植物研究雑誌 J. J. Jpn. Bo t. 79: 79: 186-195 (2004)

New Combinations in (Compositae-Anthemideae) of of Asia with a of List J apanese Species

Hiroyoshi Hiroyoshi OHASHl a and Koji YONEKURA b

aBotanical aBotanical Garden ,Graduate School of Science ,Tohoku University ,Sendai , 980-0862 JAPAN; bThe bThe M t. Hakkoda Botanical Laboratory ,Graduate School of Science ,Tohoku University ,Aomori ,030 ・0111 JAPAN

(Received (Received on Janu 紅 y 10 ,2004)

New combinations or new names for two sections , 28 species and the infraspecific taxa taxa are proposed under the conserved generic name Ch η santhemum L. The genus is cir- cumscribed cumscribed here almost as identical with Dendranthema of Kitamura (1 978) including Ajα nia ,Arctanthemum , Dendranthema and Ph αeostigm αin the sense of Bremer and Humphries Humphries (1993). A list of the Japanese species of Chrysanthemum is provided.

Key words: Asia ,Chrysanthemum ,Dendranthema ,new combinations ,new names.

The generic name Chrysanthemum L. is of Dendranthema as (A. pacifica conserved conserved with a conserved type C. indicum (N akai) K. Bremer & Humphries and A. L. L. (Brummitt 1998 , Greuter et al. 2000). The shiwogiku (Kitam.) K. Bremer & Humph- generic generic name Dendranthema Des Mou l. be- ries). However , morphological distinction

C 創 ne a synonym of Chrysanthemum ,al 国 and genetic isolation between them are con- though the name had been recently accepted sidered to be insufficient to recognize them in in E. Asia (Ki tamura 1978 ,1981 ,Shih and as different genera. Many hybrids 紅 e known Fu 1983 ,Barkalov 1992 ,Koyama 1995 , between Ajania and Chrysanthemum in Peng et al. 1998) and also in a generic mono- Japan (Table 1). The chromosomes of Ajania graph by Bremer and Humphries (1 993). and Chrysanthemum 訂 e similar to each other Bremer and Humphries (1993) discussed (Nakata et al. 1987). Distinctness of Ajania relations relations between Ch ηsanthemum (as from Chrysanthemum was already suspected

Dendranthema) and its satellite genera in 町 by Tzvelev based on extensive examination cluding cluding Ajania , Arctanthemum and of Chinese material (see Bremer and

Phaeostigma , which were included in Humphries 1993). We reg 紅 d Ajania as a

Dendr α, nthema by Ki tamura (1978). The section of Ch ηsanthemum. This was already genus i全jania Poljak. (Poljakov 1955) based proposed by Kitamura (1 978) under on A. pallasian α(Turcz. ex Besser) Poljak. Dendranthem α. Ph α eostigma is sep 紅 ated has has been accep 旬d by many authors ,espe- fromAjani αby Muldashev (1981 ,1983) , but cially cially in Russia and China , as distinct from Shih and Fu (1983) included it in ベjania. We Chrysanthemum or Dendranthem α(Tzvelev in c1 ude Ph αeostigm αinto Chrysanthemum. 1961 , Shih 1979 ,1994 ,Muldashev 1983 , Chrysanthemum arcticum L. (including Shih and Fu 1983 ,Barkalov 1992 ,Bremer Dendranthema kurilense Tzvelev) is often and Humphries 1993). Bremer and regarded as comprising the genus Humphries (1993) kept both genera separate , Arctanthemum Tzvelev (1985) based on its and treated two disciform J apanese species distinc t1 y ribbed cypselas without myxogenic

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Table Table 1. Intersectional hybrids between Chrysanthemum sec t. Chrysanthernurn (= Dendranthema) and sec t. Ajania (= Ajania) found in J apan

Hybrid (sec t. Chrysanthernurn x sec t. Ajania)

C. C. makinoi x C. paU αsian ωη = Chrysanthemum xkonoanum Makino トガクシギク 仁 morifolium x C. pacificum = Chrysanthemum xmarginatum (Miq.) N. E. Br. ハナイソギク C. C. indicum x C. kinokuniense = Ch ηsanthemum xogawae Ki tarn. ヒノミサキギク C. C. makinoi x C. rupestre = Chrysanthemum xtodaiense H. Ohashi & Y onek. * リュウノウイワインチン C. C. makinoi x C. kinokuniense 仁 morifolium x 仁 kinokuniense C. C. makinoi x C. shiwogiku C. C. morifolium x C. shiwogiku

*New narnes see text

cells cells (Bremer and Humphries 1993). This Chrysanthemum sec t. Arctanthemum species species is ,however , very similar to C. (Tzvelev) H. Ohashi & Yone k., comb. nov. yezoense yezoense Maek. which is often reg 紅 ded as a Dendranthema sec t. Arctanthemum subspecies subspecies of C. arcticum as subsp. Tzvelev , F l. URSS 26: 385 (196 1). maekawanum Ki tam. More precise analyses Arctanthemum (Tzvelev) Tzvelev in on achene characters are needed for C. Novi t. Sys t. Pl. Vasc. 22: 274 (1985); arcticum , but we consider Arctantherum Barkalov in Kh arkev. , P l. Vasc. Orien t. Extr. should should be treated as a section of Chrysanthe- Sovie t. 6: 109 (1 992); Bremer & Humphries mum. in Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): 114 114 (1993). Type species: A. arcticum (L.) New combinations for sections in Tzvelev. Chrysanthemum Chrysanthemum sec t. Aj ania (Poljak.) New combinations for species and H. Ohashi & Yonek. ,comb. nov. infraspecific taxa in alphabetical order of Ajania Ajania PO りak. in No t. Sys t. Herb. Ins t. the specific epithet Bo t. Acad. Sci. URSS 17: 419 (1955) , p. p.; Chrysanthemum abolinii (Kovalevs k.) Tzvelev , F l. URSS 26: 398 (1961); C. Shih H. Ohashi & Yonek. ,comb. nov. & G. X. Fu , F l. Reipub l. Pop. Sin. 76 (1): 102 Ajania abolinii Kovalevs k. in Novi t. Sys t. (1 983); Barkalov in Kh arkev. , P l. Vasc. Pl. Vasc. 23: 247 (1 986). Orien t. Extr. Sovie t. 6: 162 (1 992); Bremer Distr.: C. Asia. & Humphries in Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): 115 (1 993). Type species: A. Chrysanthemum alabasium (H. C. Fu) pallasiana pallasiana (Fischer ex Besser) Polja k. H. Ohashi & Yone k., comb. nov. Dendranthema sec t. Ajania (Poljak.) Ajania alabasia H. C. Fu , F l. Intramong. Kitam. Kitam. in Acta Phytotax. Geobo t. 29: 168 6: 325 ,t. 34 (1982) , ed. 2, 4: 585 (1993). (1978). (1978). Distr.: China (Inner Mongolia). Phaeostigm αMuldashev in Bo t. Zhum. SSSR 66: 586 (1 981); Bremer & Humphries Chrysanthemum arcticum L. , Sp. Pl. 2: in in Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): 889 (1 753). 115 115 (1 993). subsp. yezoense (Mae k.) H. Ohashi & Yone k., sta t. nov. 188 188 植物研究雑誌第79 巻第3号 平成16 年6月

Chrysanthemum arcticum var. yezoense Chrysanthemum arcticum is va r. yezoense Mae k. in Trans. Sapporo Na 1. His 1. Soc. 8: Mae k. (1921) , but the name is confusable 14 ,figs. 1-4 (1921); Kitam. in Mem. Coll. with C. yezoense Mae k. (1921) (= C. Sci. Sci. Kyoto Imp. Univ. ser. B , 15(3) arcticum subsp. maekawanum Kitam.) which [Compos. [Compos. Jap. 2]: 347 (1 940) ,non C. occurs simpatrically with the former in yezoense yezoense Mae k. (1 921). N emuro Peninsula (Nishikawa and Dendranthema kurilense Tzvelev , Fl. Kobayashi 1989). URSS 26: 385 (1 961) ,nom. nov. for C. arcticum arcticum v紅 . yezoense Mae k. Chrysanthemum brachyanthum (C. Arctanthemum kurilense (Tzvelev) Shih) H. Ohashi & Yone k., comb. nov. Tzvelev Tzvelev in Novi 1. Sys 1. Pl. Vasc. 22: 274 Ajania brachyantha C. Shih in Acta (1 985); Bremer & Humphries in Bul l. Na 1. Phytotax. Sin. 17: 114 (1 979) , in C. Y. Wu , His 1. Mus. Lond. (Bo 1.) 23(2): 114 (1 993). Fl. Xizang. 4: 736 (1 985); C. Shih & G. X. Arctanthemum α ucticum (L.) Tzvelev Fu , F l. Reipub l. Pop. Sin. 76 (1): 110 ,1. 17 , subsp. subsp. kurilense (Tzvelev) Tzvelev , F l. Arc 1. fig. 2 (1 983); Bremer & Humphries in Bul l. URSS 10: 115 (1 987); Barkalov in Kh arkev. , Na 1. His 1. Mus. Lond. (Bo t.) 23(2): 115 l. Pl. Vasc. Orien 1. Extr. Sovie 1. 6: 110 , in nota (1 993). (1 992). Distr.: China (S. Tibet (Xizang)). ? Chrysanthemum arcticum var. kurilense Kudo , F l. Paramushir.: 167 (1 922). Chrysanthemum delavayanum H. Ohashi Plants of Chrysanthemum arcticum in & Yonek. ,nom. nov. Nemuro Peninsula of eastem Hokkaido , Ajania serice αC. Shih in Bull. Bo t. Lab. Kurile Kurile Islands and South Sakhalin 紅 e ch 佐 N. ーE. Fo r. Ins t. 6: 14 (1 980); C. Shih & G. acterized acterized by the smaller habit with usually X. Fu , F l. Reipub l. Pop. Sin. 76 (1): 118 ,t. simple simple stem and more striated ligules of the 19 ,figs. 1 & 2 (1 983); Bremer & Humphries ray-florets ray-florets than those of the typical form. in Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): Accordingly , they have been distinguished 115 (1 993) ,non Chrysanthemum sericeum from the typical form as a different taxon at Adam (1 805-1806) , nec Hoffm. & Link the the rank of variety (Maekawa 1921 , (1820). Kitamura Kitamura 1940) , subspecies (Tzvelev 1987 , Dist r.: China (NW. Yunnan). Barkalov Barkalov 1992) or a distinct species (Tzvelev The epithet of this species under 1961 , 1985). Ki tamura (1 967 ,1978 , 1981) Chrysanthemum is changed to avoid treated treated this taxon as a synonym of C. homonymy and renamed after J. M. Delavay , arcticum , since he regarded plants from collector of the type specimen. Shiretoko Shiretoko Peninsula of eastem Hokkaido are indistinguishable indistinguishable from the typical C. Chrysanthemum dichrum (C. Shih) H. arcticum , but we consider that the taxon is Ohashi & Yone k., comb. nov. distinct distinct because its diagnostic characters are Dendranthema dichrum C. Shih in Bull. useful useful and its distribution is separated from Bo t. Lab. N.-E. For. Ins t. 6: 8 (1980); C. the the typical form. We recognize the taxon as Shih & G. X. Fu , F l. Reipub l. Pop. Sin. 76 a subspecies of C.α rcticum. Unfortunately , (1): 47 (1 983); Bremer & Humphries in Bull. the the available epithet kurilense at the subspe- Na t. His t. Mus. Lond. (Bo t.) 23(2): 114 cies cies level can not be used under (1993). Chrysanthemum due to the existence of C. Distr.: N. China (Hebei). α rcticum var. kurilense Kudo (1 922). The This Chinese species was presumed as an earliest earliest epithet for the taxon under intersectional hybrid in the original descrip- June June 2004 Joumal of Japanese Botany Vo 1. 79 No. 3 189 tion , but the latter authors treate it as a natu- Herb. Ins t. Bo t. Acad. Sc i. URSS 17: 424 ral ral species. (1 955); C. Shih & G. X. Fu , F l. Reipub l. Pop. Pop. Sin. 76 (1): 109 , in nota (1 983); Bremer Chrysanthemum fastigiatum (C. Wink l.) & Humphries in Bull. N a t. His t. Mus. Lond. H. Ohashi & Yonek. ,comb. nov. (Bo t.) 23(2): 115 (1 993). Artemisia Artemisia fastigiata C. Wink l. in Acta Dist r.: China (N. Yunnan). Hort. Hort. Petrop. 11: 373 (1 891). Ajania Ajania fastigiata (C. Wink l.) Poljak. in Chrysanthemum khartense (Dunn) H. No t. Sys t. Herb. Bo t. Ins t. Acad. URSS 17: Ohashi & Yone k., comb. nov. 428 (1 955); Tzvelev , F l. URSS 26: 405 Tanacetum khartense Dunn in Kew Bul l. (1961); (1961); C. Shih & G. X. Fu , F l. Reipub l. 1922: 150 (1 922). Pop. Pop. Sin. 76 (1): 125 ,t. 20 ,fig. 3 (1 983); Ajania khartensis (Dunn) C. Shih in Acta Bremer & Humphries in Bull. Na t. His t. Phytotax. Sin. 17: 115 (1979) , in C. Y. Wu , Mus. Lond. (Bo t.) 23(2): 115 (1 993). Fl. Xizang. 4: 737 (1 985); C. Shih & G. X. Distr.: Distr.: C. Asia , S. Siberia , China Fu , F l. Reipub l. Pop. Sin. 76 (1): 113 ,t. 17 , (Xinjiang) (Xinjiang) and Mongolia. fig. 3 & t. 1, fig. 9 (1 983); Bremer & Humphries in Bull. Na t. His t. Mus. Lond. Chrysanthemum gracile (Hook. f. & (Bo t.) 23(2): 115 (1993). Thomson) H. Ohashi & Yone k., comb. nov. Distr.: Pamir , the Himalayas and SW. Tanacetum gracile Hoo k. f. & Thomson in China. Hook .f., Fl. Bri t. India 3: 318 (1 881). ベjania gracilis (Hoo k. f. & Thomson) Chrysanthemum kinokuniense (Shimo t. Poljak. Poljak. ex Tzvelev , Fl. URSS 26: 407 & Ki tam.) H. Ohashi & Yone k., sta t. nov. (1 961); C. Shih & G. X. Fu , F l. Reipub l. Chrysanthemum shiwogiku Ki tam. var. Pop. Pop. Sin. 76 (1): 124 , in nota (1983); Bremer kinokuniense Shimo t. & Kitam. in Kitam. in & Humphries in Bull. Na t. His t. Mus. Lond. Acta Phytotax. Geobo t. 4: 73 (1 935) & 22 (Bo t.) 23(2): 115 (1 993). (4- 6): 120 (1 967). Dist r.: C. Asia and W. Himalaya. A of Chrysanthemum sec t. Ajania growing growing in coastal region of Ki i Peninsula in Chrysanthemum grubovii (Muldashev) southwestern Honshu has been regarded H. H. Ohashi & Yone k., comb. nov. taxonomically as a variety of C. shiwogiku Ajania Ajania grubovii Muldashev in Bo t. Zhurn. distributed in Shikoku (Kitamura 1935 , SSSR 67: 1529 (1 982); Bremer & 1940 ,1967 ,1978 , 1981). Shimotomai et al. Humphries in Bull. N a t. His t. Mus. Lond. (1 968) revealed it as being a hybrid deriva- (Bo t.) 23(2): 115 (1 993). tive between octoploid C. shiwogiku and Distr.: Distr.: Mongolia. decaploid C. pac がcum distributed in Kanto and and Tokai regions. In Ki i Peninsula ,how- Chrysanthemum hypoleucum (Y. Ling ever , the plants occurring in east of Cape ex C. Shih) H. Ohashi & Yone k., comb. nov. Shionomisaki , the tip of Ki i Peninsula , are Ajania Ajania hypoleuca Y. Ling ex C. Shih in principally decaploid , whereas those grow-

Acta Phytotax. Sin. 32: 366 (1 994). ing in west of the cape 紅 e usually octoploid , Distr.: Distr.: China. although many aneuploid plants were ob- served served especially at and around the tip Chrysanthemum junnanicum (Poljak.) (Shimotomai et al. 1968). Nakata et al. H. H. Ohashi & Yone k., comb. nov. (1 987) regarded the octoploid plants as C. Ajania Ajania junnanic αPoljak. in No t. Sys t. shiwogiku and decaploid as C. pacificum ex- 190 190 植物研究雑誌第79 巻第3号 平成16 年6月 clusively clusively based on their ploidy differences. This variety is dubious in morphological Their Their recognition ,however ,makes it diffi- distinctness from var. orn αtum in Satsuma cult cult to discriminate the two species from Peninsula. Due to its remote geographical each each other , because their morphological dis- separation we tentatively recognize it as it tinctions tinctions are often continuous among the was described (Hotta et a1. 1996). plants plants in Kii Peninsula. We prefer recogniz- ing ing the plants of Kii Peninsula as an inde- Chrysanthemum przewalskii (Poljak.) pendent pendent species derived from a hybrid origin H. Ohashi & Yonek. ,comb. nov. distinct distinct from the parents. Ajania przewalskii Poljak. in No t. Sys t. Herb. Herb. Ins t. Bo t. Acad. Sci. URSS 17: 422 Chrysanthemum kokanicum (Kr asch.) (1955); C. Shih & G. X. Fu , F 1. Reipub 1. H. Ohashi & Yonek. ,comb. nov. Pop. Sin. 76 (1): 109 (1983); Bremer & Tanacetum kokanicum Kr asch. in No t. Humphries in Bull. Na t. His t. Mus. Lond. Sys t. Herb. Hor t. Petrop. 4: 7 (1 923). (Bo t.) 23(2): 115 (1 993). Ajania Ajania kokanic α(Kr asch.) Tzvelev , F l. Dist r.: China (Sichuan ,Qinghai , E. Gansu URSS 26: 408 (1 961); Bremer & Humphries and Ningxia). in in Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): 115 115 (1 993). Chrysanthemum purpureiflorum H. Distr.: Distr.: C. Asia. Ohashi & Yonek. ,nom. nov. Ajania Ajania purpurea C. Shih in Acta Phytotax. Chrysanthemum korovinii (Kovalevs k.) Sin. 17(2): 115 (1 979) , in C. Y. Wu , F 1. H. H. Ohashi & Yonek. ,comb. nov. Xizang. 4: 738 (1985); C. Shih & G. X. Fu , Ajania Ajania koro νinii Kovalevs k. in N ovi t. F1. Reipub 1. Pop. Sin. 76 (1): 115 (1 983); Sys t. P1. Vasc. 23: 246 (1986). Bremer & Humphries in Bull. Na t. His t. Distr.: Distr.: C. Asia. Mus. Lond. (Bo t.) 23(2): 115 (1 993) ,non Chrysanthemum purpureum Pers. (1807). Chrysanthemum nitidum (C. Shih) H. Distr.: China (Tibet (Xizang)). Ohashi Ohashi & Yone k., comb. nov.

Ajania Ajania nitid αC., Shih in Bull. Bo t. Lab. Chrysanthemum ramosum (C. C. N.-E. N.-E. For. Ins t. 6: 15 (1 980); C. Shih & G. Chang) H. Ohashi & Yone k., comb. nov. X. X. Fu , F 1. Reipub 1. Pop. Sin. 76 (1): 123 ,t. Chrysanthemum variifolium C. C. Chang 20 ,fig. 2 (1 983); Bremer & Humphries in var. ramosum C. C. Chang in Sinensia 5: 163 Bull. Bull. Na t. His t. Mus. Lond. (Bo t.) 23(2): 115 (1 934). (1 993). Aj, α nia ramos α(C. C. Chang) C. Shih in Distr.: Distr.: China (SW. Sichuan). Acta Phytotax. Sin. 17: 114 (1979) , in C. Y. Wu , F l. Xizang. 4: 735 (1 985); C. Shih & G. Chrysanthemum ornatum Hems 1. in X. Fu , F 1. Reipub 1. Pop. Sin. 76 (1): 108 Curtis's Curtis's Bo t. Mag. 130: t. 7965 (1904). (1983). var. var. tokarense (M. Hotta & Y. Hirai) H. Phaeostigma variifolium (C. C. Chang) Ohashi Ohashi & Yonek. ,comb. nov. Muldashev var. ramosum (C. C. Chang) Dendranthema ornatum (Hems 1.) Ki tam. Muldashev in Bo t. ZhUffi. SSSR 66: 587 V 紅 . tokarense M. Hotta & Y. Hirai in Acta (1981). Phytotax. Phytotax. Geobo t. 47: 103 (1 996) ,“ tokaren- Distr.: China (E. Tibet (Xizang) and W. sis" sis" . Sichuan). Distr.: Distr.: Japan: S. Kyushu (l s1. Yaku-shima , 1. 1s 1. Kuchinoerabu サima and Tokara Islands). Chrysanthemum rhombifolium (Y. Ling June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 191

& C. Shih) H. Ohashi & Yone k., comb. nov. [Iconog r. Cormoph. Sin. 4: 510 ,fig. 6433 Dendranthema rhombifolium Y. Ling & (1 975) , comb. nud.] ex Ki tam. in Acta C. C. Shih in Bull. Bo t. Lab. N.-E. For. Ins t. 6: Phytotax. Geobo t. 29: 167 (1978); Kitam. in 2 (1 980); C. Shih & G. X. Fu , F 1. Reipub 1. Satake & a 1., Wild Flow. Jap. Herb. P1. 3: Pop. Pop. Sin. 76 (1): 31 ,t. 5, fig. 1 (1 983); 167 (1 981); Bremer & Humphries in Bu11. Bremer & Humphries in Bull. N a t. His t. Na t. His t. Mus. Lond. (Bo t.) 23(2): 114 Mus. Lond. (Bo t.) 23(2): 114 (1 993). (1993); H. Koyama in K. Iwats. & a1., F1. Distr.: Distr.: China (Chongqing). Jap. IIIb: 94 (1 995). Dendranthema lavandulifolium (Fisch. ex Chrysanthemum roborowskii (Mulda- Trautv.) Kitam. va r. seticuspe (Maxim.) C. shev) shev) H. Ohashi & Yone k., comb. nov. Shih in Bu11. Bo t. Lab. N.-E. For. Ins t. 6: 6 Ajania Ajania roborowskii Muldashev in Bo t. (1 980) , p. p.; C. Shih & G. X. Fu , F 1. Zhum. SSSR 67: 1528 (1 982); Bremer & Reipub 1. Pop. Sin. 76 (1): 42 (1 983) , p. p. Humphries in Bu11. Na t. His t. Mus. Lond. Dendranthema seticuspe (Maxim.) Ki tam. (Bo t.) 23(2): 115 (1 993). f. boreale (Makino) Ki tam. in Acta Phytotax. Distr.: Distr.: SW. China. Geobo t. 38: 381 (1 987); Ki tam. & a 1., Co 1. var. var. tsinghaicum (Muldashev) H. Ohashi 11 1. Herb. P1. Jap. (63th printing) 1: 61 ,t. 20 , & Yonek. ,comb. nov. no. 144 (1 987). Ajania Ajania roborowskii var. tsinghaicum Dist r.: Japan (Honshu and N. Kyushu) , Muldashev in Bo t. Zhum. SSSR 67: 1529 Korea and NW. China.

(1 982). Chrysanthemum seticuspe was rarely cul 同 Dist r.: China (Qinghai). tivated in the Edo era , but considered as ex- tinct tinct in the Meiji period (Kitamura 1940 , Chrysanthemum seticuspe (Maxim.) 1967). Recently it was rediscovered in the Hand.-Mazz. ,Symb. Sin. 7(4): 1111 (1 936). Imperial Palace in Tokyo (Ki tamura 1987). 五boreale (Makino) H. Ohashi & Yone k., Kitamura (1987) considered it as merely a comb. nov. cultivated form of wild C. boreale distrib- Chrysanthemum indicum L. var. boreale uted in Japan ,Korea and China and made the Makino in Bo t. Mag. Tokyo 16: 89 (1902). combination Dendranthema seticuspe f. Chrysanthemum bore α le (Makino) Maki- boreale. We agree with Ki tamura's pinion. no in Bo t. Mag. Tokyo 23: 20 (1909); This species has the same chromosome num- Matsum. , Index P1. Jap. 2(2): 636 (1912); ber and similar karyotypes with Chinese C.

Kitam. Kitam. in J. Jpn. Bo t. 13: 167 (1 937) , in lavandulifolium Turcz. ex Trautv. As the dis 帽 Mem. Co11. Sci. Kyoto Imp. Univ. se r. B ,15 tinction between the species are often criti- (3) (3) [Compos. Jap. 2]: 365 ,t. 1 ,fig. 2 (1940) , cal , further studies are needed on C. in in Acta Phytotax. Geobo t. 22( 4- 6): 129 seticuspe and C. lavandulifolium. (1967); (1967); H. H 紅 a,Enum. Spermatoph. Jap. 2: 157 157 (1952); Ohwi , F 1. Jap.: 1187 (1 953); Chrysanthemum shihchuanum H. Ohashi Kitam. Kitam. & a1., Co 1. 111. Herb. P1. Jap. 1: 61 ,t. & Yonek. ,nom. nov.

20 , no. 144 (1 957); Ki tag. ,Neo ・Lineam. F1. Ajania latifolia C. Shih in Bu11. Bo t. Lab.

Mansh.: Mansh.: 633 (1979); Ohwi & Ki tag. ,New F1. N. 闇 E. For. Ins t. 6: 12 (1 980); C. Shih & G. Jap.: Jap.: 1500 (1982). X. Fu , F 1. Reipub 1. Pop. Sin. 76 (1): 106 Chrusanthemum seticuspe v紅 . bore αle (1983); Bremer & Humphries in Bull. Na t. (M 北ino) Hand.-Mazz. ,Symb. Sin. 7(4): His t. Mus. Lond. (Bo t.) 23(2): 115 (1 993) , 1111 1111 (1 936). non Chrysanthemum lat ゆlium (DC.) Dendranthema boreale (M 紘 ino) Y. Ling Baksay (1 957). 192 192 植物研究雑誌第79 巻第3号 平成16 年6月

Distr.: Distr.: China (SW. Sichuan). subsp.α rcticum (diploid) アキノコハマ The specific epithet under Chrysanthe- ギク mum is renamed after Shih Chu , author of subsp. yezoense (Maek.) H. Ohashi. & the the basionym , to avoid homonymy. Y one k. (diploid) チシマコハマギク

Chrysanthemum xtodaiense H. Ohashi II. Sect. Chrysanthemum & Yone k., nom. nov. 2. Chrysanthemum crassum (Ki t副 n.) Chrysanthemum makinoi Matsum. & Kitam. (decaploid) オオシマノジギク N akai x C. rupestre Matsum. & Koidz.: 3. Chrysanthemum indicum L. Kitam. Kitam. in Acta Phytotax. Geobo t. 22(4-6): var. indicum (tetraploid and hexaploid) 113 113 (1 967) ,cum descr. latin. シマカンギク Type: Type: Japan: Honshu , Nagano Pref. V 訂 . albescens Makino シロパナハマカ (Prov. (Prov. Shinano) , Kamiinagun , Hasemura , ンギク Todai ,1400 m , in locis calcareis. T. Shimizu var. iyoense (te 住aploid) イヨアブラギ 5501 , 21 Sep. 1959 (KYO-holotype). ク This This is. an intersectional hybrid between var. maruyamanum Kitam. (tetraploid) sec t. Chrysanthemum and sec t. Ajania. オッタチカンギク var. var. tsurugisanense Ki tam. (tetraploid) Chrysanthemum trilobatum (Poljak.) H. ツルギカンギク Ohashi & Yone k., comb. nov. 4. Chrysanthemum japonense Nakai Ajania Ajania trilobata Poljak. , F l. URSS 26: 880 V 征 .j αrponense (hexaploid) ノジギク (1961); (1961); C. Shih & G. X. Fu , F l. Reipub l. var. ashizuriense Ki tam. (hexaploid) ア Pop. Pop. Sin. 76 (1): 116 ,t. 18 ,fig. 3 (1983); シズリノジギク Bremer & Humphries in Bull. Na t. His t. 5. Chrysanthemum makinoi Matsum. & Mus. Lond. (Bo t.) 23(2): 115 (1 993). Nakai (diploid) リユウノウギク Dist r.: NW. China (Xinjiang). 6. Chrysanthemum okiense Ki tam. (tetraploid) オキノアブラギク Chrysanthemum tripinnatisectum (Y. N akata et al. (1987) doubted its taxonomic Ling & C. Shih) H. Ohashi & Yone k., comb. status because it has the same chromosome nov. nov. number as C. indicum. Ajania Ajania tripinnatisecta Y. Ling & C. Shih 7. Ch η lsanthemum orn αtum Hems l. in in C.Shih in Bull. Bo t. Lab. N.-E. For. Ins t. V 紅 . omatum (octoploid) サツマノギク 6: 6: 14 (1980); C. Shih & G. X. Fu , F l. var. tokarense (M. Hotta & Y. Hirai) H. Reipub l. Pop. Sin. 76( 1): 118 ,t. 19 ,fig. 5 Ohashi & Y one k. (octoploid) トカラノギク (1 983); Bremer & Humphries in Bull. Na t. 8. Chrysanthemum seticuspe (Maxim.) His t. Mus. Lond. (Bo t.) 23(2): 115 (1993). Hand.-Mazz. Distr.: Distr.: China (NW. Sichuan). f. seticuspe (only known in cultivation) カモメギク A List of the Japanese species of f. boreale (Makino) H. Ohashi & Ch η'santhemum Y onek. (diploid) キクタニギク

All All the J apanese taxa 紅 e listed here with 9. Chrysanthemum wak α, saense Shimo t. ex the the co 町 ect name , ploidy in parenthesis ,and Ki tam. (tetraploid; derived from C. makinoi J apanese name. x C. seticuspe f. boreale or C. indicum) ワ カサハマギク 1. 1. Sect. Arctantherum 10. Chrysanthemum weyrichii (Maxim.) 1. 1. Chrysanthemum arcticum L. Miyabe & T. Miyake (octoploid; see Nakata June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 193 et et al. 1987) ピレオギク Hybrids in sec t. Chrysanthemum This This species was synonymised with C. 15. Chrysanthemum x aphrodite Ki tam. (= zawadskii zawadskii (Ki tamura 1967 ,1981) , but is dis- C. indicum x C. morifolium) サンインギク tinct tinct from the cytogenetical and mo 中hologi- Ki tamura (1 967) regarded this plant as a cal cal points of view. distinct hexaploid species closely related to 11. 11. Chrysanthemum yezoense Mae k. C. indicum , but N akata et al. (1987) and (decaploid) コハマギク Nakata and Takeuchi (1998) considered the This This plant has been frequently regarded as type specimen of this species is a hybrid de- a subspecies of C. arcticum in sec t. rivative between hexaploid C. indicum and Ar ctantherum (Kitamura 1935 ,1940 ,1967 , cultivated C. morifolium. The hybridity of 1981 ,Koyama 1995) , but is quite distinct as this taxon is supported from morphological revealed revealed by Nishikawa and Kobayashi point of view.

(1 989). It is rather ne 征 er to C. zawadskii and 16. Chrysanthemum x cuneifolium Ki tam. C. C. weyrichii in achene characters (Tzvelev (= C. indicum x C. yoshinaganthum) ワジキ 1961 ,Hind 1988). ギク 12. 12. Chrysanthemum yoshinaganthum 17. Chrysanthemum x leucanthum (Maki- Makino ex Ki tam. (tetraploid) ナカガワノギ no) Makino (= C. makinoi x C. seticuspe f. ク boreale) シロバナアブラギク 13. 13. Chrysanthemum zawadskii Herbich 18. Chrysanthemum miyatojimense Ki tam. (hexaploid (hexaploid and octoploid) イワギク (=仁 morifolium x 仁 yezoense) ミヤトジマ Hexaploid Hexaploid plants sporadically distributed ギク in in Honshu , Shikoku and Kyushu 紅 'e app 紅- 19. Chrysanthemum x shimotomaii M 紘 i- ently ently relictal in Japan. Japanese hexaploids no (=仁 indicum x C. japonense) ニジガハ dominantly dominantly have dissected leaves with n訂- マギク rower rower lobes , but in some localities plants with with wider lobes of leaves are growing. 11 1. Sect. Ajania These These plants were often referred to the vari- 20. Chrysanthemum kinokuniense ety ety of continental Asia , va r. latilobum (Shimo t. & Kitam.) H. Ohashi & Yone k. Ki tam. , but are here considered as lying (octoploid and decaploid; derived from C. within within the range of C. zawadskii as stated by pacificum x C. shiwogiku) キノタニシオギ Ki tamura (1 967 , 1981). ク The octoploid plants recently found in 21. Chrysanthemum pacificum N akai Kitakami Kitakami Mountains in N. Honshu (N akata (decaploid) イソギク and Kumagai 1999) seem specifically dis- 22. Chrysanthemum pallasianum '(Fisch. tinct tinct not only from C. zawadskii but also ex Besser) Kom. (hexaploid) オオイワイン from C. weyrichii. These plants were sus- チン pected pected as hybrid derivatives between 23. Chrysanthemum rupestre Matsum. & hexaploid hexaploid C. zawadskii and decaploid C. Koidz. (diploid) イワインチン yezoense yezoense (Nakata and Kumagai 1999). 24. Chrysanthemum shiwogiku Kitam. (octoploid) シオギク Cultivated Cultivated species 14. 14. Chrysanthemum morifolium Rama t. IV. Intersectional hybrids between sect. (hexaploid (hexaploid probably derived from a hybrid Chrysanthemum and sect. Ajania C. C. indicum x C. zawadskii) キク 25. Chrysanthemum xkonoanum M 紘 ino (C. (C. makinoi x C. pallasi αnum) トガクシギク 26. 26. Chrysanthemum xmarginatum (Miq.) 194 194 植物研究雑誌第79 巻第3号 平成16 年6月

N. E. B r. (仁 morifolium x C. pacificum) ハ Ohwi J. ,Ki tamura S. , Watari S. and Torninari T. ナイソギク (eds.) Wild Flowers of Japan , Herb. Pl. 3: 156-235. 27. 27. Chrysanthemum xog αwae Ki tam. (C. Heibonsha Ltd. ,Tokyo (in Japanese). 一一一 1987. On three species of Sympetalae. Acta indicum indicum x C. kinokuniense) ヒノミサキギク Phytotax. Geobo t. 38: 380-382 (in Japanese). 28. 28. Chrysanthemum xtodaiense H. Ohashi Koyama H. 1995. Tribe 5. Anthemideae. In: Iwatsuki & Yonek. (C. makinoi x C. rupestre) リユ K., Yamazaki T. , Boufford D. E. and Ohba H. ウノウイワインチン (eds.) Fl ora of Japan 3b: 77-97. Kodansha Lt d. , 29. 29. Chrysanthemum makinoi x C. kino- Tokyo. Maekawa T. 192 1. On the wild Chrysanthemum of kuniense kuniense north north Japan. Trans. Sapporo Na t. His t. Soc. 8: 9- 30. 30. Chrysanthemum morifolium x C. kino- 18. kuniense kuniense Muldashev A. A. 198 1. A new genus Phaeostigma

31. 31. Chrysanthemum makinoi x C. shiwo ・ () from the East Asia. Bo t. Zhum. SSSR giku giku 66(4): 58 4- 588 (in Russian). 一一 1983. A critical review of the genus Ajania (As 32. 32. Chrysanthemum morifolium x 仁 teraceae-Anthemideae). teraceae-Anthemideae). Bo t. Zhum. SSSR 68(2): shiwogiku shiwogiku 207-214 (in Russian). Nakata Nakata M and Takeuchi M. 1998. Variations in References References Dendranthema indicum var. aphrodite population Barkalov Barkalov V. Y. 1992. Asteraceae (Compositae). In: in Oozakai ,Himi City ,Toyama Pre f. Bull. Bo t. Charkevicz Charkevicz (Kh arkevicz) , S. S. (ed.) Plantae Gard. Toyama no. 3: 1-16 (in Japanese). Vasculares Vasculares Orientis Extremi Sovietici 6: 427 pp. 一一一 and Kumagai A. 1999. Octoploid cytotype of Nauka ,S t. -Petersburg (in Russian). Dendranthema zawadskii (Asteraceae) and its im- plications plications in evolutional history. Bul l. Bo t. Gard. Bremer Bremer K. and Humphries C. J. 1993. Generic mono 匂 graph graph of the Asteraceae-Anthemideae. Bull. Na t. Toyama no. 4: 1-15 (in Japanese). His t. Mus. Lond. (Bo t.) 23(2): 71-177. 一一一, Tanaka R., Taniguchi K. and Shimotomai N. Brummitt Brummitt R. K. 1998. Report of the committee for 1987. Species of wild in Japan: Spermatophyta: Spermatophyta: 46. Taxon 47: 443 -4 44. cytological and cytogenetical view on its entity. Greuter Greuter W. , McNeill J. , Barrie F. R., Burdet H. M. , Acta Phytotax. Geobo t. 38: 241-259 (in Japanese). Demoulin Demoulin V. ,Filgueiras T. S. , Nicolson D. H. , Nishikawa T. and Kobayashi H. 1989. Chromosome Silva Silva P.C. ,Skog J. E. , Trehane P, Turland N. J. , number and distribution of Chrysanthemum and and Hawksworth D. L. 2000. Intemational Code of arcticum L. J. Jpn. Bo t. 64: 77-84 (in Japanese). Botanical Botanical Nomenclature (Saint Louis Code). Peng C. I., Chung K. F. and Li H. L. 1998. Koeltz Koeltz Scientific Books ,Konigstein. Compositae. In: Huang T. C. et al. (eds よFl ora of Hind D. J. N. 1988. Dendranthema yezoense. Kew Taiwan 2nd ed. 4: 807-110 1. Editorial Committee Mag. 5: 55-60. of the Fl ora of Taiwan , Department of Botany , Hotta Hotta M. ,Yamakawa N. ,Hirai Y. and Shiuchi T. 1996. National Taiwan University ,Taipei. Taxonomical Taxonomical notes on plants of southem Japan 111. Poljakov P. 1955. Duo gen 町 e novae fam. Compositae. Distribution Distribution and of the Dendranthema No t. Sys t. Herb. Ins t. Bo t. Ak ad. Sci. URSS 17: ornatum ornatum group around southem Kyushu. Acta 418 -4 3 1. Phytotax. Phytotax. Geobo t. 47 (1): 91-104 (in Japanese). Shih C. 1979. Species et combinationes novae tribus Ki tamura S. 1935. (Taxonomy of Japanese chrysanthe- Anthemidearum familiae Composit 訂 um Tibeticae. mums). mums). Acta Phytotax. Geobo t. 4(1): 35-38 (in Acta Phytotax. Sin. 17(2): 13-116 (in Chinese). Japanese). Japanese). 一一一 1994. A revision of Ajania potaninii (Kr asch.) 一一一 1940. Compositae Japonicae. Pars secunda. Poljak. group (Compositae). Acta Phytotax. Sin. 32 Mem. Coll. Sci. Kyoto Imp. Univ. ser. B ,15(3): (4): 365-368. 285 -4 46. 一一一 and G.-X. Fu. 1983. Angiospermae-Dicotyledon 一一一 1967. Report on the distribution of the wild chry- eae-Compositae (3) ーAnthernideae. In: Ling Y. and santhemums santhemums of Japan. Acta Phytotax. Geobo t. Shih C. (eds.) ,Fl ora Reipublicae Popularis Sinicae 22( 4- 6): 109-137 (in Japanese). 76(1): 1-149. Science Press ,Beijing (in Chinese). Shimotomai Shimotomai N. , Adachi S. and Matsumori S. 1968. 一一一 1978. Dendranthema et N 伊, 'P onanthemum. Acta Phytotax. Phytotax. Geobo t. 29: 165-170. Cytological ,mo 中hological and geographical stud- 一一一 198 1. Compositae (Asteraceae). In: Satake Y. , ies on Ch ηsanthemum shiwogiku var. kinokuni- June June 2004 Journal of Japanese Botany Vo l. 79 No. 3 195

ense. ense. Bo t. Mag. Tokyo 81: 498-505 (in Japanese). Leningrad (in Russian). Trehane Trehane P. 1995. (1 172) Proposal to conserve 一一一 1985. Arctanthemum. Novi t. Sys t. Pl. Vasc. 22: Chrysanthemum L. with a conserved .type 274 (in Russian). (Compositae). (Compositae). Taxon 44: 439 -4 41. 一一一 1987. Arctantherum (Tzve l.) Tzve l. Fl. Arc t. Tzvelev Tzvelev N. N. 196 1. Fl ora URSS 26. Nauka , URSS 10: 11 4- 117 (in Russian).

大橋広好a,米倉浩司b .アジア産キク科キク属の 新組み合わせと日本産種一覧 キクの仲間はかつては広義の Chrysanthemum L. ことが知られている.このため,北村四郎博士は の下に分類されてきたが 近年はこれを複数の属 これらの属をキク属に含め それぞれ節として取 に細分することが多く,細分した場合の属名とし り扱った.われわれは北村博士のキク属の概念を てはこれまで Dendranthema (DC.) Des Moulins が 採用する. Bremer and Humphries (1 993) は, Ajania 用いられてきた.しかし, 1995 年に,これまで、シュ とキク属との差異が小さいことを認めながらも, ンギク Chrysanthemum coronarium L.をタイプと 両者を合一することで必要となる多数の新組合せ してきた Chrysanthemum L.をシマカンギク仁 を避けて両属を維持し,かえって日本産のイソギ indicum indicum L.をタイプとして保存することが提案さ クとシオギクを Ajania に組み替えた.しかし,こ れ,これがセントルイスの第16 回国際植物学会議 こでは両属を同一属として扱ったので,中国から で認可された.この結果 ,Chrysanthemum は現行 中央アジアにかけて最近記載された多数の Ajania の国際植物命名規約付則 111 の保存属名中に掲載 の種を Chry ・santhemum に組み替えることが必要と されている (Greuter et al. 2000). このため,キク なった. この群に属するキノクニシオギク仁 属の学名は Chrysanthemum に戻さねばならない. shiwogiku var. kinokuniense Shimo t. & Ki tam. は, 日本産のキク属植物は大部分 Chrysanthemum の下 イソギクとシオギクの交雑由来の植物であること で記載されてきたので,新組み替えの必要がある が下斗米直昌博士らの研究により既に明らかになっ ものは少ないが,近年に Dendranthem αの下で記 ているので種として扱った. 載されたトカラノギクや古い栽培植物のカモメギ この他,チシマコハマギクの学名の変更と,リユ クと品種関係にあることが分かつたキクタニギク ウノウイワインチン(リユウノウギク×イワイン については組み替えが必要となった. チン)に対する新名の提唱を行った.チシマコハ 中国やロシアでは,頭花に舌状花を欠くイワイ マギクに対して新たに定めた学名は,類似するコ ンチンの仲間を Ajan 臼や Phaeostigma ,痩果の性 ハマギクの学名と紛らわしいが,命名法上やむを 質が少し異なるチシマコハマギクを Arctanthemum えない. として別属で扱うことが多く,これらの属は世界 最後に,日本産のキク属植物の学名を Ch η'san- のキク亜科植物の属レベルのモノグラフをまとめ themum の下で整理した一覧を示した. た Bremer and Humphries (1993) にも採用されてい (a 東北大学大学院理学研究科 る. しかし,これらの属の識別形質は小さい差異 附属植物園津田記念館,

であり,また,イワインチンの仲間は日本ではキ b東北大学大学院理学研究科 クの仲間とふつうに交雑して遺伝的な隔離はない 附属植物園八甲田山分園)