebracteate (exceptH. kawano i). Flowers pedi- c.5 per scape. Tepals elliptic to oblanceo-

cellate ， cup-shaped or campanulate ， with a late ...・ H ・.....・ H ・.....・ H ・H ・H ・..…...・ H ・..…...・ H ・... 3 strong to weak fragrance. Tepals 6 ， 3. Undivided columnar part of style to c.l.5 oblanceolate ，elliptic ， narrowly elliptic ，nar- mm long. Flowers c.5-15 per scape. Seeds 6 rowly oblong or linear-oblanceolate ， equal in c.0.5-0.7 mm broad ) .. 2) H. yunnanensis length ， 4-16.5 mm long ，nectariferous on the 3. Undivided part of style to c.0.5 mm long. ventral ventral side near base ，persisten t. Neighbour- Flowers c.30-72 per scape. Seeds c.O.8-1.0

ing ing tepals connate or free at base. 6， mm broad ....・ H ・...・ H ・..…...・ H ・.. 1) H. bullata April1998 April1998 Journal of Japanese Botany Vo I. 73 No. 2 105

4. 4. Anthers unilocular. Inner filaments adnate 342 (1 753) ， ed. 2， 1: 485 (1762) ，p.p. ，exc 1.

basally basally to ovary ....・ H ・....・ H ・. 4) H. thibetica Plukenet; Amoen. Acad. 3: 12 ，t .1， f.1， (1 756); 4. 4. Anthers virtually bilocular (thecal conflu- Willd. ， Sp. P1. 2(1): 273 (1 799) ，p.p. ，exc 1. ence ence at apex rudimentary). Inner filaments Plukenet; Redoute ，Liliac. 1: 13 (1 802) ，p.p. ，

free free from ovary ....・ H ・-…....・ H ・-…....・ H ・-…… 5 exc 1. Plukenet; Aiton ，Hort. ，Kew. ed. 2 ，2: 330 5. 5. Filaments not adnate to their corresponding (1 811); Kunth ，Enum. P1. 4: 174 (1 843); Baker tepals. tepals. Neighbouring tepals free from each in J. Linn. Soc. 17: 468 (1 879); Britton & A.

other...... ・H ・.....・ H ・..…...・ H ・..…...・ H ・.....・ H ・.... 6 B r.， 11 1. F1. N. States & Canada ， ed. 2， 1: 488 5. 5. Filaments adnate basally to their correspond- (1 913); Gleason ，I11.Fl .NE. U.S.l: 406 (1 952); ing ing tepals. Neighbouring tepals distinctly Radford et a1.， Man. Vasc. F1. Carolinas ，299

connate connate at base …...・ H ・..…...・ H ・.....・ H ・..…… 8 (1 964); Rickett ，Wild Flow. U. S. 1(1); 40 ， 6. 6. Fl owering in late summer to autumn. Tepals p1. 8 (1 965); Utech in Ann. Carnegie Mus. 47: usually usually less than 6 mm long …- 169 (1 978).

…7)H. kα， wanoi Veratrum (Americanum) rαcemo 6. 6. Flowering in spring. Tepals usually more simplicissimo ，cae t. Miller ， Gardners Diction-

than than 6 mm long ....・ H ・....・ H ・.....・ H ・.....・ H ・..… 7 aη" ed. 8， n .4 (1768). 7. 7. Inflorescence racemose. Flowers usually Veratrum scapo fistuloso squamoso ， spica nodding ，campanulate ，white ，occasionally stricta. Trew ，P1. Selec t. n.4 1，t. 77 (1 771) ，p.p. become tinged with pink after a peak of Helonias latifolia Michx. ， F 1. Bor.-Amer. flowering. flowering. Fragrance very weak. Tepals 1: 212 (1 803); Pers. ， Syn. P1. 1: 398 (1 805); narrowly narrowly elliptic to narrowly oblong ，9.5- Spreng. ，Sys t. Veg. ed. 16 ， 2: 144 (1 825). 14.5 14.5 mm long ，slightly reflexed at apex. Distribution: Eastern North America. Leaves lustrous above ， to 31 cm long ，5 cm Note: Chromosome number 2n = 34 (Utech

wide.... ・H ・...・ H ・....・ H ・.....・ H ・. 5) H. leucantha 1980 ，Tanaka unpub ふ 7. 7. Inflorescence umbellate (occasionally Specimens examined. subumbellate). subumbellate). Fl owers not necessarily nod- NorthAmerica. New Jersey. Morris Co. ，Budd's lake ding ，cup-shaped ， white to pink ，fragran t. K. K. Mackenzie 1209 ，May 7，1905 ，fl. ， June 11 ，1905 ， 仕.， E); Hort. Edin. [cul t. plants at Royal Bo t. Gard. Tepals Tepals oblanceolate ，8-12 mm long ， not Edinb. ，May 14 ，95 (1 895 ?)， fl. E]; Cul t. at Royal necessarily necessarily reflexed at apex. Leaves to c.16 Bo t. Gard. Edinb. ，acces. n.19694729. Transplanted at cm long ， 2.2 cm wide .….. 6) H. umbellata Teikyo Univ. ，Tokyo (N. Tanaka s.n. ， Jun. 20 ，1996 ， 8. 8. Leaves relatively thick (papery to Bio 1. Lab. ， Teikyo Univ.). subcoriaceous) ， lustrous above. Flowers 2) Helonias yunnanensis (W.W.Sm. & usually usually pin k. Bottom of nectary sac usually Jeffrey) N.Tanaka ，comb. nov. located located below the level of ovary base. Tapels Ypsilandra yunnanensis W.W.Sm. & Jeffrey inNotes inNotes Roy. Bo t. Gard. Edinb. 9: 143 (1 916); 9.5-16.5 9.5-16.5 mm long ....・ H ・-… 8) H. orientalis 8. 8. Leaves relatively thin (papery) ， often Hand.-Mazz. ，Symb. Sin. 7: 1192 (1 936); minutely minutely undulate on margins. Flowers Anonym. ，Ic on. Cormophy t. Sin. 5: 425 ，t. 7680 white white or tinged slightly with pin k. Bottom (1 976); S.C. Chen ， F1. Reip. Pop. Sin. 14: 17 ， of of nectary sac usually located above the t. 5 (1 980). level level of ovary base. Tepals 8.3-1 1. 7 mm }均 7ヤ'JS ila 仰ndra αyunn αnensi 臼sW va 訂r. micrantha Hand.-Mazz. in Anzeig. Akad. long..... ・H ・.....・ H ・-…...・ H ・... 9) H. breviscapa Wiss. Wiss. WienMath. Naturw. K 1. 60: 155 (1923); Enumeration of the species Anonym. ， Icon. Cormophyt. Sin. 5: 425 3 1) 1) Helonias bullata L. )， Sp. P1.， ed. 1， 1: (1 976). 106 106 植物研究雑誌第73 巻第2号 平成10 年4月

均silandrayunnanensis W.W.Sm. & Jeffrey above Tseku and Tsehchung (J .F. Rock 10091 ，1923 ，孔， va r. himalaica Hara in H. Hara et al.， Enum. E) ，Nepa l. Kasuwa Kh ola ，13000 ft (L. W. Beer 25356 ， Aug. 21 ，1975 ，fr.， TI); ibid. ，14000 ft (J.D.A. Stainton Flow. Flow. Pl. Nepall: 80 (1978); Noltie ， F l. Bhu- 7063 ，Jun. 1，1974 ，fl. ，TI); MiddleNepa l. Southof Anna tan tan 3(1): 90 (1994). puma ， Kaski District ， above Siklis ， 18 km NNE of Distribution: Distribution: ， Bhutan， Tibet ， Pokhara ， al t. c.4 300 m (R. G. Troth 219 ，May 23 ，1972 ， and . fl. ，TI). Note: Note: Hara (1978) distinguished the dwarf 3) Helonias alpina (Wang & Tang) N. and small flowered form of Ypsilandra Tanaka ，comb. nov. yunnanensis yunnanensis as var. himalaica ， but this form Ypsilandra alpina Wang & Tang in Bull. seems not to be clearly distinguished from the Fan Mem. Ins 1. Bio l. Bo 1. 7: 81 (1936); S.C. typical typical form (v 訂. yunnanensis) due to the Chen ， F l. Reip. Pop. Sin. 14: 18 (1 980). presence presence of intermediate forms. For instance ， Distribution: Tibet and Yunnan. as as for tepals ，Hara (1978) reported that they are Specimen examined. FrontierofTibetandBurma ，Sarighkuwang ，28 0 8'N ， 2-3.5 2-3.5 mm long in var. himalaica. N oltie (1994) 9T24'E ，al t. 14000ft(F. Kingdon Ward 7084 ，9.7.1926 ， reported reported that the tepals of var. himalaica are 自.-Type ，E.). (2.5-) (2.5-) 3-4 .3 mm long. In my observations ， the 4) Helonias thibetica (Franch.) N.Tanaka ， tepals tepals of a syntype of Y. yunnanensis (G. comb. nov. Forrest Forrest 12055 ，E) were c.4 .5-5.5 mm long. Ypsilandra thibetica Franch. in N ouv. Arch. Therefore ， there is no appreciable difference Mus. His 1. Na 1. Paris ser. 2， 10: 93 ，1. 17 (1 887); in in the length between the syntype (va r. Pl. David. 2: 132 ，t.1 7 Diels (1888); in Eng l. yunnanensis) yunnanensis) and var. himalaica. As a matter Bo 1. Jahrb. 29: 240 (1900); Anonym. ，Ic on. of of fact ， in the specimens 1 examined tepals the Cormophy 1. Sin. 5: 425 ，1. 7679 (1 976); S.C. varied varied in length from c.2.7 to 6.5 mm almost Chen ， F l. Reip. Pop Sin. 14: 17 ，1. 4 (1980); continuously. continuously. Further ， no otherqualitative dif- G.H. Yang ， F l. Sichuan. 7: 11 ， p l. 3， f.I-3 ference ference was observed between the two varie- (1991). ties. ties. Accordingly ，var. himalaica appears to be Ypsilandra cavaleriei H. Lev. & Vaniot in inseparable inseparable from the typical form. Mem. Pon t. Accad. Rom. Nuovi Lincei 23: In In the meantime ，a similar dwarf and small 375 (1 905); Anonym. ，Ic on. Cormophy 1. Sin. flowered flowered form of Y. yunnanensis was reported 5: 425 (1976); S.C. Chen ， F l. Reip. Pop. Sin. by Handel-Mazzetti (1 923) from the north- 14: 17 (1 980). westem part of Yunnan ，China ，and named as 均silandra kansuensis R.N. Zhao & Z.X. Y. Y. yunnanensis var. micrantha Hand.-Mazz. Peng in Ac 1. Bo 1. Bor.-Occ. Sin. 7( 1): 57 ，f .l According According to him ， the perianth and stamens of (1987). this this variety are 3-5 mm long and the flowering Distribution: China; ，Sichuan ， plants 紅 e not more than 6 cm in heigh 1. This ， ， and . variety variety was later reduced to Y. yunnanensis by Note: Accordingto Chen (1980) ，均 silandra Handel-Mazzetti Handel-Mazzetti himself (1936) and by Chen thibetica and Y. cavaleriei have some differ- (1 980) ，and 1 also agree with this reduction by ences in floral characters. For example ， the them. them. two species are reported to differ in the extent Representative Representative specimens examined. of lobation of ovaries. That is ， the of lobation China. China. Yunnan ，Mekong-Salween divide ， la t. ovaries ovaries of Y. thibetica is deeper than that of Y. 28 0 lO 'N ，alt. 12000 ft (G. Forrest 14961 ，Sep t. 1917 ，fr.， E); E); Yunnan ， Shweli-Salween divide ， la t. 25 0 20'N ， al t. cavaleriei ， the lobed part of the former species 9000 ft (G. Forrest 12055 ，Aug. 1913 ，fl.-Syntype ，E.); being 2/5- 1/ 3 of the entire ovary length ， while N.W. Yunnan ，Mekong-Salween watershed ， mountains that of the latter being 1/4-1/5. However ， in April1998 April1998 Journal of Japanese Botany Vo l. 73 No. 2 107

my observations there was no significant diι in Fig. 3 of my paper (Tanaka 1997 e) does not ference ference in the extent of the ov 紅 y lobation in c1 ude the locality in Gansu. between the two species. Chen (1 980) also Chromosome number 2n = 34 (Tanaka reported reported that at flowering time the stamens and unpub ふ pistils pistils of Y. thibetica far exceed the perianth ， Representative specirnens exarnined. wh i1 e those of Y. cavaleriei only sligh t1 y ex- China. Sichuan. M t. Orne i. cul t. at Nikko Bo t. Gard. ceed ceed the perianth. In the specimens I examined Univ. Tokyo [May 12 ，1996 ，(日 .)-fr. MAK]; Guizhou ， Pin-fa ，rnontagnes ， liux hurnides (J. Cavalerie 1291 ， similar a similar tendency was present ， but at the same Mars 10 ，1902 ，fl. ， E); Guizhou. al t. 1450 rn (J. Esquirol time ， there were also some individuals that 2013 ， Avril 1910 ，日.， E); S. E. Guizhou ， environs de show the intermediate state conceming the Touchan ， dans la rnontagne (J. Cavalerie 2638 ， Juin 3， lengths lengths of these floral organs (e.g. ，Cavalerie 1899 ，fr. - Type of Ypsilandra cavaleriei ，E); Wild origin 1291 ， E). As for the floral size ，Chen (1 980) unknown (plants obtained frorn cultivated sources in Japan) ，cultivated at Teikyo University (N.Tanaka s.n. ， reported reported that the tepals of Y. cavaleriei are 4- Apr. 6，1986 ，fl. - only flowers; Jun. 8，1986 ，fr.; March 6 mm long ， while those of Y. thibetica are 6- 6，1988 ，乱; Ap r. 24 ，1988 ，fl.-f r.; March 14 ，1989 ，fl.; 10 10 mm long. This report indicates that the May 20 ，1994 ，fr.， Bio l. Lab. Teikyo Univ.). variation variation of the tepallength is virtually con- 5) Helonias leucantha (Koidz.) N. Tanaka ， tinuous tinuous between the two species. In fact ， in my comb. nov. material material the tepallength varied from 4 to 10.6 Sugerokia leucantha Koidz. in Acta mm without any significant gap. Judging from Phytotax. Geobo t. 6: 46 (1937). all all these observations ， Y. thibetica and Y. Heloniopsis leucantha Koidz. in Acta cavaleriei cavaleriei seem not to have a marked differ- Phytotax. Geobo t. 6: 46 (1 937) ， pro syn.; ence ence to be distinguished from each other in Masam. ，Enum. Tracheophy t. Ryukyu Insu 1. specific specific ran k. 9: 103 (1957); Hatus. & Amano ， F 1. Okinawa Zhao and Peng (1987) described Y. 141 (1 958); Walker ，F1. Okinawa & S. Ryukyu kansuensis kansuensis as a new species from southem Is 1. 304 (1 976). Gansu ， China. According to them ，this species Heloniopsis leucantha (Koidz.) Honda in differs differs from Y. thibetica in having deflexed Bo t. & Zoo 1. 6: 1680 (1 938); Hatus. & Amano ， linear-oblanceolate linear-oblanceolate leaves being 3-9 mm wide ， F1. Ryukyus ， South of Amami Is 1. 198 (1977) ， a flowering scape shorter than leaves ，a ra- ed. 2，303 (1 994); Shimab. ，Check List Vasc. ceme with secund flowers ， pedicels shorter F1. Ryukyu Is 1. 520 (1 990). than than the perianth ， stamens nearly as long as Heloniopsis leucantha (Koidz.) Hatus. ， F1. tepals ，obtriangular capsules which are 1 cm in Ryukyus 786 (1 971) ， ed. added & corrected ， diameter ，triparted almost to the central axis ， 786 (1 975) ，comb. nud.

and of which the lobes 訂 e patent ，etc. How- Japanese name: O-shiro 寸 lojobakama

ever ，many of these characters 紅 e also often (Koidzumi 1937). seen seen in Y. thibetica. As far as the original Distribution: The Ryukyu Islands description description and figure of Y. kansuensis were (Tokunoshima ，Okinawa ，Ishigaki and Iri omote surveyed ， Y. kansuensis seems not to have a Islands). sufficient sufficient difference worthwhile of its specific N ote: Hatusima (197 J， 1975) noted that delimitation delimitation from Y. thibetica. Heloniopsis leucantha and H. acutifolia may It It was in the course of preparing this paper not be different species. But ， the latter differs that that I knew of the existence of a report on Y. from the former in having a more slender kansuensis kansuensis by Zhao and Peng. Accordingly ， scape ， fewer flowers a scape ，an umbellate the the geographical range of Y. thibetica depicted inflorescence ，oblanceolate tepals ，etc. These 108 108 植物研究雑誌第73 巻第2号 平成10 年4月 two taxa can be regarded as distinct species. In Masam. ，Enum. Tracheophy t. Ryukyu Insu l. this this paper ，however ， H. acutifolia was treated 9: 103 (1 957) ，p.p. as as conspecific with Helonias umbellata [see Heloniopsis taiwaniana Ying ，Co l. 111. Herb. the the next entry (6)]. Chromosome number 2n = Pl. 1: 222 ， f.108 (p.67) (1980).

34 (Tanaka unpub.). Japanesen 仰 e: Hime-shojδbak 翻 a(Haya ね Representative Representative specimens examined. 1917) ，Shima-shojobakama(Kawakami 1910). Japan. Japan. Okinawa Pre f. Okinawa Is 1.: Kunigami-gun ， Distribution: Taiwan. Haneji-son ， M t. Taniyu-dake (T. Kanashiro 162 ，Jan. 5， Note: According to Hayata (1 920) ， 1938 ，自.， RYU 2508); Haneji-son ，M 紘iya-ku ，Uishiga- yama (S. Nakamine 322 ，Nov. 29 ，1954 ，RYU 41828 ， Heloniopsis acutifolia differs from H. TNS 116547 ， 116548); Ogimi-son ，Nuhaogawa (T. umbellata in having longer anthers and acute Amano 6331 ，Aug. 16 ，1951 ，TNS 129705); Nago-shi. leaves. But ， in my observations these two Meijiyama (H. Higa ， Jan. 31 ，1975 ，fl. ，RYU 35723); species do not differ significantly in these near near Meijiyama Bo t. Gard. (no collector name ，自.， RYU 10041).lshigakils l.: (Y. Niiro ， Aug. 1958 ，RYU 19301). characters ，and coincide well with each other lriomote lriomote Is 1.: M t. Komidake ， al t. 400 m. (T. Yamazaki ， in many characters. Therefore ，1 regard H. June June 17 ，1971 ，Tl) ，ibid. (S. Hatusima 35249 ， Aug. 3， αcutifolia as conspecific withH. umbellata ， as 1973 ，RYU 12555); Urauchi R.， al t. 150m( T. Yamazaki ， Chen (1 980) has already done. There also June June 13 ，1971 ，Tl). seems to be no ground to accept Heloniopsis 6) 6) Helonias umbellata (Baker) N.Tanaka ， arisanensis arisanensis Hayata ex Honda as a distinct comb. nov. species. species. This species was described as differ- Heloniopsis Heloniopsis umbellata Baker in J. Bo t. 12: ing ing from H. umbellata in having very short- 278 (1 874) ，“ Heleniopsis"; in J. Linn. Soc. pedicellate pedicellate flowers. But ， in many respects this Bo t. 17: 460 (1 879); Wright in J. Linn. Soc. species species accords well with H. umbellata ，and Bo t. 36: 140 (1 903); Matsum. ， Ind. Pl. Jap. falls falls under the variation range of. the latter 2( 2( 1): 197 (1905); Matsum. & Hayata in J. Co l. species. species. The flowers of the type specimen of Sci.Imp. Sci.Imp. Univ. Toky022:442 (1 906);Makino H. H. arisanensis still seem to be in the young & Nemoto ， F l. Jap. ，1260 (1 925) ， ed. 2，1547 stage. Both pistils and stamens are short. The (1 931); Anonym. ， Icon. Cormophy t. Sin. 5: anthers anthers are stiU not dehisced ，and the stigmas 426 ，t. 7681 (1976); Liu & Ying ， F l. Taiwan appe 紅 to be somewhat immature. The short 5: 5: 55 (1 978); S.C. Chen ，Fl. Reip. Pop. Sin.14: pedicels pedicels of this specimen may partly be due to 15 ，t. 3 (1 980). this this early stage of developmen t. Meanwhile ， Heloniopsis Heloniopsis acutifolia Hayata ， Ic. Pl. Heloniopsis Heloniopsis taiwaniana was described by Ying Formos. 9: 144 ，f. 53 (1 920); Makino & (1980) (1980) as a new species near H. orientalis. But ， Nemoto ， F l. Jap. 1259 (1 925) ， ed. 2，1547 judging judging from his description and colour photo- (1 931). graph ，it appe 紅 s that his species is nothing but Sugerokia Sugerokia acut 扮lω(Hayata) Koidz. ， F l. H. H. umbellata. He described H. taiwaniana as Symb. Ori.-Asia t. 95 (1 930). having having greenish perianth-segments ， but his Sugerokia Sugerokia umbellata (Baker) Koidz. ， F l. specimen was collected as late as Apr i1 20 in Symb. Ori.-Asia t. 95 (1 930). the the northern part of Taiwan at altitudes be- Heloniopsis Heloniopsis arisanensis Hayata ex Honda tween ca. 300-1000 m. 1t is a general habit that in in Bo t. & Zoo l. 6(10): 1679 (1938). the the perianths of H. umbellata turn greenish Sugerokia Sugerokia arisanensis (Hayata ex Honda) after after anthesis. The flowers of H. taiwaniana in Koidz. Koidz. inActaPhytotax. Geobo t. 8: 53 (1 939). his his photograph are apparently after a peak of Heloniopsis Heloniopsis orientalis (Thunb.) C. Tanaka flowering. flowering. Therefore 1 also regard H. va r. yakusimensis auc t. non (Masam.) Ohwi: taiwaniana taiwaniana as conspecific with H. umbellata. April1998 April1998 Journa1 of Japanese Botany Vo l. 73 No. 2 109

As far as I am aware ， Helonias in Taiwan is Hatus. ，F 1. Ryukyus 786 (1 971) ， ed. added & represented represented by only one endemic species ，H. corrected ，786 (1 975); Hatus. & Amano ，F 1. umbellata. umbellata. Chromosome number 2n = 34 Ryukyus ， South of Amami Is 1. 198 (1 977). (Chuangeta 1. 1962 ，Hsu 1971 ，Tanakaunpub ふ Japanese name: Ko-shojobakama (Koidz.

Representative Representative specimens examined. 1937) - Okinawa 同 shojδbakama (Sakaguchi Taiwan. Taiwan. Taipei Co.: Yanmingshan to M t. Chihsing ， 1924). t. al t. 600-1100 m (T. Shimizu 11117 ， Feb. 26 ，1961 ，fl. ， Distribution: The Ryukyu Islands (Amami- TI); TI); M t. Chihsing (Shichiseizan) (N. Fukuyama 3015 ， Jan. Jan. 25 ，1932 ，fl. ，TNS 59639) ，ibid. (S. Sasaki & T. oshima ，Tokunoshima ，Okinawa ，Miyako ， Soma ， Feb. 21 ，1921 ，fl. ，TNS 321215); M t. Tatun Ishigaki and Iriomote Islands). (Daitonzan) (Daitonzan) (S. Sasaki ， Apr. 20 ，1932 ，fr.， TNS 321209) ， Note: This species has occasionally been ibid. ibid. (T. Sato 127 ，Oc t. 26 ，1935 ， TI); Tatunshan- confused with Helonias umbellata ofTaiwan ， Hsinpeitou Hsinpeitou (1. Sasaki 831 ，Oc t. 11 ，1965 ，TI). Il an Co.: but but these two species can clearly be distir ト t. M t. Taiping (al t. 2000 m ，M. Tamura et a1. 21439 ， Ju 1. 30 ， 1963 ，fr.， TI) ，ibid. (C.c. Chuang&M.T. Ka04088 ， Mar. guished from each other ，and can be regarded 2，1966 ，fl. ，TI) ，ibid. (M.T. Kao 6796 ， Mar. 4，1967 ，fl. ， as different species. Chromosome number 2n TI); TI); M t. Bonbon-zan (May 10 ，1917 ，fl. - Type of = 34 (Tanaka unpub.). Heloniopsis Heloniopsis acutifolia Hayata ，TI). Taichung Co.: M t. Representative Representative specimens examined. Lee-shan Lee-shan (C. S. Feung & M.T. Kao 4554 ， Apr. 20 ，1962 ， Japan. Japan. Kagoshima Pref. Amami-oshima Is 1.: M t. 仕.， TI). Chiayi Co.: M t. Alishan (Arisan) ，2500 m (U. Yuwan-dake (T. Amano 6500 ，May 20 ，1951 ，RYU Faurie Faurie 942 ，Jun. 1914. - Type of Heloniopsis arisanensis 25059). 25059). Okinawa Pref. Okinawa Is 1.: Kunigami ，Aha ， Hayata ex Honda ， TI). Taitung Co.: Chinshuiying Tanaga-gumui (T. Amano 7474 ， Dec. 21 ，1954 ，RYU (Sinsuiei) (Sinsuiei) (S. Sasaki ， Feb. 10 ，1925 ，fl. ，TNS 321217). 28030 ，TNS 116896) ，ibid. ， al t. 100 m (S. Hatusima County unspecified: Central mountains ， Abi-sen (T. 18309 ，Jun. 24 ，1955 ，fr. ， TI; T. Yamazaki ，Jun. 23 ，1971 ， Kawakami & S. Sasaki ，Jan. 2，1911 ，fl. ，TI). TI). TI). Miyako Is 1.: T. It o 1202 ，Aug. 9，1894 ，TNS 60738). 7) 7) Helonias kawanoi (Koidz.) N.Tanaka ， Ishigaki Is 1.: M t. Omoto-dake (Y. Niiro ，Nov. 5，1958 ， comb. nov. late fl. ，RYU 19300). Ir iomote Is 1.: Gozadake (T. Heloniopsis kawanoana Koidz. in Yamazaki ， Dec. 26 ，1963 ，fr.， TI ，RYU 27147); Otomi (M. (M. Furuse 1080 ，Sep t. 19 ，1972 ，fl. ，RYU 5724); Urauchi S.Sakag. ， Gen. Index F 1. Okinawa 86 (1 924) ， R.， R.， by the upper stream (M. Furuse 4414 ，Oc t. 26 ，1973 ， nom. nud. RYU 17010). Sugerokia Kal γanoi Koidz. in Acta 8) Helonias orientalis (Thunb.) N.Tanaka ， Phytotax. Phytotax. Geobo 1. 6: 46 (1937). comb. nov. Heloniopsis Heloniopsis K ω， vanoi Koidz. in Acta Scilla bifolia auc 1. non L.: Thunb. ，F 1. Jap. Phytotax. Phytotax. Geobo 1. 6: 46 (1 937) ， pro syn.; 138 (1 784). Hatus. Hatus. & Amano ，F 1. Okinawa 141 (1 958). Scilla orientalis Thunb. in Trans. Linn. Heloniopsis Heloniopsis Kawanoi (Koidz.) Honda in Soc. 2: 334 (1 794); Willd. ， Sp. Pl. 2(1): 129 Bo 1. & Zoo 1. 6: 1680 (1 938); Walker ，F 1. (1 799); Pers. ， Syn. Pl. 1: 365 (1805); Thunb. ， Okinawa & S. Ryukyu Is 1. 304 (1 976) ，exc 1. Pl. Jap. Nov. Sp. 6 (1 824); Schul 1. & Schult .f.， syn. “? H. orientalis va r. yakusimensis Sys 1. Veg. 7(1): 570 (1829); Kunth ，Enum. P 1. (Masam.) Ohwi"; Shimab. ，Check List Vasc. 4: 330 (1843). F 1. Ryukyu Is 1.， 520 (1 990); Hatus. & Amano ， Heloniopsis pauciflora A. Gray in Mem. F 1. Ryukyus ， South of Amami Is 1.， ed. 2，303 Amer. Acad. Art. Sci. new ser. 6: 416 (1 859); (1994). (1994). Maxim. in Bull. Acad. Imp. Sci. S1. Petersb. Heloniopsis Heloniopsis orientl αlis (Thunb.) C. Tanaka 11: 436 (1 867); Franch. & Sav. ，Enum. P1. Jap. var. var. yakusimensis auc 1. non (Masam.) Ohwi: 2: 87 (1 877)2) ，“ Helionopsis"; Baker in J. Masam. ，Enum. Tracheophy 1. Ryukyu Insu 1. Linn. Soc. 17: 460 (1 879) ，exc l. Miq. 9: 9: 103 (1 957) ，p.p. Sugerokiajaponica Miq. in Ann. Mus. Bo 1. Heloniopsis Heloniopsis umbellata auc 1. non Baker: --‘nu EEA 植物研究雑誌第73 巻第2号 平成10 年4月

Lugd.-Ba t. 3: 145 (1 867)1) p.p.? Sugerokia orientalis (Thunb.) Koidz. ， F l. Heloniopsis Heloniopsis japonica (Miq.)7) Maxim. in Symb. Ori-Asia t. 95 (1 930). Bull. Bull. Acad. 1mp. Sci. St. Petersb. 11: 437 Heloniopsis japonica (Thunb.) Maxim. var. (1867); (1867); Franch. & Sav. ，Enum. Pl. Jap. 2: 87 purpurea Nakai ， et var. sanguin ωNakai in (1877)2) “Helionopsis" ，p.p.; Baker in J. Linn. Bo t. Mag. Tokyo 47: 243 (1 933).

Soc. Soc. 17: 460 (1 879) ，p.p.; in Bo t. Mag. 114: t. Heloniopsisjaponica (Thunb.) Maxim. v紅. 6986 (1888) ，p.p.; WrightinJ. Linn. Soc. Bo t. grandiflora N akai in Bo t. Mag. Tokyo 47: 243 36: 36: 140 (1903) ，p.p.; M 紘 ino in 1inuma ，Y. ， (1933) ，quoad basionym. Somoku-Dzusetsu2: 48 ，p1. 48 (p.378) (1 910); Heloniopsis japonica (Thunb.) Maxim. f. Miyabe and Kudo ， F l. Hokkaido & Sagha l. 3: lateriti αNakai ex Honda in Bo t. & Zoo l. 309 (1 932) ，p.maj.p.; Nakai in Bo t. Mag. To- 6(10): 1679 (1938). kyo 47: 242 (1933) ，p.p. ， ut auc t. (Thunb.); Heloniopsis acutifolia auc t. non Hayata: Honda in Bo t. & Zoo l. 6: 1677 (1938) ，p.p.; Honda in Bo t. & Zoo l. 6(10): 1678 (1938). Makino ，111. Fl. Jap. 758 ，t. 2274 (1940) ， ed. Sugerokia acutifolia auc t. non (Hayata) rev. ，758 ，t. 2274 (1 949). Koidz.: Honda in Bo t. & Zoo l. 6 (10): 1678 Heloniopsis Heloniopsis grandiflora Franch. & Sav. ， (1938). Enum. Pl. Jap. 2: 88 (1 877)2) ，nom. nud. ， et Heloniopsis orientalis (Thunb.) Koidz. var. 529 (1 878)2) ，“ Helionopsis" . purpurea Nakai inBull. Na t. Sci. Mus. 31: 146 Heloniopsis Heloniopsis breviscapa auc t. non Maxim.: (1 952); Ki m ，Wild Flow. Korea 450 (1988). 恥tlakino in Bo t. Mag. Tokyo 5: 126 (1891); Heloniopsis orientalis (Thunb.) C. Tanaka Matsum. 1nd. Pl. Jap. 2(1): 197 (1 905) ，p.p. f. lutea Mochizuki in 1. Geobo t. 26(1): 14 Heloniopsis Heloniopsis pauciflora A. Gray var. rubra (1978). H. H. Koidz. in Shinano Kyoiku ， no. 421: 15 Japanesename: Shojobakama(K. It o 1829). (1921) ，cum desc r. jap. Distribution: Southern Sakhalin ，Hokkaido ， Heloniopsis Heloniopsis orientalis (Thunb.) T. Tanaka Honshu ，Shikoku ， northern Kyushu and Ko- (= (= C. Tanaka) in Bul t. Sci. Terku l. Kjusu Imp. rea. Univ. Univ. 1: 200 (Sep t. 1925); Ohwi ， F l. Japan ， Note: This species usua11y has pinkflowers ， 285 (1 953) ， ed. Eng l.， 284 (1 965) ，ed. rev. 336 but it is known that there is some variation (1975) ，p.p. ，quoad var. orientalis; Kitam. in regarding the floral colour; e.g. ，brick-red ， Kitam. Kitam. eta l.， Co l. 111. Herb. Pl. Jap. (Monocot.) blood-red ，red ，blue ， pale or deep pink ，pur- 151 151 (1 964) ，p.p.; Y.N. Lee ，Ill. Fl. & Fauna of plish pink ，white ， pale yellow ， or yellow. Chro- Korea 18: 156 ，781 ， p l. 9， fig. 1 (1 976); mosome number 2n = 34 (Sato 1942 ，Nakamura Okuyama & S. Kobay. in Okuyama ed. ， 1967 ，Tanaka unpub ふ Terasaki' Terasaki' s Ill. Fl. Jap. 874 ，t. 3314 (1977); T.B. Representative specimens examined. Lee ，Ill. Fl. Korea ，2nd prin t.， 196 (1 982); Japan. Hokkaido. Soya. Utanobori (M. Yamanoi ， Satake Satake in Satake et al.， Wild Flow. Jap. Herb. May 6，1972 ，fl. ，TNS 319544). Sarobetsu Genya (H. Hara & S. Kurosawa ，Jun. 11 ，1976 ，fl. ，TI). Kamikawa. l. Pl. 1: 26 (1 982) ，p.p. ，quoad var. orientalis; Taisetsu Mts. ， M t. Asahi-dake (H. Koidzumi ，Aug. 3， H. Ohba in Kimura et Leonov ，Thunb. Draw. 1917 ，fl. ，TNS 315850 ， 315851). Hidaka. M t. Apoi (H. Jap. Jap. Pl. 445 (1994) ，p.p. Hara ，May 26 ，1933 ，fl. ，TI). Heloniopsis Heloniopsis orientalis (Thunb.) Koidz. in Honshu. Tohoku: Aomori Pref. M t. Hakkoda (K. Hasegawa ， Jun. 30 ，fl. ，TI). Aki ta Pref. Senpoku-gun ， Bo t. Mag. Tokyo 39: 312 (Dec. 1925) ，exc l. t. M t. Komagatake ， al t. 1300 m (Y. Tateishi 2123 ， Ju l. 5， syn. syn. Maxim.; Makino & Nemoto ， F l. Jap. ed. 1975 ，fl. ，TI). Iwate Pref. Hienuki-gun ， M t. Hayachine ， 2，1547 (1931); Terasaki ，Nippon Shokubutsu Odagoe ， al t. 1200 m (S. Okuyama 12327 ， Ju l. 18 ，1967 ， Dzufu 93 (1 933) ，“ Helonoipsis". fr. ，TNS 177907). YamagataPre f. M t. Gassan (R. Yatabe ， l4 咽'. 目目 April1998 April1998 Journal of Japanese Botany Vo l. 73 No. 2 A

1. Ju 1. 23 ，1887 ，fl. ，TI). Kanto: Gunma Pre f. M t. Haruna- J. Jpn. Bo t. 21: 148 (1 947); Okuyama ，Co l. 111. san san (N. Maruyama ，May 6，1948 ，fl. ，TNS 81854 ， 81855). Wild Pl. Japan (1 0th prin t.) 1: 29 (1974); Ibaraki Ibaraki Pre f. Taga-gun ， Kawajiri-kaigan (M. Komatsuzaki ， Apr. 10 ，fl. ，MAK 137629). Tokyo. Okuyama & S.Kobay. in Okuyama ed. ， Kozushima Is 1.， M t. Te 吋o-san ， al t. 400 m (T. Yamazaki ， Terasaki's Ill. Fl. Jap. 874 ，t. 3315 (1 977). Apr. Apr. 14 ，1976 ，fl. ，TI). Chubu: Niigata Pref. M t. Naeba Heloniopsis japonica auc t. non Maxim.: (S. (S. Okuyama ， Ju 1. 19 ，1936 ，fl. ，TNS 53019). Nagano Franch. & Sav. ，Enum. Pl. Jap. 2: 87 (1 877)2) ， Pref. Pref. M t. Senjo-dake (H. Koidzumi ， Ju 1. 23 ，1923 ，自.， “ TNS 315696). Fukui Pref. Takefu ，Murakuniyama ， al t. Helionopsis ぺp.p.; Makino in Bo t. Mag. To- 150 150 m (T. Yamazaki ，M 訂. 26 ，1987 ，日.， TI). Ki nki: kyo 5: 126 (1 891); Matsum. ，Ind. Pl. Jap. 2(1): Kyoto Pref. Takeno-gun ，Yasaka-cho ，Midono ， al t. 300 197 (1 905) ，p.p. m (G. Murata ，20968 ，fl. ，TNS 304009). Mie Pref. M t. Sugerokia nipponica Ohwi in Bo t. Mag. Fujiw 紅 a-dake (S. Okuyama ，fl. ，TNS 102335 ， 102338). Tokyo 44: 566 (1930). Chugoku: Tottori Pre f. Saihaku-gun ，Daisen-machi ， M t. Daisen Daisen (N. side) (K. Midorikawa ，2215 ，May 14 ，1981 ， Heloniopsisjaponica(Thunb.)Maxim. var. fl. ，TI). Yamaguchi Pref. Hagi-machi (J. Nikai ，Ap r. 22 ， flavida Nakai in Bo t. Mag. Tokyo 47: 243 1929 ，fl. ，TNS 49293). (1 933); HondainBo t. &Zoo l. 6: 1679 (1 938). Shikoku. Shikoku. Kagawa Pref. M t. Otakiyama (Y.切 ike ， Heloniopsis Heloniopsis japonica (Thunb.) Maxim. v紅. Apr. Apr. 28 ，1939 ，fr. ，TNS 79928). Tokushima Pre f. M t. Tsurugisan Tsurugisan (H. Ki mura ， Jun. 5，1956 ，fl. ，TNS 139424). grandiflora Nakai in Bo t. Mag. Tokyo 47: 243 Kochi Pref. Takaoka-gun ，Sakawa-cho ，Ogawa ， (1 933) ，p.p. Minezaka (T. 乱1akino ，May 21 ，1889 ，fl. ，MAK 137670). Heloniopsis japonica Maxim. va r. Korea. Korea. Pyongannamdo. Yangdok (Yotoku) (T. Nakai yakusimensis Masam. in Mem. Fac. Sci. & 12478 ， Jun. 15 ，1928 ，fr. ， TI). Kyongsangnamdo. M t. Chiri-san ， c.1600-1700 m (K. Uedaet a1. 1220 ， Jun. 24 ， Agr. Taihoku Imp. Univ. 11: 551 (1 934). 1979 ，fr. ，TI 1313106 ， 1314994). Heloniopsis nipponica (Ohwi) Nemoto ，Fl. 9) 9) Helonias breviscapa (Maxim.) N. Jap. Supp l. 1058 (1 936); Hiyamain Yaso 5(1): Tanaka ，comb. nov. 15 (1 939); Okuyama ，Co l. 111. Wild Pl. Jap. 8 Scilla Scilla japonica Thunb. ) ex Murray ，Sys t. (7th prin t.) 6: 12 (1 974). 9 Veg. ， ed. 14 ，329 (Mai-Jun. ) 1784) ， ed. 15 ， HeloniopsisjaponicaMaxim. var. alb ♂ ora 349 (1 798); Thunb. ，Fl. Jap. ， 137 (Aug. 1784); Honda ， et var. breviscapa (Maxim.) Honda in Icon. Icon. Pl. Jap. 4 ，t. 4 (1802); Gmelin ，Sys t. Na t. Bo t. & Zoo l. 6: 1678 (1 938). 2(1): 2(1): 552 (1 791); Wi11d. ， Sp. Pl. 2(1): 127 HeloniopsisjaponicaMaxim. v ぽ .tessellata (1 799); Pers. ， Syn. Pl. 1: 366 (1805); Thunb. ， (Nakai) ex Honda in Bot. & Zoo l. 6: 1679 l. Pl. Jap. Nov. Sp. ，6 (1824); Schul t. & Schul t. f. (1938). Sys t. Veg. 7(1): 580 (1829); Kunth ，Enum. Pl. Heloniopsis yakusimensis (Masam.) Honda 4: 4: 330 (1 843). in Bo t. & Zoo l. 6: 1679 (1 938). Sugerokia Sugerokia japonica Miq. in Ann. Mus. Bo t. Sugerokia yakusimensis (Masam.) Koidz.

Lugd. ーBa t. 3: 145 (1 867) 1)， p.p.?; Koidz. ， F l. in Acta Phytotax. Geobo t. 8: 53 (1 939). Symb. Ori.-Asia t. 95 (1 930). Heloniopsis brevisc αpa Maxim. va r. Heloniopsis Heloniopsis pauciflora auc t. non A. Gray: yakusimensis (Masam.) H. Hara in 1. Jpn. Bo t. Miq. Miq. in Ann. Mus. Lugd.-Ba t. 3: 146 (1 867) ， 21: 148 (1 947). Helionopsis". “Helionopsis". Heloniopsis breviscapaMaxim. var.flavida Heloniopsis Heloniopsis breviscapa Maxim. in Bull. (Nakai) Hara ， et var. albiflora (Honda) H. Acad. Acad. Sci. St. -Pe t. 11: 436 (1 867); Franch. & Hara in 1. Jpn. Bo t. 21: 149 (1 947). Sav. ，Enum. P 1. Jap. 2: 87 (1877)2¥ Heloniopsis orientalis (Thunb.) C. Tanaka Helionopsis"; “Helionopsis"; Baker in 1. Linn. Soc. 17: 459 var. breviscapa (Maxim.) Ohwi ， F l. Jap. 285 (1 879) ，exc l. syn. Franch. & Sav.; Matsum. ， (March 1953) ，comb. nud.; in Bul l. Na t. Sci. Ind. Ind. Pl. Jap. 2(1): 197 (1905) ，p.p.; H. Hara in Mus. Tokyo 33: 68 (J uly 1953); Fl. Jap. ed. 112 植物研究雑誌第73 巻第2号 平成 10 年4 月

Eng l. 284 (1 965) ， ed. rev. 336 (1 975); Satake yakusimensis ，v 訂 .α lbiflora and v紅 .flavida ，

in in Satake eta l.， WildFlow. Jap. Herb. Pl. l: 26 besides the typical form (v 訂 . breviscapa). (1 982). But ， in my observations these forms cannot Heloniopsis Heloniopsis orientalis (Thunb.) C. Tanaka clearly be distinguished from each other ，due

V 紅 .y αkusimensis (Masam.) Ohwi ， F l. Jap. to the presence of intermediate forms. 285 (March 1953) ，comb. nud.; in Bu11. Na t. Helonias breviscapa has its range in the Sci. Sci. Mus. Tokyo 33: 68 (J uly 1953); Fl. Jap. south of the range of its closest species ， H. ed. ed. rev. 336 (1 975); Masam. ，Enum. orientalis ， with some overlap (Tanaka 1997b ， Tracheophy t. Ryukyu Insu l. 9: 103 (1 957) ， 1997 e; under the generic name of H eloniopsis). p.p. ，quoad pl. ex Yakushima. The area predominated by H. breviscap αIS Heloniopsis Heloniopsis orientalis (Thunb.) C. Tanaka genera11y characterized by a warmer climate var.flavida var.flavida (Nakai) Ohwi ， F l. Jap. 285 (March in winter with less snowfa11 than the area 1953) ，comb. nud.; in Bu11. Na t. Sci. Mus. predominated by H. orientalis. Helonias Tokyo 33: 68 (July 1953); Fl. Jap. ed. Eng l. breviscapa has thinner leaves thanH. orientalis 284 (1965) ， ed. rev. 336 (1 975); Ki tam. in (Tanaka 1997d ， 1997e). It seems likely that Ki tam. et al.， Co l. Ill. Herb. Pl. Jap. 3: 151 this interspecific difference in the leaf texture (1964); (1964); Satake in Satake et al.， Wild Flow. Jap. is related to the difference in the winter climate Herb. Herb. Pl. 1: 26 (1 982). covering the ranges of the two species. Chro- Heloniopsis Heloniopsis orientalis (Thunb.) C. Tanaka mosome number 2n = 34 (Nakamura 1967 ， subsp. subsp. breviscapa (Maxim.) Ki tam. in Ki tam. Tanaka unpub.). et et al.， Co l. 111. Herb. Pl. Jap. 3: 151 (1964) ， Representative specimens examined. nom. inva l.; Ki tam. & Muratain ActaPhytotax. Japan.Honshu.Kanto:ChibaPref.Awa-gun ，Amatsu- Geobo t. 22(3): 72 (1966). kominato-cho ， M t. K.i yosumi (T. Makino ， Apr. 1896 ，fl. ， MAK 137585 ，137586) ， M t. Ki yosumi (T. Nakai ， Apr. Heloniopsis yakusimensis (Masam.) 1937 ，fl. - Type of Heloniopsisjaponica var. tessellata Masam. ，Co l. Ill. Fl. Nippon 8: 11 (1 969). Nakai ex Honda ，TI) ， Mt. K.i yosumi (H. Motomiya ， Apr. Heloniopsis Heloniopsis orientalis C. Tanaka ssp. 5，1970 ，fl. ，TNS 280174). Tokyo. Tama-shi ， Sekido (T. breviscapa breviscapa Ki tam. f. satsumensis Doi ，Enum. Sakai ，Ap r. 3， 1938 - Type of Heloniopsisjaponica var. lbiflora αlbiflora Honda ，TI). Kanagawa Pref. ，Tanzawa Mts. ， l. Pl. in Pre f. Hiroshima Spon t. Nascent. 135 Miyagase (T. Saito ， Apr. 21 ，1963 ，late fl. ，TNS 156935 ， (1 983) ，nom. seminud. ，cum descrip. jap 156936). Hakone-machi ， Hatajuku -M t. Shirogane- (p.143). (p.143). yama(Y. Tateishi&M.Fukuda2054 ，Jun.18 ，1975 ，TI). Heloniopsis Heloniopsis orientalis auc t. non (Thunb.) Chubu: Shizuoka Pref. Hakone ，Higane-yama (H. Hara ， Tanaka: Tanaka: H. Ohba in Ki mura & Leonov ， Thunb. Apr. 15 ，1956 ，fl. ，TI). K.i nki: Mie Pref. Suzuka-gun ， Seki-cho ，Bandoyama ， al t. 250 m (H. Koyama & N. Draw. Jap. Pl. 445 (1994) ，p.p. Fukuoka 185 ， Apr. 3，1963 ，fl. ，TNS 179700). Kyoto. M t. Japanese name: Kocho-shojobakama Hiei (T. Nakai ， Mar. 29 ，1911 ，fl. ， TI). Osaka. M t. (newly (newly named; after the J apanese name ‘Ko り0' Kongo-san (M. Togashi ，Ap r. 25 ，1963 ，fl. ，TNS 105921). recorded recorded by Thunberg (1784) under Scilla Wakayama Pref. It o-gun ，Koya-cho ，ne 訂 Otaki (K. Deguchi Deguchi 6077 ，日.， TNS 594690). Kobe-sh i. M t. Rokko japonica) ，Shirobana-shojobakama (T. Iwasaki (S. (S. Kuriyama ，Jun. 8，1928 ，fl. ，MAK 137594). Chugoku: 1824) ， Tsukushi-shojobakama (Honda 1938) ， Hiroshima Pref. Sandankyo (H. Hara & S. Kurosawa ， Yakushima-shojobakama (Honda 1938). Apr.4 ，1976 ，TI). Yamaguchi Pref. Abu-gun ，Mutsumi-

Distribution: Distribution: J apan. Honshu (southern p紅 t mura ， Kawahira (N. Miake ，Ap r. 7，1971 ，fl. ，MAK of of the Kanto district and westward to the 130266). Shikoku. Shikoku. Tokushima Pref. Otakijiyama (K. Abe Chugoku district) ， Shikoku and Kyushu. 12166 ，fl. ，TNS 132384). Myodo-gun ，Sanagochi-mura ， Note: Note: Under Heloniopsis breviscapa ，Hara Sagamyo (J. Nikai ， Apr. 30 ，1913 ，仕.， TNS 79930). (1947) (1947) recognized three varieties; va r. Kochi Pref. Nagaoka-gun ，Otoyo-mura ，Tachikawa (T. April1998 April1998 Journal of Japanese Botany Vo l. 73 No. 2 113

Makino ，May 5，1893 ，late fl. ，MAK 137569). 1 could not refer to “h在orison" in this study. Kyushu. Fukuoka Pre f. Kitakyushu-shi ，Moji ， 4) 4) Adanson also cited “Morison" (see endnote Adachiyama (T. Hashimoto ，Mar. 13 ，1952， fl. ，Tl). Oita Pre f. Beppu (H. Koidzumi 42537 ，日.， TNS 315693). 3). Saga Pref. Ogi-gun ， M t. Hiko-dake (T. Hashimoto ，Ap r. 5)Year of publication followed Parr ， Leussink 22 ，1980 ，fr. ，Tl).Nagas 北 iPref. M t. Unzen (1. Matsumura and Stafleu ，Index N ominum Genericorum & G. Koidzumi， Apr. 1911 ，fl. ，Tl). Kumamoto Pref. M t. (Plantarum) 2: 917 (1 979). Aso ， near Akamizu (H. Koidzumi 42822 ，日.， TNS 6)In 6)In this study one fruiting specimen of 315694). 315694). Miyazaki Pref. Minami-naka-gun ， obi-cho (M. N agasawa ， Apr. 1946， late fl. ，TNS 132586). Kagoshima Helonias yunnanensis was newly available Pref. Pref. M t. Kaimon-dake (T. Sakata ，Mar. 30 ，1964 ，fl. ， (Beer 25356 ， TI). It s seeds were ca.2.3-3.2 TNS 161902). Yakushima Is l. (G. Masamune ， Apr. 6， mm long and 0.5-0.7 mm broad. 1927 ，fl. ，Tl). 7)Por the basionym of this name ， see Hara Besides Besides the above specimens ，1 examined a (1947). (1947). photograph photograph of the type specimen of Sugerokia 8)The 8)The name Scilla japonica Thunb. cannot be nipponica nipponica Ohwi [N ara pre f. (Y oshino) ，be- used used as the basionym in Helonias ， since in this tween Kashiwagi and Obagamine ，J. Ohwi ， genus genus there has been an earlier homonym May 19 ，1928 ，KYO] preserved at TNS (n. Helonias? Helonias? japonica (Willd.) Schul t. & Schul t. 320902). 320902). f. f. [Sys t. Veg. 7(1): 1567 (1 830)] ，which is a synonym of Chionographisjaponica (Willd.) 1 thank the staff of the. following herbaria Maxim. for for providing me with opportunities to exam- 9)Month of publication followed Stafleu and ine ine the specimens: E ，MAK ，RYU ，TI and Cowan ，Taxonomic literature ， ed. 2， 3: 670 TNS. 1 am also very grateful to the staff ofthe (1 981). Botanic Botanic Garden ofthe University ofTokyo for allowing allowing me to refer to some classical botani- References cal cal books in the library ，and to the anonymous (See also the literature cited in the tex t) reviewers reviewers of my manuscripts for advisory com- Baker J. G. 1874. On a new species of Heloniopsis from ments. ments. Formosa. J. Bo t. Bri t. & For. 12: 278. 一一一一一 1879. A synopsis ofColchicaceae and the aber- rant rant tribes of Li liaceae. Heloniopsis. J. Linn. Soc.17: Endnotes 459 -4 60. l)Mique l' s description of Sugerokia was pub- BenthamG.1883.Heloniopsis. In: BenthamG. & Hooker lished lished also in Vers. Med. Koni k. Ak ad. Wetens. J. D. ，Genera Plantarum 3(2) ， p. 827. L. Reeve ， London. r. se r. 2， 2: 88 (1 867) ，and in its reprint (1 866) ， Chen S- c. 1980. Heloniopsis. 均silandra. In: Wang F.ー but but 1 could not refer to either paper. Por the T. and Tang T. (ed よFl ora Reipublicae Popularis ye 紅 s of publications of these papers see Stafleu Sinicae 14: 15-16. Science Press ， Beijing (in Chi- F. F. A. ，Wentia 16: 84 (1 966). nese). 2)Year 2)Year of publication followed Stafleu and Chuang T. ，Chao C. Y.， Hu W. W. L. and Kwan S. C. 1962. 1962. Chromosome numbers of the vascular plants of Cowan ，Taxonomic literature ， ed. 2 ，1: 869- Taiwan. 1. Taiwania 8: 51-66. 870 (1976). Engler A. 1888. Heloniopsis. In: Engler A. und Prantl K.， 3)Linnaeus 3)Linnaeus (1753) and some other authors cited Die N aturlichen Pflanzenfamilien. 2(5): 22. Wilhelm Engelmann ，Leipzig. “Morison ，His t. 3. p. 606 ，s. 15 ，t. 2 ，f. 1." as

Franchet Franchet A. 1887. Plantae Davidianae ex Sinarum lmpe 伺 its its plant being identical with How- H. H. bullata. rio. Part II. 均silandra. Nouv. Arch. Mus. His t. Na t. ever ， Morison' s species is excluded from H. Paris ser. 2， 10: 93-95 ，t. 17. bullata bullata in Curtis' s Bo t. Mag. [20 ，t. 747 (1804)] ， 一一一一& Savatier L. 1877. Enumeratio Plantarum in Schul t. & Schul t. f. (1830) ，etc. Unfortunately Japonia Sponte Crescentium. 2. Helionopsis ， pp. 87- 88. 88. F. Savy ，bibliopolam ，Paris. 114 114 植物研究雑誌第73 巻第2号 平成10 年4月

Gray A. 1859. Diagnostic characters of new species of N akai T. 1933. N otulae ad plantas J aponiae & Koreae 43. phanerogamous plants ，collected in J apan by Charles Heloniopsis japonic α. Bo t. Mag. Tokyo 47: 242- Wright ，botanist of the U.S. north Pacific exploring 244. expedition. expedition. Heloniopsis. Mem. Amer. Acad. n.ser. 6: N akamura T. 1967. Cytological studies in family Liliaceae 416. 416. of Japan. II. The karyotype analysis in the genus Hara H. 1947. Annotationes miscellaneae ad plantas Heloniopsis. La Kr omosom071: 2316-2321 (inJapa- Asiae-Orientalis Asiae-Orientalis (III). J. Jpn. Bo t. 21: 143 ー150 (in nese). Japanese). Japanese). Ohwi J. 1953. Fl ora of Japan. Heloniopsis ， pp. 285-286. Honda M. 1938. Species of Heloniopsis. Bo t. and Zoo 1. Shibundo ，Tokyo (in Japanese). 6: 6: 1675-1680 (in Japanese). 一一一一 1965. Fl ora of J apan. English ed. He loniopsis ，p. Hsu C. C. 1971. Preliminary chromosome studies on the 284. Smithsonian Institution ，Washington ，D. C. vascular vascular plants of Taiwan (IV). Counts and some Sato D. 1942. K 訂 yotype alteration and phylogeny in systematic systematic notes on some monocoty ledons. Taiwania Li liaceae and allied families. Jap. J. Bo t. 12: 57-16 1. 16: 16: 123-136. Tanaka N. 1997 a. Taxonomic significance of some floral Kawano S. and Masuda J. 1980. The productive and characters in Helonias and 浄silandra (Liliaceae). J. reproductive reproductive biology of flowering plants VI I. Re- Jpn. Bo t. 72: 110-116. source source allocation and reproductive capacity in wild 一一一一一 1997b. Evolutionary significance of the varia- populations populations of Heloniopsis orientalis (Thunb.) C. tion ofthe floral structure of Heloniopsis. J. Jpn. Bo t. Tanaka (Li liaceae). Oecologia 45: 307-317. 72: 131-138. KoidzumiG.1930. 日orae Symborae Orientali -Asiaticae. 一一一一一 1997c. Phylogenetic and taxonomic studies on Sugerokia ， pp. 94- 96. Kyoto. Helonias ，Ypsilandra and Heloniopsis. 1. Compari- 一一一一 1934. Shojobakama zoku ni tsuite [on 血egenus son of character states (1). J. Jpn. Bo t. 72: 221-228. Heloniopsis]. Heloniopsis]. Acta Phytotax. Geobo t. 3: 42 -4 3 (in 一一一一 1997d. Phylogenetic and taxonomic studies on Japanese). Japanese). Helonias ，玲silandra and Heloniopsis. I. Compari- Kr ause K. 1930. Heloniopsis. In: Engler A. und Prantl K. son of character states (2). J. Jpn. Bo t. 72: 286-292. (ed よDie Naturlichen Pflanzenfamilien 15a: 259- 一一一一 1997e. Phylogenetic and taxonomic studies on 260. 260. Wilhelm Engelmann ，Leipzig. Helonias ，浄silandra and Heloniopsis. 11. Evolution Maximowicz C. J. 1867. Diagnoses breves plant 訂 um and geographical distribution. J. Jpn. Bo t. 72: 329- novarum J aponiae et Mandshuriae (III). H e loniopsis. 336. Bull. Bull. Acad. Imp. Sci. S t. -Petersb. 11(4): 436 -4 37. UtechF.H.1980.SomatickaryotypeanalysisofHelonias Miquel F. A. G. 1867. Prolusio Florae Iaponicae. buZZata L. (Liliaceae) ， with a comparison to the Asian Sugerokia. Sugerokia. Helionopsis. Ann. Mus. Bot. Lugd.- Heloniopsis orientalis (Thunb.) C. Tanaka. Ann. Batav.3: Batav.3: 144-146. Carnegie Mus. 49: 153-160.

田中教之: Helonias ，Ypsilandra ，Heloniopsis の系統 と分類 III. 分類学的検討 これまでに得られた観察結果 (Tanaka 1997a-e) マとツクシショウジョウパカマの2群(あるいはヤ によると，表題の3属は属のランクで区別するほど マシマショウジョウパカマも区別して3群)に区別 の顕著な差異を持たないと判断される.従って，こ されることが多いが，私はこれらを区別せず一種 れら 3属を 1属に統合することを試みた.Yj フsilandra として見なした.これらは明確な差異をもって互 属と Heloniopsis 属は Helonias 属に統合される.検 いに区別されるというものではない.かつては，九 討の結果， Helonias 属には9種が認められた.この 州のものも含めて，上記の分布域に生育する白色 内の 1種は北米東部に，他の8種はアジア東部に分 系統のものはシロバナショウジヨウパカマと呼ば 布している.本稿には，学名の新組合せによる新 ばれていた(牧野 1891 ，Matsumura 1905 ， Koidzumi 名が8 含まれている . Heloniopsis 属の和名である 1930). 従って，この種 (Helonias breviscapa) の和 ショウジョウパカマ属という名称は， 3属を統合し 名としてはシロパナショウジョウバカマがふさわ た場合のHelonias 属の和名としても使えると思う. しいと考える.しかし，この和名は，紅紫色の花を 日本産のショウジョウパカマ属 (Helonias) で， つける(普通の)ショウジョウパカマ (H. orientalis) 花は白色または少し帯紅紫色，葉はやや薄質で縁 の白色花をつけるものとこれまでしばしば混同さ がしばしば細かく波状となり，やや黄緑色のもの れてきた.この混同は和名だけに留まるものでは が，関東南部から本州西部，四国，九州、|に分布して なかった.名称、に由来する混乱を避ける意味から， いる.この種類は今日シロパナショウジョウバカ H. breviscapa の和名として新にコチョウショウ April1998 April1998 Journal of Japanese Botany Vo l. 73 No. 2 115

ジョウパカマ(胡蝶狸々袴)の名を与えることに 種のうち，日本に分布するのは次の 4 種である: する[この名は Thunberg (1784) がかつて Scilla Helonias leucantha オオシロショウジョウバカマ， japonic α( = H. breviscap α) の和名として採録した H. kawanoi コショウジヨウノてカマ ， H. orientalis ‘Ko り0' に因む] .ショウジョウバカマの白花品に ショウジョウノ T カマ， H. breviscapa コチョウショウ 対してはシロショウジョウバカマ(槍山 1939) の ジョウノ T カマ. 和名がある.ただし，本稿では花色の変異体を分 (帝京大学文学部教育学科) 類学的に区別することはしなかった.アジア産 8

第 72 巻 6 号正誤 Errata ofVo l. 72 No. 6

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329 329 ↓ 14 to paraphyletic to be paraphyletic 329 329 is be presumed is presumed 335 335 ↓ 16 shinnka shinka