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J. Jpn. Bot. 73 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 tepals ， while those of the two species plants. 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 taxonomy 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).
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