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A Taxonomic Revision of L. Subgenus Anemonanthea (DC.) (DC.) J uz. sensu lato () 1

Svetlana Svetlana ZIMAN¥ Carl S. KEENER b， Yuichi KADOTA C， Elena BULAKH¥ Olga TSARENKO a and Bryan E. DUTTON d

aM. aM. G. Kholodny lnstitute of Botany of the National Academy of Sciences ， Tereshchenkivska Tereshchenkivska st r. 2，Kiev ，01601 UKRAINE; b208 b208 Mueller Laboratory ， Pennsylvania State University ， University University Park ，Pennsylvania ，16802 U. S. A.; CDepartment CDepartment of Botany ， National Science Museum ， Amakubo 4- 1-1 ，Tsukuba ，lbaraki ， 305-0005 JAPAN; dDetartment dDetartment of Biology ，Western Oregon State University ， Monmouth ，Oregon ，97361 U. S. A.

(Received on July 11 ， 2003)

As treated here ，Anemone subgen. Anemonanthea includes three sections (viz. ， Anemonanthea ，Rosulantes sec t. nov. and Tuberosa). Two sections (viz. ，Stolonifera and

Keiskea) 紅 e sep 訂 ated from the foregoing ones and placed in a different subgenus ，viz. ， Stolonifera ， comb. and sta t. nov. We elaborated a conspectus of subgenus Anemonanthea s. s. 1.， together with a very short mo 中hological delimination of two subgenera ，five sec- tions tions and 11 問 ries ，included latin diagnoses of seven new infrageneric taxa. A key for determination of subgenera ，sections ，series and species is included. For delimitation delimitation of the taxa within Anemonanthea s. 1.， although we used the morphological characters characters as given by Tamura (1 995) ， in our research we have included additional essen- tial tial characters' (viz. ，chromosome numbers ， types of pollen grains and ca 中els ，germina- tion tion type and time of basal development ， types of ， shoot structure ， shape and and size of basal and involucralleaves ， number ，shape ，venation and villosity.

In In this Part 1 the species No. 1 to No. 13 紅 'e taxonomically revised.

Key words: Anemone ，morphology ，phylogeny ， subgenus Anemonanthea ， .

Anemone nemorosa L. and its allied spe- species ，its taxonomic rank ，and state and re- cies cies (sec t. Anemonanthea sensu DC.) 紅 e lationships among its members. The aim of mainly geophytic ephemeroids with long our paper is to examine the taxa of Anemone sympodial rhizomes ， ternate or 3-sected subgenus Anemonanthea ，and specify their basal basal ， l-few-flowered scapes ，and status and relationships. short-puberulent short-puberulent achenes with short persis- tent tent styles and which are distributed in shady Materials and Methods localities localities within the temperate floras of the Our 佐eatment is based mainly on herbar- Northem Hemisphere. Beginning with ium .material involving about 2000 speci-

Pritzel Pritzel (1841) ， the author of the first mono 同 mens contained in more than 20 collections graph graph on Anemone ， until the present (BM ，CLU ，E ，GH ，J ，K ，KW ，LE ，NY ，TNS ， time ，there 訂 'e many different opinions on US ， VLA ， W ， WU ， etc.; abbreviations the the size and structure of the above group of according to Holmgren et al. 1990). From

--4 3ー 44 植物研究雑誌第79 巻第1号 平成16 年2月 these these specimens ， both and fruit sam- species ， with elongate rhizomes and petiolate ples ples (about 400) were studied. Standard involucral leaves) ， rhizomes and petiolate anatomical anatomical techniques were employed ， involucral leaves) ，and Stolonifera Ulbr. (A. including including light and scanning microscopy (the baicalensis Turcz. and al1 ied species ， with latter latter for examination of achenes). We also stolon-like rhizomes and mainly sessile re- examined ca. 300 collected mainly in duced involucralleaves). 1990-1997 in ca. 20 natural populations of The Russian taxonomist Juzepchuk later five five taxa. raised the rank of sec t. Anemonanthea to a subgenus subgenus with three sections (which co 町 e- Literature Literature Survey spond to the Ulbrich's subsections) plus sev- Anemone nemorosa L. and three allied eral series (Juzepchuk 1937). species species (A. trifolia L. ， A. ranunculoides L. Tamura (1967) was the first to regard the and A. quinquefolia L.) were described by A. nemorosa complex as sec t. Anemone. In Linnaeus Linnaeus (1753). Initially these taxa were in- subsequent papers ，Tamura (1 991 ， 1995) fol- cluded cluded in sec t. Anemonanthea DC. (Candolle lowed mainly the work of Juzepchuk in ac- 1817 ，1824) ，together with plants having tu- cepting this group of taxa as subgen. berous berous rhizomes (viz. ， A. coronaria L.) ，and Anemonanthea ， but he correc t1 y recognized as as circumscribed by Candolle this section A. nemorosa and allied species as sec t. initia l1 y consisted of plants with both lanate Anemonanthea (not Anemone nor Sylvia) and subglabrous achenes. because of De Candolle 's priority. According Shortly Shortly after Candolle' s taxonomic 位eat- to Tamura (1 995) ， subgenus Anemonanthea ment ，Gaudin (1 828) circumscribed the A. includes four sections ，viz. ，Anemonanthea ， nemorosa-group nemorosa-group as sec t. Sylvia. A number of Stolonifera ， Tuberosa ， plus Keiskeana authors authors have followed Gaudin's treatment Tamura. (viz. ，Spach 1838 ， Janczewski 1892 ， Chater On the other hand ， there were several at- 1973 ，Hoot et al. 1994) ， but other authors tempts to regard this group of species as (viz. ， Pritzel 1841 ， Prantl 1887 ， Ulbrich genera -A nemonoides Mill. (Miller 1754) or 1906 ，Femald 1928 ，Wang 1974 ， 1980) sim- Anemonanthea (DC.) S. F. Gray (Gray 1821). ply ply left this group the name Anemonanthea. Several years ago these proposals were sup- Pritzel Pritzel (1841) produced first monograph ported by Holub (1 973) and later by on Anemone in which he classified within Starodubtsev (1 989 ， 1991). Moreover ， sec t. Anemonanthea several allied species Starodubtsev proposed to sep 紅 ate two small (A. (A. nemorosa and A. ranunculoides) ， but also groups of species of Anemonanthea as gen- A. A. richardsonii Hoo k. and A. antucensis era (Tamuria Starod. and Arsenjevia Starod). Poepp. ， both of which are rather unlike the Hoot et al. (1 994) recen t1 y proposed a two former species. According to Finet and classification scheme of Anemone quite diι Gagnepain (1904) ， A. nemorosa ， A. nikoensis ferent from the preceding authors in that ，ac- Maxim. and other closely related species cording to them ， the A. nemorosa and A. were included in a so called fourth section ， baicalensis groups belong to diverse subgen- together together with A. hepatica L. and other spe- era: the A. nemorosa complex to subgen. cies cies which were app 紅 en t1 y distant from this Anemone (plants with x = 8) ，sec t. Anemone group. group. (A. nemorosa ， A. baldensis L. ， A. multifida The second comprehensive monograph on Poi r.， etc.) ，and the A. baicalensis complex Anemone was by Ulbrich (1906) who di- was placed in subgen. Anemonidium (plants vided vided sec t. Anemonanthea into three subsec- with x = 7) ，sec t. Keiskea (A. baicalensis ， A. tions: tions: Sylvia Gaudin (A. nemorosa and al1 ied keiskeana T .l to ex Maxim. ，etc ふ February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 45

On the basis of new molecular data ， with distinct or hardly recognizable styles ， Ehrendorfer Ehrendorfer and Samuel (2001) confirmed and linear or dilated ， sometimes subcapitate the the affinities of the A. nemorosa and A. stigmas) ， and germination type (epigeal or blanda blanda groups. hypogeal; Tamura et a1. 1977 ，Ziman 1985 ， Barykina Barykina and Potapova 1994 ，etc.). We used Biological Biological and Morphological these characters ， in addition to those men- Characteristics Characteristics tioned by Tamura ， for delimitation of taxa All All plants within subgenus Anemonanthea within the subgen. Anemonanthea s. 1. 紅 e characterized by prostrate ，more or less According to Tamura ，sec t. Anemonan- elongate ， sometimes tuberous rhizomes ， few thea is characterized by 3-5-sected ，ternate long-petiolate long-petiolate basal leaves (sometimes ab- to bitemate petiolate basal leaves ，well- sent sent at the base of stem) ， 3-1eaved involu- developed 3-sected to temate involucral cres ， one to four with 5 to 20 ， leaves ，1-3 イlowered cymes ，many ovoid to sessile sessile or rarely stalked ，sc 紅白ly compress- ellipsoid ， hardly comp 印 ssed ， short- ed ，short-hirsute or rarely glabrate achenes pubescent ， sessile achenes with short or with with curved or straight styles and usually slightly elongate styles ， and elliptic or capitate capitate stigmas (cf. Tamura 1995). Flower- capitate stigmas ， and achenes a町 anged in ing ing occurs mainly in spring (ephemerals and globose heads. geophytes). geophytes). The plants prefer to grow in In addition to these characters ， the taxa of shady shady forests or under bushes ， and hypogeal sec t. Anemonanthea have several important germination germination is found in many species. characteristics in common: several basal Our comp 紅 ative analysis confirmed the scale-like and persistent leaves ， in addition presence presence in these taxa of most of the forego- to solitary leaves with long na 町 ow petioles ing ing characters. However ， there is a rather and distinct blades which develop after distinct distinct differentiation within subgen. anthesis on rhizomes distinct from reproduc- Anemonanthea sensu Tamura of several es- tive shoots (which are sometimes regarded as sential sential characters such as the of type absent because they soon perish) ， hypogeal (l ong thin ，short-nodulose or tuberous) ， shoot germination ， and 3-colpate pollen grains. structure structure (non-rosetteous or semirosetteous) ， However ，it is necess 訂 Y to sep 訂 ate sev- time time of basal leaf development (before or eral species from sec t. Anemonanthea be- after after anthesis) ， shape of basal parts of the cause they have several basal leaves with basal basal leaf petioles (na 町 ow ， vaginated or distinct blades developing before anthesis ， sharply sharply dilated and scale-like) ， types of basal lack of scale-like leaves (but basal remnants leaves leaves (l ong-petiolate with distinct blades or of old leaf petioles) ， epigeal germination ，

small small and scale 四 like) ， size and shape of and pantocolpate pollen grains. These spe- involucral involucral leaf petioles and blades ， and tepal cies are classified under sec t. Rosulantes. number ， shape ， venation and villosity. Within both sections there are plants with Moreover ，plants within this subgenus differ either a monomorphic or a dimorphic in in their chromosome numbers (x = 8 or x = perianth ， dilated or filiform filaments ， 7; 7; Kurita 1955 ，1956 ， Baumberger 1970 ， puberulent or subglabrous ovaries and Starodubtsev Starodubtsev 1991 a， 1991 b， unpublished achenes (some with na 町ow lateral ribs) ， with data data of Qin- Er Yang ，etc よ types of pollen linear or subcapitate stigmas. Therefore we grains grains (3-colpate ，pancolpate or polycolpate; regard these sections as including several se- Kumazawa 1936 ， Huynh 1964 ， 1970 ， ries: Anemonanthea with five series Savitski Savitski 1982 ，etc.) ，mo 中hological variation [Anemonanthea (= Hylalectron) ， Reflexae ofthe ofthe ca 叩els and achenes (sessile or stalked ， Ulbrich ，Altaicae Starod.] ， and two new ones 46 植物研究雑誌第79 巻第1号 平成16 年2月

(Nikoenses (Nikoenses and Quinquefoliae ，series nov よ 13 tepals ，dilated filaments and few stalked and and Rosulantes with two series (Rosulantes compressed achenes with distinct styles. and and Exiguae ，series nov ふ With Starodubtsev (1991) ，we believe that A. According According to Tamura ， plants of sec t. keiskeana is allied to A. deltoidea ， but we re- Tuberosa Tuberosa have tuberous rhizomes ，3-5-secte g訂 d this group of species as a section ， not a d or temate basal leaves ，well-developed sep 紅 ate genus. In our opinion ， the both spe- petiolate petiolate involucral leaves similar to basal cies belong to the diverse monotypic series leaves ，solitary f1 0wers with 8-20 linear- which differs by the seasonal rhythms as oblong oblong tepals ，sessile achenes ， these mainly noted above ， shape of involucral leaves ， and ovoid ，hardly compressed and shortly pubes- specific tepal and filament characteristics. cen t. In In our opinion ， the following characters Conspectus are are also essential for sec t. Tuberosa: basal Anemone L. ， Sp. Pl. 538 (1 753). leaves leaves which develop before anthesis ，their Type: A. coronaria L. petioles petioles basally sharply dilated (“ ears") and Subgenus Anemonanthea (DC.) Juz. ， F l. scale-like ， germination epigeal ， and 3- URSS 7: 241 (1 937). colpate colpate pollen grains. In addition ， these Type: A. nemorosa L. plants plants have a dimorphic perianth ， and the Genus Mil l.， Gard. Dic t. subglabrous subglabrous ovaries and achenes. Abr. ed. 4. 28 (1 754). As stated by Tamura ，sec t. Stolonifera is Genus Anemonanthea (DC.) S.F.Gray ， characterized characterized by f1 eshy or thin rhizomes ，1 to Na t. Arr. Bri t. Pl. 2: 724 (1 821). few basal 3-5-sected to temate leaves ，two Sec t. Stephanomata Spreng. ，Sys t. Veg. 1: or or three sessile involucral leaves (often re- 660 (1 825). duced duced and basally connate) ，solit 紅 y f1 0wers Rhizomes long ，白t hi 血n and 泊shor 抗tι. 幽幽目幽幽幽但 with with 5-6 tepals ， and sessile ，scarcely com- sometimes tuberous. Basal leaves with pressed ， mainly ovoid and shortly pubescent temate blades or p征 tly scale-like. Involucral achenes achenes with short styles and nearly sessile leaves petiolate. Ovaries and achenes sessile ， capltate capltate st1 gmas. with distinct styles and small ， mainly linear However ，we are adding to these charac- stigmas; basic chromosome number x = 8. ters ters the presence of short stout or long stolon-like stolon-like rhizomes ，partly reduced involuc- Sec t. Anemonanthea DC. ，Sys t. Na t. 1: ral ral leaves (bracteoles) ，epigeal germination ， 196 (1 817). and and polycolpate pollen grains. Within sec t. Sec t. Sylvia Gaudin ， F l. Helv. 3: 490 Stolonifera Stolonifera s. st r.， we recognize two series as (1 828) ，p. described described by Juzepchuk (Stoloniferae or Subsec t. Sylvia (Gaudin) Ulbr. in Bo t. Flaccidae Flaccidae and Baicalenses). We believe they Jahrb. 37: 187 (1906). differ differ mainly by the presence or absence of Sec t. Anemone sensu Tamura in Sci. Rep. basal basal scales and rosetteous leaves with dis- Osaka Univ. 16: 25 (1967) ，p. p. ， nom. nud. tinct tinct blades ， and the time of their seasonal Rhizomes monomorphic or dimorphic ， but development ， whether before or after non tuberous. Basal leaves 2-3 ，scale-like ， anthesis. anthesis. persistent; solitary leaves with distinct peti- According According to Tamura ，sec t. Keiskea is oles and blades developing on rhizomes monotypic monotypic and it is characterized by f1 eshy ap 紅 t from f1 0wering stems after anthesis. elongate elongate rhizomes ， one to few basal temate Cymes l-few- f1 owered. Tepals 5-15 ，mono- leaves ， sessile involucI 叫 leaves slightly morphic or dimorphic. Pollen grains 3- connate connate at the base ，solitary f1 0wers with 5- colpate. Germination hypogea l. February February 2004 Joumal of Japanese Botany Vo l. 79 No. 1 47

Ser. Ser. Anemonanthea. pseudoaltaica var. gracilis (H.Hara) H.Ohba Sec t. Hylalectron Irmisch in Bo t. Zei t. 14: var. katonis H.Ohba 19 19 (1856). 13. A. raddeana Regel Ser. Ser. Hylalectron (Irmisch) Ulbr. ， l. c. (1 906). Ser. Nikoenses Ziman ，Kadota & Bulakh ， Subsec t. Hylalectron (Irmisch ex Ulbr.) se r. nov. Juz. ，l. c. (1937). Type: A. nikoensis Maxim. Ser. Ser. Nemorosae Tamura ，l. c. (1 967) ，nom. Sec t. Anemonanthea DC. ，l. c. (1 817) ， p. nud. nud. p. Genus Anemonoides sec t. Anemonoides Rhizomata dimorphica. Foliola in Starod. Starod. p. p. ，subsect t. Anemonoides ，Ranun- petiolulatum 3-10 mm longa. Involucri folia culoides culoides St 紅 od. ，sec t. Umbrosa Starod. in in petiolum 3-5 mm lata. Perianthii tepala 5， Bot. Bot. Zhum. 74: 1346 (1 989). anastomosi nervi 5-9 ， dense pilosa. Rhizomes monomorphic ，long ， 1-3 mm in Rhizomes dimorphic. Leaflet petiolules 3- diameter ，branched ， mainly pros 甘ate. Leaflet 10 mm long. Involucralleaf petioles 3-5 mm petiolules petiolules mainly 1-3 mm ， rarely 3-5 (-10) wide. Tepals 5， with 5-9 vein anastomoses ， mm long. Involucral leaf petioles mainly 1- densely puberulen t.

2 mm wide. Tepals 5-8 (r 訂 ely-9 ，10) ， 14. A. nikoensis Maxim. spreading ， generally lacking vein anasto- moses ， pubescent or glabrous. Ser. Re f1 exae Ulbr. ，1. c. 194 (1 906). 1. 1. A. nemorosa L. Lectotype: A. reflexa Stephan ex Willd. 2.A. α murensis (Korsh.) Kom. Subsec t. Reflexa (Ulbr.) Juz. ，l. c. 250 3. 3. A. caerulea DC. (1 937). 4. 4. A. uralensis DC. Genus Anemonoides sec t. Anemonoides 5. 5. A. ranunculoides L. subsec t. Reflexae (Ulb r.) Starod. ，Vetrenytsy: 6. 6. A. udensis Trautv. & C.A.Mey. Sys t. Evo 1. 123 (1 991). 7. 7. A. trifolia L. ，v 紅 . tr ぴolia ， var. albida Rhizomes dimorphic. Leaflet petiolules 5- (M 紅 iz) Ulbr. ，v 紅 . brevidentata Ub aldi & 15 mm long. Involucralleaf petioles 1-2 mm Puppi wide. Tepals 5-7 ， without vein anastomoses ， 8. 8. A. umbrosa C.A.Mey. puberulen t. 9. 9. A. soyensis Boiss. 15. A. reflexa Stephan ex Willd. 10. 10. A. debilis Fisch. ex Turcz. Ser. Ser. Quinquefoliae Ziman ，Kadota & Ser. Ser. Altaicae (Starod.) Ziman ，Kadota & Bul 北 h ，ser. nov. Bulakh ，comb. et sta t. nov. Type: A. quinquefolia L. Type: Type: A. α[t， α ica Fisch. ex C.A.Mey. Sec t. Anemonanthea subsec t. Hylalectron Genus Anemonoides sec t. Anemonoides p. (Irmisch) Juz. ，1. c. 241 (1 937) ， p. p. p. ，subsec t. Altaicae Starod. ，l. c. Genus Anemonoides sec t. Anemonoides (1989). (1989). subsec t. Anemonoides St 征 od. ，1. c. (1 991). Rhizomes dimorphic ， short ， thick Rhizomata dimorphica. Foliola in (nodulose) ，and long ，thin. Leaflet petiolules petiolulatum 1-2 mm longa. Involucri folia 2-5 (-10) mm long. Involucralleaf petioles in petiolum 1-2 mm lata. Perianthii tepala mainly 3-5 mm wide. Tepals 8-15 ， with 3- imprimis 5， anastomosi nervi imprimis nulla ， 5 vein anastomoses ，glabrous. glabra. 1. 11. A. altaica Fisch. ex C.A.Mey. Rhizomes dimorphic. Leaflet petiolules 1- 12. 12. A. pseudoaltaica H.Hara v紅. 2 mm long. Involucral leaf petioles 1-2 mm 48 植物研究雑誌第79 巻第1号 平成16 年2月 wide. wide. Tepals mainly 5， without vein Ser. Exiguae Ziman ，Kadota & Bulakh ， anastomoses ，glabrous. se r. nov. 16. 16. A. quinquefolia L. subsp. quinquefolia ， Sec t. Anemonanthea DC. ，1. c. (1 817) ， p. subsp. subsp. minima (DC.) Frodin p. 17. 17. A. lancifolia Pursh Type: A. exigua Maxim. 18. 18. A. piperi Britton ex Rydb. Folia involucria in petiolum 0.5-1.5 cm 19. 19. A. grayi Behr & Kellogg subsp. grayi ， longa ， bracteola nulla. Caula florifera susp. susp. lyallii (Britton) B.E.Dutton & a1. solitaria ，floris solitaria. Tepala subglabra ， 20. 20. A. oregana A.Gray anastomosi nervi nulla ， staminodia nulla. Involucral Involucral leaf petioles 0.5-1.5 cm long; Sec t. Rosulantes Ziman & Kadota ，sec t. bracteoles absen t. Scapes solitary ， 1- nov. nov. flowered. Tepals subglabrous ， lacking vein Type: Type: A. stolon 俳ra Maxim. anastomoses; staminodes absen t. Sec t. Anemonanthea DC. ，1. c. (1 817) ， p. 23. A. gr if.f ithii Hook .f. & Thoms. p. p. 24. A. exigua Maxim. Genus Anemonoides sec t. Stolonifera 25. A. scabriuscula W.T.Wang (Ulbr.) (Ulbr.) Starod. ，1. c. 122 (1 991) ， p. p. Rhizomata longa ramosa. Folia radicalia Sec t. Tuberosa Ulb r.， 1. c. (1906). 2-5 ， evolvuntur ante anthesin ， in petiolum in Genus Anemonoides sec t. Tuberosa p紅 s basalis gradatim dilatatus ， basi residius (Ulbr.) Starod. in Bo t. Zhurn. 76: 123 petiolorum petiolorum vestiti. Inflorescentia pauciflorae (1 989). or or flores solitarii. Tepala 5-7 ，monomo 中ha. Lectotype: A. apennina L. Grana pollinis pantocolpata. Germinatio Rhizomes tuberous. Basalleaves 1-3 ，de- eplgaea. eplgaea. veloping before anthesis. Basal leaf petioles Rhizomes long ， branched. Basal leaves 2- basally sharply dilated ，scale-like. Cymes 1- 5， developing before anthesis. Basal leaf flowered. Tepals 8-20 ， dimorphic. Pollen petioles petioles basally gradually dilated ， sur- grains 3-colpate. Germination epigea 1. rounded by fibrous remnants. Cymes l-few- 26. A. apennina L. flowered. flowered. Tepals 5-7 ，monomorphic. Pollen 27. A. blanda Schott & Kotschy grains grains pantocolpate. Germination epigea 1. 28. A. caucasica Willd.

Ser. Ser. Rosulantes. Subgen. Stolonifera (Ulbr. ex Juz.) Sec t. Anemonanthea DC. ，1. c. (1 817) ， p. Ziman ，Kadota & Bulakh ，sta t. nov. p. p. Subsec t. Stolon{ 作ra Ulb r.， 1. c. (1 905) ， p. Type: Type: A. stolon{ 作ra Maxim. p. Folia Folia involucria in petiolum 1-3 cm longa ， Sec t. Stolonifera (Ulbr.) Juz. ，1. c. (1937). bracteola bracteola parva adsun t. Caula florifera 2-3. Sec t. Keiskea Tamura ，1. c. (1967). Inflorescentia Inflorescentia pauciflorae. Tepala pilosa ， Lectotype: A. baicalensis Turcz. ex Ledeb. anastomosi anastomosi nervi adsunt ut nulla ， staminodia Rhizomata longa aut brevita ， non adsun t. tuberifera. Folia involucria sessiles. Ovaria Involucralleaf Involucralleaf petioles 1-3 cm long; small et achenia estipitata ut stipitata ， stylodii bracteoles bracteoles presen t. Scapes 2-3 ， few- conspicua ut vix visibiles ， stigmata imprimis flowered. flowered. Tepals puberulent ， having or lack- subcapitata. Grana pollinis polycolpate. n = ing ing vein anastomoses; staminodes presen t. 7. 21. 21. A. stolonifera Maxim. Rhizomes long or short ， but non-tuberous. 22. 22. A. davidii Franch. Involucral leaves sessile. Ovaries and February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 49 achenes achenes sessile or stalked ， with distinct or Sec t. Keiskea Tamura emend. Ziman & hardly hardly recognizable styles and mainly Kadota. capitate capitate stigmas. Pollen grains polycolpate. Sec t. Anemonanthea DC. ，1. c. (1 817) ， p. Basic Basic chromosome number x = 7. p. Subsec t. Sylvia Gaudin ，1. c. 193 (1 905) ， p. Sec t. Stoloniferae. p. Genus Arsenjevia Starod. ，1. c. (1 989). Genus Tamuria Starod. ，1. c. (1 991). Lectotype: Lectotype: A. baicalensis Turcz. ex Ledeb. Lectotype: A. keiskeana T .I to ex Maxim. Rhizomes short ， stout and long ，stolon- Rhizomes long ， branched. lnvolucral like. like. lnvolucral leaves several (part of them leaves 3 ，reduced. Cymes l- f1 owered. Tepals reduced). reduced). Cymes 1-3- f1 owered. Tepals 5-7. 5-13. Ovaries and achenes stalked ， covered Ovaries Ovaries and achenes sessile. Styles indis- with hairs 0.1-0.3 mm long; styles distinct; tinct; tinct; stigmas mainly capitate. stigmas liner or slightly dilated. Pollen Ser. Ser. Stoloniferae. grains 3-colpate or polycolpate. Ser. Ser. Baicalenses Juz. ，1. c. (1937). Subsec t. Stolonifera Ulbr. ，1. c. (1 906) ， p. Ser. Keiskea. p. p. Sec t. Keiskea Tamura in Sci. Rep. Osaka Lectotype: Lectotype: A. baicalensis Turcz. ex Ledeb. Univ. 16: 26 (1 967). Scales-like Scales-like basal leaves absent; basal Lectotype: A. keiskeana T .I to ex Maxim. leaves leaves with distinct blades and basally Basal leaves several ，3-sected ， with dis- vaginate vaginate petioles developing before anthesis. tinct petioles and blades ， developing before Carpel Carpel stigmas subcapitate ， achene stigmas anthesis. lnvolucral leaves 3-10bed. Tepals slightly slightly dilated. 10-22 ，linear-lanceolate ，puberulent; ovaries 29. 29. A. baicalensis Turcz. ex Ledeb. ， var. and achenes compressed. baicalensis ， v征 . kansuensis (W.T.Wang) 33. A. keiskean αT.I to ex Maxim. W.T.Wang ， var. saniculiformis (C.Y.Wu) Ziman & B.E.Dutton ， var. litoralis Litw. ， Se r. De It oidea Ziman ，Kadota & Bulakh ， V 紅 . glabrata Maxim. ser. nov. 30. 30. A. prattii Huth ex Ulbr. Sec t. Anemonanthea DC. ，1. c. (1 817) ， p. p. p. Ser. Ser. Flaccidae Juz. ，1. c. (1 937). Type: A. deltoidea Hook. f. ex Douglas Subsec t. Stolonifera Ulb r.， 1. c. (1 906) ， p. Folia radicalia imprimis scariosa et p. p. solitaria e laminae evolvuntur ante anthesin. Lectotype: Lectotype: A. flaccida F. Schmidt lnvolucri folia integerrima. Perianthii tepala Scale-like Scale-like basal leaves present; basal 5，ovata ， glabra; ovaria et achenia non leaves leaves with distinct blades and n紅 TOW peti- compressa. oles oles developing after anthesis. Carpel stig- Basal leaves few ，scale-like ，persistent ， mas capitate or subcapitate ， achene stigmas and solitary ， with distinct petioles and blades subcapitate subcapitate or dilated. developing after anthesis. lnvolucral leaves

1. 31. A. flaccid αF， .Schmidt ，v 紅 . flaccida ， undivided. Tepals 5，ovate ，glabrous; ovaries var. var. hirtella W.T. Wang ，v 紅 . hofengensis and achenes not compressed. (W.T.Wang) Ziman & B.E.Dutton ， var. 34. A. deltoidea Hook .f. ex Douglas anhuensis anhuensis (W. T. Wang) Ziman & B .E.Dutton 32. 32. A. delavayi Franch. ，var. delavayi ，var. Key to the species of Anemone subgenus oligocarpa oligocarpa (S .J .Pe i) Ziman & B.E.Dutton Anemonanthea a. 1a. Rhizomes long ， thin and short nodulose ， 50 植物研究雑誌第79 巻第1号 平成16 年2 月

sometimes sometimes tuberous; involucral leaves ...・ H ・... ..1. A. nemorosa petiolate; petiolate; ovaries and achenes sessile ， with 6b. Central leaflets of basal leaf blades 3- distinct distinct styles and mainly linear stigmas; sected or 3-parted ，rhombic ， with semicor- basic basic chromosome number n = 8 (subgen. date bases; tepals 5-8 ，monomorphic ，basally

Anemonanthea) Anemonanthea) ...・ H ・..…...・ H ・...・ H ・...・ H ・H ・H ・..2 narrowed ，lacking vein anastomoses; achene

lb. lb. Rhizomes long or short ， but non- stigmas sublinear... ・H ・.....・ H ・.2. A. amurensis tuberous; tuberous; leaves involucral sessile; ovaries 7 a. Tepals basally narrowed ， mainly blue or and and achenes sessile or stalked ， with distinct whitish ， densely puberulent; filaments di- or or hardly recognizable styles and mainly lated basally …...・ H ・..…...・ H ・....3. A. cα erulea capitate capitate stigmas; basic chromosome number 7b. Tepals basally rounded ， mainly rose or n = 7 (subgen. Stolonifera)... ・H ・.....・ H ・..…・ .29 red ，sparsely puberulent; filaments dilated

2a. 2a. Basal leaves several ，scale-like ，persis- apically...... ・H ・.....・ H ・..…...・ H ・..4. A. uralensis

tent; tent; solitary leaves with distinct blades and 8a. Cymes l-few 町 flowered; tepals pubes- long long petioles developing on rhizomes after cent ，usually with vein anastomoses; achenes

anthesis; anthesis; achene styles mainly 1- 1. 5 mm slightly compressed ， with lateral ribs ...・ H ・..9 long long (sec t. Anemonanthea) …...・ H ・.....・ H ・..… 3 8b. Cymes 1- flowered; tepals glabrous or 2b. 2b. Basalleaves several ， with distinct blades subglabrous ， lacking vein anastomoses; and and long petioles developing on reproductive achenes compressed or not compressed. … 11 shoots shoots before anthesis; achene styles mainly 9a. Basal and involucral leaflets 2-3-10bed

0.5-0.7 0.5-0.7 mm long ・… H ・H ・.....・ H ・...・ H ・H ・H ・-… 22 (sometimes parted); involucral leaf petioles 3a. 3a. Rhizomes monomo 中hic ， long and 1-3 3-5 cm long; tepals dimorphic ，basally nar- mm in diameter ，branched ， mainly horizon- rowed; achene styles 0.8-1.5 mm long ・…・ tal; tal; tepals 5-8 (rarely 9-10) (ser. Anernonan- … 5. A. ranunculoides thea) thea) ...・ H ・H ・H ・..…...・ H ・.....・ H ・.....・ H ・..…...・ H ・..4 9b. Basal and involucral leaflets undivided; 3b. 3b. Rhizomes dimorphic ， short ， thick involucral leaf petioles 1-3 cm long; tepals

(nodulose) (nodulose) or long ，thin; tepals 5-12... ・H ・.13 monomo 叩hic ， basally rounded; achene

4a. 4a. Involucral leaf petioles 3-5 mm wide; styles 1.5-2.5 mm long ・…...・ H ・...・ H ・...・ H ・..10 carpel carpel styles ca. 1 mm long; stigmas 10a. Cymes l-flowered; tepals 5， mainly

subcapitate subcapitate or slightly dilated …...・ H ・.....・ ....5 pilose ， having 5-9 vein anastomoses.... ・H ・-…

4b. 4b. Involucral leaf petioles 1-2 mm wide; ...・ H ・H ・H ・.6. A.. udensis carpel carpel styles longer than 1 mm; stigmas lin- 10b. Cymes few-flowered; tepals 5-8 ， ear...... 8 ear...... 8 glabrous ， having solitary vein anastomoses...

5a. 5a. Involucral leaf petioles 1-3 cm long; …H ・H ・..7. A. trifolia involucral involucral leaf blades similar to the basal lla. Central segments of basalleaf blades 3- leaf leaf blades; cymes 1- flowered; tepals 5-8 ， lobed; tepals 5; achene hairs 0.2-0.3 mm

glabrous; glabrous; achene bodies (2-) 3-5 mm long … long...... ・H ・..…...・ H ・..…...・ H ・..… 8. A. umbrosa .....6 .....6 11 b. Central segments of basal leaf blades 5b. 5b. Involucral leaf petioles 3-5 mm long; se 町 ate or coarsely toothed; tepals 5-7;

involucral involucral leaf blades larger than the basal achene hairs ca. 0訓 mm long ...・ H ・H ・H ・...1 2 leaf leaf blades; cymes 1-2-flowered; tepals 5， 12a. Tepals 10-15 mm long ， basally pubescent ， achene bodies 1-2 mm long ・… .7 rounded ， white; achenes not compressed ， 6a. 6a. Central leaflets of basal leaf blades 3- without ribs ， 4- 5 mm long ， sp 訂 sely

lobed ， ellipsoid-oblong ， with wedge-like puberulent ...・ H ・....・ H ・..… H ・H ・-… 9. A. soyensis bases; bases; tepals 6-10 ， mainly dimo 中hic ， 12b. Tepals 4-8 mm long ，basally narrowed ， basally basally rounded ， having 3-5 vein white-greenish; achenes slightly compressed ， anastomoses; anastomoses; achene stigmas slightly dilated with narrow ribs ，2 -4 mm long ， densely February February 2004 Journal of Japanese Botany VoL 79 No. 1 51

puberulent puberulent ...・ H ・..…...・ H ・.....・ H ・..10. A. debilis puberulent; achene hairs less than 0.1 mm

13a. 13a. Tepals 8-15 ， glabrous ， spreading; long (ser. Reflexae) ...・ H ・....・ H ・.15. A. reflexa cymes 1- flowered (se r. Altaicae) …… H ・H ・..14 17b. Cymes 1-flowered; bracteoles absent; 13b. 13b. Tepals 5-8 ， glabrous or pubescent ， tepals 10-25 x 4-10 mm ，spreading ，glab- spreading spreading or bent; cymes 1-few-flowered. … rous; achene hairs 0.1-2 mm long (ser.

...1 6 Quinquefoliae) …...・ H ・..…....・ H ・.....・ H ・...・ H ・.18 14a. 14a. Basal leaf petiolules 5-10 mm long; 18a. Tepals 5， having solitary vein tepals tepals dimorphic; filaments filiform; ovaries anastomoses; involucral leaf petioles 0.5-2 and and achenes basally narrowed ， with na 町 ow cm long ， achene styles sparsely puberulent... lateral lateral ribs; achene hairs 0.1 -0 .3 mm long; ..19 achene achene styles puberulent; ca 中el stigmas 18b. Tepals 5-7 ，lacking vein anastomoses; subcapitate; subcapitate; achene stigmas slightly dilated … involucralleaf petioles 1-3 cm long; achene

… 11. A. αltaica styles glabrous ...・ H ・.....・ H ・.....・ H ・.....・ H ・..… .20 14b. 14b. Basal leaf petiolules 2-5 mm long; 19a. Lateralleaflets often deeply two-lobed; tepals tepals monomorphic; filaments basally di- basal leaf blades puberulent; involucral leaf lated; lated; ovaries and achenes basally narrowed blades and stems puberulent or subglabrous; or or rounded ， with ribs or lacking them; filaments filiform; ovaries and achenes co ト

achene achene hairs 0.2-0.5 mm long; achene styles ered with hairs 0.5-1 mm long.... ・H ・...・ H ・.. glabrous; glabrous; ca 叩el and achene stigmas linear or ...... 16. A. quinquefolia sublinear sublinear ...・ H ・.....・ H ・.....・ H ・.....・ H ・.....・ H ・..….1 5 19b. Lateral leaflets all lanceolate ，undi- 15a. 15a. Basal and involucral leaflets deeply 3- vided; leaves and stems glabrous; filaments lobed; lobed; involucral leaf petioles 10-20 x 3-5 basally dilated; ovaries and achenes covered mm ，distinctly dilated; tepals 8-12 ， having with hairs 0.1 -0.2 mm long.... 3-5 3-5 veins anastomoses; achenes basally ..17. A. lancifolia rounded ， not compressed ，styles 0.5-1 mm 20a. Scapes few ， puberulent; rhizomes long long …...・ H ・.....・ H ・.....・ H ・.12. A. pseudoaltaica mainly vertical; ovaries and achenes densely

15b. 15b. Basal and involucral leaflets medially covered with hairs 1-2 mm long ....・ H ・・- 3-10bed 3-10bed or toothed; involucral leaf petioles ..… 18. A. piperi 5-12 x 1-2 mm ， basally slightly dilated; 20b. Stems solit 紅 y ，glabrous; rhizomes hori- tepals tepals 9-15 ， having 1-3 veins anastomoses; zontal or ascending; ovaries and achene bod- achenes achenes basally narrowed ， slightly com- ies covered with hairs 0.1-0.2 mm long...21 pressed ，styles 1- 1. 5 mm long.... 21a. Tepals 7-10 x 4-6 mm ， mainly white or ..13. ..13. A. raddeana blue; achenes basally narrowed ， slightly 16a. 16a. Involucral leaf petioles 3-5 mm wide; compressed ， with na 町ow lateral ribs... … tepals tepals 5 (-6) ，spreading ， with n征 row bases …… 19. A. grayi and and apices ， 15-30 mm long ， having 5-9 21b. Tepals 10-20 x 5-10 mm ， red to blue; veins veins anastomoses ， sparsely puberulent; achenes basally rounded ， not compressed ， achenes achenes densely puberulent (ser. Nikoenses) without ribs ...・ H ・.....・ H ・H ・H ・.. ..20. A. oregana

...・ H ・...14. A. nikoensis 22a. Rhizomes non-tuberous; basalleaf peti-

16b. 16b. Involucral leaf petioles 1-2 mm wide; oles basally gradually dilated; cymes 1-few 四 tepals tepals 5-7 ，spreading or bent ， with rounded flowered ， tepals 5; achenes ovoid ， bases bases and apices ， 5-20 mm long ， mainly subglabrous or glabrous (sec t. Rosulantes) … without without vein anastomoses ， subglabrous; ...23

achenes achenes sparsely puberulent..... ・H ・....・ H ・-… 17 22b. Rhizomes tuberous; basal leaf petioles 17a. 17a. Cymes few-flowered; bracteoles pre- basally sha 中ly dilated ，scale-like; cymes 1- sent; sent; tepals 5-7 x 1-2 mm ，bent ，sp 紅 sely flowered; tepals 8-12; achenes ellipsoid ， 52 植物研究雑誌第79 巻第1号 平成16 年2月

sparsely sparsely puberulent (sec t. Tuberosa)... ・H ・..27 ribs ，scarcely pubescent... ・.. .26. A. apennina 23a. 23a. Scapes 2-3 ，few-flowered; involucral 28b. Basal leaf blades 3-sected ， with leaf leaf petioles 1-3 cm long; bracteoles present ， petiolules 1-2 mm long (frequently absent); small; small; tepals puberulent ， with or without tepals of outer circle with 5-9 veins vein vein anastomoses; staminodes present; anastomoses; filaments linear; carpels and achenes achenes not compressed ， without ribs (ser. achenes with lateral ribs and pubescent only

Rosulantes) …...・ H ・..…...・ H ・.....・ H ・H ・H ・..…… 24 at the base... …...・ H ・.....・ H ・H ・H ・-・ 27. A. blanda 23b. 23b. Scapes solitary ， 1- flowered; involucral 29a. lnvolucral leaves several (several re- leaf leaf petioles 0.5-1.5 cm long; bracteoles ab- duced); cymes 1-3-flowered; tepals 5-7; sent; sent; tepals subglabrous ， lacking vein achenes sessile; styles hardly recognizable; anastomoses; anastomoses; staminodes absent; achenes stigmas dilated or subcapitate (sec t.

slightly slightly compressed ， with na 町 ow ribs (ser. Stolonifera) ...・ H ・H ・H ・.....・ H ・..…...・ H ・.....・ H ・..30

Exiguae) Exiguae) ...・ H ・H ・H ・.....・ H ・.....・ H ・.....・ H ・..…… 25 29b. lnvolucralleaves 3 ，all reduced; flowers 24a. 24a. Tepals 5-10 mm long ， lacking vein solit 訂 y; tepals 5-13; achenes stalked ，styles anastomoses; anastomoses; staminodes between tepals and distinct ，stigmas linear (sec t. Keiskea) …… 33 ; stamens; stigmas linear …… 21. A. stolonifera 30a. Scale-like basal leaves absent; basal 24b. 24b. Tepals 15-20 mm long ， having 5-15 leaves with distinct blades and basally veins veins anastomoses; staminodes between sta- vaginate petioles developing before anthesis;

mens and c紅 pels; stigmas subcapitate ...・ H ・.. C 訂 pel stigmas subcapitate ， achene stigmas

…・・ 22. A. davidii slightly dilated (ser. Stoloniferae) …...・ H ・..31 25a. 25a. lnvolucral leaf petioles 3-5 mm wide ， 30b. Scale-like basal leaves present; basal basally basally connate; ovaries and achenes leaves with distinct blades and na 町 ow peti- sparsely sparsely puberulen t....・ H ・.....・ H ・.23. A. exigua oles developing after anthesis; ca 中el stigmas 25b. 25b. lnvolucral leaf petioles 1-2 mm wide ， capitate or subcapitate ， achene stigmas free; free; ovaries and achenes glabrous.... ・H ・...26 subcapitate or dilated (ser. Flaccidae) …… 32 26a. 26a. lnvolucral leaf blades temate ， larger 31a. Rhizomes short ， 8-10 mm in diameter ， than than those in basal leaves; styles nearly ab- with long underground stolons; tepals 10-15 sent; sent; stigmas subglobose ...... 24. A. griffithii mm long ， with 3-5 veins anastomoses; 26b. 26b. lnvolucral leaf blades 3-sected ，smaller achene bodies basally narrowed ， slightly than than those in basal leaves; styles distinct; compressed ， with na 町 ow ribs.

stigmas stigmas linear...... ・H ・..…. .25. A. scabriuscula ..29. A. baicalensis 27a. 27a. Rhizomes cylindroid; basalleaf petioles 31b. Rhizomes short ， 5-7 mm in diameter ， 10-20 10-20 cm long ，involucral petioles 1. 5-3 cm with long above-ground stolons; tepals 8-10 long; long; tepals 12-14 ， 15-20 mm long; achenes mm long ， without vein anastomoses; achene 3.2-3.6 3.2-3.6 mm long ，its ribs ca. 0.2 mm or ab- bodies basally rounded ， not compressed ，

sent; sent; stigmas linear …...・ H ・.....・ H ・.....・ H ・-… .28 without ribs ...・ H ・.....・ H ・.....・ H ・.. ..30. A. prattii 27b. 27b. Rh izomes subspherical; basalleaf peti- 32a. Basalleaves having blades 2 -4; scapes oles oles 5-8 cm long ，involucral petioles 0.5-1.5 1-3 ，cymes few-flowered; bracteoles pre- cm long; tepals 8-10 ， 8-10 mm long; sent; tepals 5-10 mm long ， with 7-9 veins achenes achenes 2-2.2 mm long ， with ribs ca. 0.5 anastomoses; achenes sp 訂 sely puberulent or mm and slightly urved styles; stigmas dilated glabrate ，hairs 0.1 mm long...3 1. A. flaccida … 28. A. caucasica 32b. Basal leaves having blades solitary; 28a. 28a. Basalleaf blades temate ， with petiolules scapes solit ぽ y，cymes 1- flowered; bracteoles 3-5 3-5 mm long; tepals of outer circle with 1- absent; tepals 7-15 mm long ， lacking vein 3 vein anastomoses; filaments basally di- anastomoses; achenes densely puberulent ， lated; lated; c紅 pels and achenes without lateral hairs ca. 0.5 mm long...... ・H ・.32. A. delavayi February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 53

33a. 33a. Rhizomes 1-2 mm in diameter; tepals Anemone nemoros αL. subsp. europaea 10-22 ，linear-lanceolate ， having few veins Ulb r. in Bo t. Jahrb. 37: 225 (1 906). anastomoses ， basally sparsely puberulent; Anemone nemorosa L. subsp. かpica achene achene stalks 1.5-2 mm long ， achene bodies Korsh. in F1. Vos t. Evr. Ross. 1: 59 (1 892). long-obovate ， sparsely puberulent ， stigmas Anemone nemorosa L. var. purpurea et linear-lanceolate linear-lanceolate (ser. Keiskea) … V 紅 . rosea DC. ，Prodr. 1: 20 (1 824). …・ 33. A. keiskeana Anemone nemoros αL. va r. coerulea Pritz. 33b. 33b. Rhizomes 2-5 mm in diameter; tepals in Linnaea 15: 652 (1 841). Syntype: “ Prope 5， elliptic ， without vein anastomoses ， Dachland ad Rh enum" ， Zeyher M. glabrous; glabrous; achene stalks 0.5-0.7 mm long ， Anemone nemorosa L. var. hirsuta achene achene bodies subglobose ， hispid proxi- Wierzbicki ex Pritz. ，1. c. (1 841). Type: mally ， stigmas triangular-subulate (ser. “Mons Magdalenae prope Brixiam It aliae" ，

Deltoidea) Deltoidea) ...・ H ・H ・H ・..…...・ H ・. .34. A. deltoidea Brach t. Anemone nemorosa L. v訂 . parviflora Taxonomic analysis Schlechtd. ex Pritz. ，1. c. (1 841). Characteristics Characteristics of all taxa are given below ， Anemone nemorosa L. var. rubra Pritz. ，1. followed followed by a discussion of their interrela- c、(1 84 1). tionships. tionships. We excluded from the descriptions Anemone nemorosa L. var. semiplena [except [except the first one within the each sub- Pritz. ，1. c. (1841). group group (section or series)] the characters in Anemone nemorosa L. var. sulj フhurea common to all taxa ，viz. ， the characteristics Pritz. ，1. c. (1 841). of of the scale-like basal leaves ， whether dis- Anemone nemorosa L. var. latiloba tinct tinct basal leaf blades develop after anthesis ， J. Kickx f. in Bull. Soc. Roy. Bo t. Belg. 4: the the 3 involucral leaves ，monomorphic tepals ， 204 (1 865). rounded rounded apices of anthers ca. 1 mm long ， Anemone nemorosa L. var. anandra Hua conic conic glabrous carpel and achene styles ， in Bull. Soc. Bo t. Fr. 26: 255 (1 889). subglobose subglobose achene heads ca. 1 cm in diame- Anemone nemorosa L. var. bastardii Rouy ter ，neither do we mention the various types & Foucaud ， F 1. Fr. 1: 44 (1 893). (holo- ，lecto- and others) ，if they 紅 e not des- Anemone nemorosa L. var. grandiflora ignated. ignated. Rouy & Foucaud ，1. c. (1 893). Anemone nemorosa L. va r. tenuiflora Rouy Subgen. Subgen. 1. Anemonanthea. & Foucaud ，1. c. (1 893). Sec t. 1. Anemonanthea DC. Anemone nemorosa L. var. bracteata Hegi Ser. Ser. 1. Anemonanthea. in 11 1. F1. Mitte 1. Eur. 3: 524 (1 912). 1. 1. Anemone nemorosa L. ， Sp. P1. 541 Anemone nemorosa L. var. monoica Hegi ， (1 753). 1. c. (1912). TYPES: “F1. suec. 450. Anemone nemorosa L. var. cyanopsis sylvarum. sylvarum. Habitat in Europaea asperis ，duris ， Lagerh. in Svensk Bo t.“ insula Varmdo nemoribus". nemoribus". N 710.15 (l ectotype designated prope Stockholmiam ，1896 ， Netzel Suniv". here-LINN!). “W : Herb. Clifford Anemone nemorosa L. var.α rpetala Salisb. 224 ，Anemone 3， Ranunculus phragmites in Ann. Bo t. (Oxford) 30: 528 (1916). albis" albis" (BM!). Holotype: “ Great Britain. Oak-Hombeam Anemone nemorosa-alba Crantz ， Stirp. woods of Hertfordshire" ， Worsdell (K!). Austr. Austr. Fasc. 1-2: 101 (1762). Anemone nemorosa L. var. robusta Anemone alba Gilib. ， F 1. Lith. 2: 275 Salisb. ，1. c. (1 916). (1 782). Anemone nemorosa L. f. b抑ora Peterm. ， 54 植物研究雑誌第79 巻第1号 平成16 年2月

l. Fl. Lips. 407 (1 838). densely puberulent (hairs 0.1-0.2 mm long); Anemone nemorosa L. f. latisect αSchur ，f. styles apically curved ，0.5-1.2 mm long ， subumbellata Schur in Enum. Pl. Transs. 4 sparselypuberulent or glabrous; stigmas (1 866). slightly dilated (Fig. 3A). Anemone nemoros αL. f. lat ザolia Ulbr. in Chromosome number: n = 15 ，16 (Baum- Bo t. Jahrb. 37: 215 (1906). berger 1970). (L.) Holub in Distribution: Europe (most countries in- Folia Folia Geobo t. Phytotax. Praha 8: 166 (1973). cluding Great Britain ， Germany ， Austria ， Rhizomes pros 住ate or ascending ， 3-5 mm It aly ， Poland ， Ukraine ， etc.); occurring in in in diameter ， long and branched. Basalleaves forests and under bushes ，but sometimes in 2-3 ，scale-like (4-6 x 5-7 mm) ，solitary with open sites ， al t. 300-1500 m. distinct distinct blades developing after anthesis; Specimens examined: UKRAINE; Ki ev Reg. ， petioles petioles 5-20 cm long ， glabrous or sp 紅 sely Boyarka ， 12 .4 .1915 ， Semenkewich (KW); puberulent; puberulent; blades temate ，pentagonal ，2- Nemeshaev ， 7.4 .1916 ， Semenkewich (KW); near 6x 2-5 cm ， glabrous or sparsely puberulent Syrets ， 23 .4 .1922 ， Semenkewich (KW); Klavdiev ， 18 .4 .1927 ，Semenkewich (KW); Trubetska ，1. 5.1924 ， along along veins; petiolules 3-5 mm long; central Lazarenko (KW); Feofania ， 10.5.1998 ， Ziman & leaflets leaflets 3-10bed ， ellipsoid-oblong; bases Bulakh (KW); Kiev ， near National Botanical Garden ， wedge-like; wedge-like; margins se 町 ate and incised ，api- 20 .4 .1999 ， Bulakh (KW); Zhitomir Reg. ， Ovruch ces ces acuminate; lateral leaflets deeply 2- Distr. ，Kolesnyki ，12 .4 .1971 ， Barbarich (KW); Olevski lobed ， ovate to slightly deltoid ， having ser- Distr. ， Sushchyni ， 25.5.1948 ，Dobrochaeva (KW); Kh melnitski Reg. ， Chemerovets Dist r.， Karachkovtsy ， rate-incised rate-incised margins. Scapes 5-20 (-30) cm 18 .4 .1968 ，Morozyuk (KW); Lviv Reg. ，Turka Distr. ， long ， glabrous or pilose; cymes 1-flowered. Ratsyni ，5.5.1985 ，Kr y司(KW); Rivne Reg. ，Verbetsky Involucralleaves Involucralleaves 3; petioles flat and dilated ， Dist r.， Dubki ，26 .4 .1954 ，Kotov (KW); Winnitsa Reg. ， 1-3 1-3 cm x 3-5 mm ， glabrous or sparsely pu- Pyatnichany ，24 .4 .1939 ， Grynj (KW); Ternopil Reg. ， bescent; bescent; blades temate ， similar to those in Bozha M t. near Kr emenets ，10.6.1998 ，Ziman (KW); basal basal leaves ， 2-5 x 2-5 cm ， sparsely Chernivtsy Reg. ， Vyzhnitski Dist r.， Lopushno ， 11. 4.1961 ， Solodkova (KW); Ivano-Frankovsk Reg. ， puberulent puberulent along veins; petiolules 2-5 mm Pereginski Dist r.， Osmoloda ，25 .4 .1941 ， Grynj (KW); long; long; central leaflets 3-10bed; lateral leaflets near Vorokhta ， M t. Pozhizhewska ，30.6.1997 ，Ziman deeply deeply 2-10bed or 2-parted (Fig. lA). & Dutton (KW); Transcarpathia Reg.: Rakhiv Dist r.， Pedicels Pedicels 2-6 (-9) cm long ，pilose. Tepals 6- Bogdan ， M t. Goverla ，1. 7.1948 ，Bylyk (KW); Jasinja ， 10 ，oblong-ovate ， white or tinged red to blue ， M t. Bliznitsa ，summit ， 1800 m ，28.6.1997 ，Ziman & Dutton Dutton (KW); Svaljava Dist r.， close PO りana ， rose rose or pink ，glabrous ， often dimorphic: 30.5.1998 ， Ziman (KW). HUNGARY; Gomor ， outer outer tepals 10-20 x 5-10 mm ， with 5-9 Tiszolcz ， Furmaniecz vogy ， 25.5.1900 ， Richter basal basal veins and 3-5 vein anastomoses; inner (CLUJ); Kr istians M t.， Nysaeter ， 8.6.1907 ， Fridtz tepals tepals 8-12 x 3-6 cm ， with 4-5 basal veins (CLUJ). ROMANIA; Muntenia Mts. ，Bucegi ， 1500 m ， and solitary vein anastomoses. Stamens 4- 8 6.8 .1 938 ，Nyarady (CLUJ); Muntenia Distr. ，Prahova ， mm long; filaments mainly filiform ， but Galma ， prope pagum Busteni ，950 m ，28 .4 .1940 ， Cretzoiu Cretzoiu (MHA); Oltenia ， Vilcea Distr. ，Govora ，500 sometimes sometimes basally dilated; anthers oblong- m ，17 .4 .1967 ， Popescu (MHA); Satu Mare ，4 .4 .1976 ， ellipsoid ， with rounded apices and wide con- Pop (CLUJ); Sibiu ， Talmacin Suvara ，410 m ，1 1. 4. nectives ，ca. 1 mm long. Ovaries ovoid ，0.7- 1980 ， Dragulescu (CLUJ). POLAND; Gda 吋 Distr. ， 1.5 1.5 mm long ， densely puberulent (hairs 0.1- Dulowa Chrzanow ，29 .4 .1952 ，Novak (MHA); Babyi 0.2 0.2 mm long); styles conic ， almost straight ， Dol ， near Mukovo ， 17.6.1978 ，Matsenko (MHA). CZECHIA; Silesia ， Slask Dolny ， Brzeznica pow. ， 0.5 -0 .8 mm long; stigmas slightly dilated Zabkowice ， 20 .4.1 952 ， Borzeta (MHA); Bohemia ， (Fig. (Fig. 2A). Achene bodies ovoid to ellipsoid ， Kunratice ，3.5.1958 ， Nitka (MHA). GERMANY; N 3.5-5 3.5-5 x 1.5-2.5 mm ，slightly compressed ， Schwielow-See ， Mark ， 10.5.1924 ， Laubert 1031 with with narrow lateral ribs (0 .1-0 .2 mm wide) ， (MHA); Dollerupholz ，Gluksburg ，14.5.1970 ， Larsen February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 55

(CLUJ); (CLUJ); Halle-Saale ， Petersberg ， 22 .4 .1980 ， Anemone nemorosa L. var.β ssa Ulbr. in Skwortsow (MHA); Harz-Gebirge ， Harzgerode ， Bo t. Jahrb. 37: 226 ， 1906. Type: CHINA: 29 .4.1 980 ， Skwortsow (MHA). DENMARK; “Sudostmandschurei: am oberen U ssuri" ， Yutlandia ，prope 可ele ，18 .4 .1969 ，Skwortsow (MHA); 1860 ，Maximowicz (LE!). Zealand ， Grib Skov ，Kagerup ，15.5.1970 ，Fredtoft & Jacobsen Jacobsen (MHA). ; S Tavastia ，Tyrvanto ， Anemonoides amurensis (Korsh.) Holub in Lepaa ， 8.5.1959 ， Pankakoski (MHA). GREAT Folia Geobo t. Phytotax. Praha 8: 165 (1 973). BRITAIN; Berkshire ， Reading ， Whiteknights ， Rhizomes prostrate or ascending ， 3-10 Wildemess ，24 .4 .1978 ， Souster (MHA). ITALY; Prov. cm x 1-3 mm ，long ，branched. Basal leaves Riet ， Lazio ， M t. Terminillo ， 1300 m ， 19.5 .1 987 ， 2-3 ， scale-like ， and solitary ， with distinct Zucconi Zucconi et al. (MHA). SP AIN; Prov. Leon ，Cordillera blades ，developing after anthesis; petioles 7- Cantabrica ， Pinar de Lillo ， 1550 m ，3.6.1980 ，Losa- Quintana Quintana (MHA). 15 (-20) cm long ，basally slightly dilated ， According According to Juzepchuk (1 937) ， Chater sparsely pilose or subglabrous; blades 1- or (1 973) ，Pignatti (1 988) and others ， the most 2-temate ，pentagonal ， 3-3.5 x 5-6 cm ，aba- important important characters of A. nemorosa are the xially glabrous ，adaxially sparsely puberu- temate temate basal and involucral leaves ，solitary lent; petiolules 5-10 (一 15) mm long; central flowers flowers with 5-7 glabrous tepals and oblong leaflets 3-sected or 3-parted ， rhombic to achenes achenes with curved styles. broadlyovate; bases subcordate to cuneate; From the time of De Candolle (1817) and margins crenate or dentate; apices acuminate Pritzel Pritzel (1 841) ， A. nemorosa was regarded as to obtuse; lateral leaflets similar to central variable a variable taxon which included several sub- leaflet ， rhombic to obovate ， but oblique; cen- species species and varieties (e.g. ，var. rubra Pritz. ， tral segments deeply 3-10bed ，rhombic ，lat- var. var. grandiflora Rouy & Foucaud ，v 紅. eral segments usually 2-clef t. Scapes (5-) bracteata bracteata Hegi). Most of these varieties were 10-25 cm long ，glabrous; cymes 1-flowered. described described on the basis of tepal size and Involucralleaf petioles 1- 1. 5 cm x 3-5 mm ， colour colour and other variable characters. dilated and narrowly winged ，pilose; blades Therefore ，we confirm the opinion of Dutton ternate ， ovate to pentagonal ，2.5 -4 x 2.5 -4 (1996) (1996) who noted the continuous nature of cm ， subglabrous or adaxially sparsely the the character variation throughout the range puberulent; petiolules 2-5 mm long; central of of this species which makes recognition of leaflets subpinnately parted ，ovate-rhombic; infraspecific infraspecific entities unwarranted. bases cuneate; margins se 町 ate; apices acuminate; acuminate; lateral leaflets 2-parted ， ovate to 2. 2. Anemone amurensis (Korsh.) Kom. in lanceolate (Fig. 1B). Pedicels 1. 5-5 cm long ， Tr. Tr. Imp. St. -Peterburgs k. Bo t. Sada 22: 262 puberulen t. Tepals 5-8 ， monomorphic ， (1 903). obovate to oblong ， with long-narrow bases TYPES: RUSSIA: “ In sylvis frondosis ad and rounded apices ，white ， 10-15 (-18) x 5- ostium ostium U ssuri prope Kazakewiczewa ヘ 7 (-10) mm ，glabrous; basal veins 3-5 ， vein 15.5.1891 ， Korzhinski (l ectotype-LE! ; anastomoses solitary. Stamens 4-7 mm long; isotype-GH! isotype-GH! ). filaments filiform or sometimes slightly di- Anemone nemorosa L. subsp. amurensis lated basally and apically; anthers ellipsoid ， Korsh. Korsh. in Tr. Imp. St. -Peterburs k. Bo t. Sada connectives wide. Ovaries cylindroid ， 2-2.5 12: 12: 293 (1 892). mm long ， densely puberulent ，hairs ca. 0.2 Anemone nemoros αL. var. kamtschatica mm long; styles straight ， 0.8-1 mm long; Kom. ， F l. Pen. Kamtsch. 2: 129 (1 929). stigmas subcapitate (Fig. 2B). Achene bodies Anemone nemorosa L. subsp. kamtscha- narrowly ovoid ， 4-5 x 1. 5-2 mm ， densely tica tica (Kom.) Starod. in Bo t. Zhum. 68: 1015 puberulent (hairs 0.1-0.2 mm long); styles (1 983). slightly curved ， 0.5-1 mm long ，glabrous; 56 植物研究雑誌第79 巻第1号 平成16 年2月 stigmas stigmas sublinear. (holotype-G). Isotypes: “ In montibus Chromosome number: n = 24 (Starodubt- altaicus frequenter in pratis humidis sev sev 1982 ， Nishikawa 1997). montosis et ad nives deliquescentes 人 Distribution: Distribution: NE : Russia (Far East ， Ledebour & Bunge. “ In pratis ad rivulum Kamchatka) ，NE China (J ilin ，Heilongjiang ， Urmuchaika prope Buchtarminsk 人 Meyer Liaoning) ，N Japan (Hokkaido) ，N Korea; (paratypes-LE!) . occurring occurring in forests or under bushes ， mainly Anemone fisheriana DC. ， Prodr. 1: 20 ln ln mountmn regions. (1 824). RUSSIA: “ In Sibiria ， prope Salair 人 Specimens Specimens examined: RUSSIA; Far East ，Primorje ， Fischer (holotype-LE!). near near Vladivostok ， 26 .4 .1902 ， Palczewski (KW); Anemone incisa Ledeb. ， Ind. Sem. Hort. 26.3.1906 ， Palczewski (LE); 3.5.1910 ， Kuznetsov Dorp. (1 824). (LE); (LE); 17.5.1910 ， Kuznetsov (KW); 4.5.1970 ，Gorovoy Anemone ranunculoides L subsp. caerulea (MHA; KW); 5.5.1980 ， Starodubtsev (MHA); Shkotovski Shkotovski Distr. ， Zmeinka ， 9.5.1973 ， Kostenko (DC.) Korsh. ， F 1. Vos t. Evr. Ross. 54 (1 892). (VLA); (VLA); Tygrovyi ，29 .4 .1980 ，Starodubtsev (MHA); Anemone caerulea DC. subsp.η pica Ulbr. Ussuri Ussuri Reserve ， Maikhinski Kluch ， 14.5.1973 ， in Bo t. Jahrb. 37: 217 (1 906). Kostenko Kostenko (VLA); 3.5.1974 ， Kharkewitch & Buch Anemonoides caerulea (DC.) Holub in (MHA); Kh abarovski Reg. ， Khasanski Distr. ， Folia Geobo t. Phytotax. Praha 8: 166 (1 973). Furugelma Furugelma Is 1.， 5.1975 ， Zdorovjeva (MHA); Kedrovaya ，9.5.1980 ， Starodubtsev (MHA); Il jinski Anemonoides fischeriana (DC.) Holub ，1. Distr. ， near Evgenievka ，Kh anka Lake ，3 1. 3.1911 ， c. (1973). Cherski Cherski (LE); near Tatarski Strait ，5.1912 ，Derbek Rhizomes pros 凶 te ，long ，branching ，2 -4 (LE); (LE); ChuguevDistr. ， Sokolovka ， 19.5.1980 ， mm in diameter. Basalleaves 2-3 ，scale-like ， Starodubtsev Starodubtsev (MHA); c1 0se Kr asnorechensk ，2.5.1983 ， and solitary ， with distinct blades; petioles 8- Shestera Shestera (MHA); Kamchatka ， Abachinska ，guba 12 12 (-15) cm long ，glabrous; blades 3-sected ， Tarvinska Tarvinska Bay ，7.6.1908 ，Komarov (LE). JAPAN; Hokkaido ， Tokachi Prov. ， Urahoro ， 21. 4.1985 ， rhombic-pentagonal ，2 -4 x3 -4 cm ，glabrous; Nishikawa Nishikawa (TNS). petiolules 1-2 mm long; central segments Korzhinski Korzhinski (1 892) initially described this narrowly oblong ， undivided or 3-1obed; species species as a subspecies under A. nemorosa bases narrowly cuneate; margins se 町 ate; api-

(as (as subsp. amurensis). However ，Komarov ces long-acuminate; lateral segments 2幽 (1 903) regarded it as a distinct species (A. P訂 ted ，oblique ， with 4- 10 lacerate lobes. amurensis) amurensis) because of its wide-dilated Scapes 15-25 (-35) cm long ， glabrous; involucral involucral leaf petioles. Juzepchuk (1937) cymes few 四 flowered. Involucral leaf petioles also also noted its longer basal leaf petiolules ， 3-10 x 3-5 mm ， sparsely puberulent or smaller smaller flowers ， and longer stamens. glabrate; blades 3-sected ， larger than the In In this treatment A. α murensis differs from basal leaf blades ，4- 6x 5-6 cm ，pilose be- A. A. nemorosa mainly by shape of the basal neath; segments subsessile; central segments leaf leaf leaflets (3-sected or 3-parted ，rhombic ， oblong or linear-lanceolate; bases narrowly with with subcordate bases) and monomorphic cuneate; margins coarsely serrate or incised- tepals tepals lacking vein anastomoses but with dentate ， 3-5 long-acute teeth distally ， long-narrow long-narrow bases. However ， both taxa (not falcate; apices acuminate; lateral segments 2- 紅白 only only A. amurensis) 訂 e characterized by di 園 P d， having several acute teeth (Fig. 1 C). lated lated invlucral leaf petioles and glabrous Pedicels 1-2 (-5) cm long ，appressed- or tepals tepals of a similar size. erect-spreading-villous. Tepals 5 (-6) ，ob- ovate-elliptic ， with n紅 row bases and wide 3. 3. Anemone caerulea DC. ，Sys t. Na t. apices ， mainly blue or rarely whitish ，6- 1: 1: 203 (1 817). 10 x4 ー7 mm ，appressed-pilose; basal veins TYPES: “ In Sibiria circa Zmeof ，"， Patrin 5-7 ， vein anastomoses solitary. Stamens 3- February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 57

5 mm long; filaments basally dilated; anthers Candolle (1 824) also described A. ellipsoid ，connectives wide. Ovaries ovoid- fischeriana a closely related species ， but ellipsoid ， basally and apically narrowed ， which later was included in A. caerulea. We densely densely puberulent ，hairs 0.1-0.2 mm long; confirm that the distinctions of A. styles styles uncinate ， 0.7-1 mm long; stigmas fischeriana from A. caerulea are too slight to

slightly slightly dilated (Fig. 2C). Achene bodies warrant recognizing A. fischeriana as a dis 由 ovoid-ellipsoid ， 1-1.5 x 1-1.5 mm ， densely tmct specles. puberulent puberulent (hairs ca. 0.2 mm long); styles straight straight or curved ， 1-1.5 mm long ，glabrous; 4. Anemone uralensis Fisch. ex DC. ， stigmas stigmas sublinear. Prodr. 1: 19 (1 824). Chromosome number: n = 8 (Baumberger TYPE: “ In montibus Uralensibus" ， 1970 ， Kartashowa et al. 1974). Fischer (holotype-G). Distribution: Distribution: NE Asia: Russia (Siberia); Anemone ranunculoides L. subsp. occurring occurring in forests and their m 紅 gins ， ur α， lensis (DC.) Korsh. ， F l. Vos t. Evr. Ross. scrubs ， and occasionally in meadows. 1: 57 (1 892). Specimens Specimens examined: RUSSIA; Tomsk ，9.5.1903 ， Anemone caerulea DC. var. uralensis Nekrasova (KW); ne 町 Tomsk ，2.5.1923 ， Kry lov (DC.) Korsh. ，Ten t. Fl. Ross. Orien t. 6 (KW); Tomsk Distr. ， Basand 司ka ， 17.6.1925 ， (1 898). Sergievskaya Sergievskaya (KW); Tomsk Reg. ， 11. 5.1939 ， Anemone caerulea DC. var. uralensis Bashmakova (KW); Novosibirsk Reg. ， Maslyaninski Distr ，Otchikha ，2 1. 5.1967 ，Kr asnoborov (KW). (DC.) Ulbr. in Bo t. Jahrb. 37: 266 (1 906). Anemone caerulea was described by De RUSSIA: “S Ural ，im Ufa bei Zlatoust ，iil Candolle Candolle (1 817) ，and ， according to the Perm bei Kr assnoufimsk ， an der Tchusovaya" ， author' author' s description ，its plants have ternate Georgi 1773 (l ectotype-LE!). long-petiolate long-petiolate basal leaves ，involucralleaves Anemonoides uralensis (DC.) Holub in with with short petioles ， 5 elliptic tepals ， and Folia Geobo t. Phytotax. Praha 8: 166 (1 973). ovoid ovoid sessile achenes. Rhizomes prostrate ，long ，branching ，2 -4 However ，Korzhinski (1 892) regarded this mm in diameter. Basalleaves 2-3 ，scale-like ， taxon taxon as a subspecies of A. ranunculoides ， or and solitary ， with distinct blades; petioles 7- in in the case of Ulbrich (1906) ，a species very 15 cm long ，glabrous; blades 3・・ sected ，2- close close to it. Ulbrich regarded both of them as 4x 2 -4 cm ，glabrous; petiolules 1-2 mm species “species collectiva". Korzhinski (1 892) and long; central segments ovate to rhombic ， Ulbrich Ulbrich (1 906) both noted the cylindric with 2-4 lacerate lobes; bases n訂 row- shape shape of the rhizomes ， the long-acuminate lycuneate; m 紅 gins se 町 ate; apices acuminate; terminal terminal leaf lobes ，1-2- flowered glabrous lateral segments similar to central ones ， stems ， and the rose or blue tepals. lanceolate ， with 4- 8 lacerate lobes. Scapes Nevertheless ， the view that A. caerule αis a 8-20 cm long ，glabrous; cymes 1-2-flowere species species allied to A. ranunculoides did not d. Involucral leaf petioles 3-5 x 3-5 mm ， change. change. glabrous or puberulent; blades 3-sected ， According According to our data ， A. caerulea is quite larger than those in basalleaves ， 3.5-6 x 3- close close to A. ranunculoides (short involucral 5 cm; petiolules 1-2 mm long; central seg 四 leaf leaf petioles and leaflet petiolules ，few- ments narrow-elliptic; bases cuneate; m 紅- flowered flowered cymes ， pubescent tepals with vein gins se 町 ate ， with 4-9 teeth; apices anastomoses). anastomoses). In our opinion ，however ，it is acuminate; lateral segments similar to central closest closest to A. uralensis (differs only by the ones. Pedicels 2-5 cm long ， or pilose villous. tepal tepal shape ， colour and haimess ， and fila- Tepals 5 (-6)， obovate-elliptic ， with wide ment shape). bases and apices ， mainly rose or red ，some- 58 58 植物研究雑誌第79 巻第1号 平成16 年2月 times times yellow ， blue or white ，6-12 x 4-7 mm ， europaea Korsh. ， F l. Vos t. Evr. Ross. 56 puberulent puberulent proximally or medially; basal (1 892). veins veins 5-7 ， vein anastomoses 3-5. Stamens Anemone ranunculoides L. subsp. 2-5 mm long; filaments apically dilated; an- jenisseensis (Korsh.) Kry lov & Steinb. in Tr. thers thers ellipsoid ，connectives wide (sometimes Bo t. Muz. Ros. Akad. Nauk 17: 60 (1 918). stamens stamens lacking anthers). Ovaries ovoid ， Anemone ranunculoides L. subsp. 0.7-1 0.7-1 mm long ， densely puberulent ， hairs wockeana Ashers. & Graebn. ， F l. Nordos t. 0.01-0.05 0.01-0.05 mm long; styles slightly curved ， Deutsch. Flach l. 332 (1 898). 0.6-1 0.6-1 mm long; stigmas slightly dilated (Fig. Anemone ranunculoides L. vars. latisecta 2D). 2D). Achene bodies subglobose ， 1- 1. 5x Schur ， subumbellata Schur ， Enum. Pl. 0.5-1.5 0.5-1.5 mm ， densely puberulent (hairs less Transsilv. 4 (1 886). than than 0.1 mm long); styles recurved ，ca. 1 mm Anemone ranunculides L. var. latifolia long ，glabrous; stigmas sublinear. Ulb r. in Bo t. Jahrb. 37: 215 (1 906). Chromosome number: n = 16 (Chupov Anemonanthea ranunculoides (L.) A. 1975). 1975). Gray in Na t. Arr. Bri t. Pl. 2: 725 (1821). Distribution: Distribution: Russia (Ural); occurring in Anemonoides ranunculoides (L.) Holub in meadows and scrubs ， in mountains. Folia Geobo t. Phytotax. Praha 11: 81 (1 973). Specimens Specimens examined: RUSSIA; Ekaterinburgh Rhizomes long and branched ，prostrate to Reg. ，C Ura l. M t. Volchikha ，25.5.1966 ， Storozheva ascending ， 2-5 mm in diamete r. Basalleaves (MHA); Didina ， 21. 5.1977 ， Storozheva (MHA); 2-3 ，scale-like ，and solit 紅 y， with distinct Chusova River ， M t. Kamenj Georgievski ，2 1. 5.1979 ， blades; blades; petioles 4-20 cm long ， glabrous; Storozheva Storozheva (MHA). According According to the author's description ， A. blades temate to bitemate ，2 -4 x2 -4 cm ， uralensis uralensis is characterized by ternate short- adaxially glabrous or sparsely pilose ， petiolate petiolate involucralleaves with linear termi- abaxially glabrous; petiolules 2-5 mm long; nal nal lobes and 5-6 oblong tepals. Pritzel central lea f1 ets 3・lobed ， ovate to rhombic; bases bases cuneate; margins crenate to se 町 ate ， (1 841) reg 紅 ded this species as close to A. caerulea ， and Juzepchuk (1 937) to A. apices acuminate; lateral segments 2-p 紅白d ranunculoides. ranunculoides. or deeply 2-10bed ， leaf petioles 3-5 cm x 1- This This species is closest to A. caerule α(as a 2 mm ， glabrous or sparsely pilose; blades sister sister species) and in addition to tepal char- temate ，3-6 x2 -4 cm ，adaxially glabrous or acteristics acteristics mentioned above ，it differs from sparsely pilose along veins; petiolules 2-5 the the latter by its restricted geographical area mm long; centrallea f1 ets 3-parted to 3-10bed ， (an (an endemic of the Urals) ，and also by num- ovate to rhombic ， bases cuneate ， margins ber ber of chromosomes (tetraploid). crenate-se 町 ate ， apices acute; lateral lea f1 ets similar similar to central ones ， but 2-parted (Fig. 5. 5. Anemone ranunculoids L. ， Sp. Pl. 541 1D). Pedicels 2-5 cm long ，pilose. Tepals 5- (1 753). 8 (-10) ， broadly ovate to obovate ，12-20 x TYPE: “ Ranunculus nemorosus luteus. 5-10 mm ，yellow ， with narrow bases and Habitat Habitat in Europe borealis pratis nemorosis ぺ rounded apices ， dimorphic: outer ones N 710.30. (l ectotype designated here-LINN!). abaxially densely pubescent ， basal veins 5- Anemone nemorosa-lutea Crantz ， Stirp. 7， vein anastomoses 1-2; inner ones sp 紅 sely Austr. Austr. 1-2: 101 (1 762). puberulent along central vein ， basal veins 2- 3 without anastomoses. Stamens 4- 6 mm Anemone flava Gilib ， F l. Li 由. 2: 274 ， (1 781). long; filaments apically slightly dilated; an- Anemone ranunculoides L. subsp. thers ellipsoid ， connectives wide. Ovaries ovoid ， 0.6-1.5 mm long ， basally narrowed ， February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 59 densely densely puberulent ，hairs 0.1-0.3 mm long; ITAL Y: Prov. Roma. Lazio. Castelli Romani. Velletri styles styles straight or curved ， 0.8-1 mm long; 27 .4 .1993 ， Milloza et al. 15993 (MHA); SPAIN; stigmas stigmas subcapitate (Fig. 2E). Achene bodies Alava ，Sierra de Entzia ，Legumba ，23 .4.1 989 ，Uribe- Echebarria Echebarria & Urrutia 206 (MHA). ovoid ovoid to ellipsoid ，slightly compressed ， with The authors who worked with A. narrow narrow ribs ， 2-4 x 1.5-2.5 mm ， densely ranunculoides ranunculoides in the local floras (Chater puberulent puberulent (hairs 0.2 -0 .3 mm long); styles 1973 1973 and others) noted its temate basal and curved ，0.8-1.5 mm long ，glabrous; stigmas involucral involucral leaves (the latter with short peti- linear linear (Fig. 3B). oles) ，bipartite lateral leaf segments ，2 -4 Chromosome number: n = 8， 24 (Chupov flowers flowers with 6-9 pubescent tepals and ob- 1975); 1975); 16 (Baumberger 1970). long long pubescent c紅 pels. Juzepchuk (1937) Distribution: Distribution: Europe (a lot of countries) ， noted noted the presence of scale-like leaves on Caucasus ， Asia Minor (Turkey); occurring in creeping creeping rhizomes ，solitary basalleaves with open open forests or shady slopes. ternate ternate blades (sometimes absent) ， the few- Specimens Specimens examined: UKRAINE; Kiev Reg. ， Goloseevo Goloseevo close Kiev ， 12 .4 .1920 ， Oksijuk (KW); flowered cymes ， yellow pubescent tepals ， 16 .4.1 923 ，Dubovik (KW); 10.5.1928 ， Polonskaya short stamens and comparatively large (KW); 24 .4 .1946 ， Majevskaya (KW); Lysa Gora near achenes with short curved styles. Kiev ， 30 .4.1 977 ， Klokov (KW); Bila Tserkva ， According to our data ， the main distinc- 29 .4 .1923 ， Grodzinski (KW); Bucha ， 1. 5.1928 ， tions of A. ranunculoides ftom A. nemorosa Polonskaya Polonskaya (KW); Borodyanka ， Babka ， 9.5.1980 ， include include short narrow involucral leaf petioles ， Andrienko Andrienko (KW); Kh arkov Reg. ，Pomerky ，9 .4 .1915 ， Kotov (KW); Sokolnyki near Kh訂 kov ，17 .4 .1922 ， non-solitary flowers ， and yellow pubescent Kozlov Kozlov (KW); Zhuravlevka ，3.5.1938 ， Kotov (KW); tepals with few vein anastomoses. In view of 3.5 .1 938 ， Mykhailenko & Osadcha (KW); Cherkasy the morphology of the flowers and achenes ， Reg. ， near Smela ，Sunky ，4 .4 .1897 ， Rakochi (KW); this species is closer to A. trifolia but differs Jablunivka ，Repyashne ，25 .4 .1924 ， Kleopov (KW); from it mainly by its monomorphic perianth ， Poltava Poltava Reg. ， Vorskla River ，Kyshe 吋ka ，9.4.1 897 ， Zinger Zinger (KW); Khorol ，Stezhky ，30 .4 .1924 ， Polonskaya shorter achene styles ， basal and involucral (KW); Sumy Reg. ， Miropolski Distr. ， Junakovka ， leaf shape (2-3-1obed or parted leaflets with 9.6.1954 ，Gryny (KW); Chernigov Reg. ， Ignyanski mainly crenate margins ， longer involucral Distr. ，Kachanovka ，3 .4 .1957 ，Gry 吋 (KW); Donetsk leaf petioles and smaller blades). Reg. ， Avdeevski Distr. ， Jakovlevka ， 19 .4 .1962 ， The variability of many morphometric Dubovik (KW); Amvrosievski Distr. ，Novo-Petrovka ， characters of A. ranunculoides was the basis 24 .4 .1962 ， Dubovik (KW); Zakarpatski Reg. ， U zhgorodski Dist r.， Antalovetska Polyana ，10.5.1956 ， for describing several interspecific taxa [viz. ， Chopik Chopik (KW); Chernivtsi Reg. ，Tsezino ，15 .4 .1961 ， subsp. jenisseensis Korsh. (1 892) ， va r. Gorokhova et al. (KW); Estonia; Polivere ，Haaprala ， latifolia Ulb r. (1 906) ， var. subumbellata 14.5 .1 962 ，Aasamaa 20152 (KW); M 訂 tra ，17.5.1962 ， Schur (1 886) ，etc.] ， but in the present treat- Aasamaa 20144 (KW); Dakvere ，Kadrina ，close village ment these 訂 'e not accepted as subspecific Vahnja ， 25.5.1962 ， Aasamaa 20131 (KW); ROMANIA; Banatus ，Timis ， Baile Lipova ，23 .4 .1943 ， taxa. Borza Borza (CLUJ); Poidureo Taucapesti ， Mitropolie Vladiceni ，13 .4 .1955 ， Cerbouiscu (CLUJ); CZECHIA; 6. Anemone udensis Trautv. & C.A.Mey. ， Bohemia ， Kutna Hora ，Solopysky ，29 .4.1 965 ，Vasak Fl. Ocho t. 6 (1 847). (MHA); SLOVENIA; Kobic ， Kobarid ， 25 .4 .1980 ， TYPE: RUSSIA: “Kh abarovski Reg. ， Chiapella Chiapella 10186 (MHA); GERMANY; Halle ，Saale ， Udskoe ， Siberi orient 人 18.6 .1 844 ， 25 .4 .1980 ， Skwortsov (MHA); Gotha et Eisenach ， 3.5 .1 980 ， Skwortsow (MHA); FRANCE; Dep. Middendorf (l ectotype-LE!). Pyrenees-Atlantiques ， Vallee d'Ossau ， 16.5.1986 ， Anemonoides udensis (Trautv. & C.A. Lazare Lazare 13055 (MHA); Dep. du Hault-Rhin ，Colmar ， Mey.) Holub in Folia Geobo t. Phytotax. 27 .4 .1978 ，Rastetter 9169 (MHA); Dep. du Bas-Rhin ， Praha 8: 166 (1 973). Soufflenheim ， 18 .4 .1979 ， Schneider 9170 (MHA); 60 60 植物研究雑誌第79 巻第1号 平成 16 年2月

Rhizomes long-pros 甘ate or ascending ，2- Shreter (MHA); ne 訂 Vladivostok ，Bagrationa Station ， 6 mm in diameter ， having sometimes thick- 28.5.1976 ， Nechaeva (MHA); Sputnik ， 1. 6.1980 ， ened ened nodes. Basal leaves several ，scale-like Jakubov (MHA); 5.6.1992 ， Kharkevich & Buch (5-10 (5-10 mm long) ， and solitary ， with distinct (MHA); Sikhote-Ali 吋Reserve ，Medvezhyi ，8.6.1958 ， Mutsertony Mutsertony (MHA); Kavalerov Distr. ， Povorot ， blades; blades; petioles 5-15 (-20) cm long ，villous; 23.5.1980 ，Starodubtsev 6104 (MHA); Te りukhe Distr. ， blades blades temate ，rhombic ，3 -4 x 3.5 -4 .5 cm ， Tetjukhe ，30.6.1964 ， Chizhevskaya (MHA); CHINA; adaxially adaxially glabrous ， abaxially sp 紅 sely Uizimi ，10.6.1905 ， Syuzev (LE). villous; villous; petiolules 3-5 mm long; centralleaf- The authors of this species noted its long lets lets undivided ，obovate; bases cωunea 剖t臼e町; ma 紅f- and thin stolon-like rhizomes ， 1紅 ge gms crenate to incαised-d involucral leaves ， and white pubescent tωo tωo obtuse; lateral leaflet 臼s similar to central tepals ， but they found no basal leaves. ones ， but oblique ， ovate to obovate. Scapes However ， Juzepchuk (1 937) described the 15-25 15-25 (-4 0) cm long ，spreading-pubescent; long petioles ，their short petiolules ， and l-flowered. l-flowered. Involucral leaf petioles 2-3 solitary flowers. cm x 1-2 mm; blades temate ， similar to The most distinct characters of this taxon those those in basal leaves ， but larger ，rhombic- are the narrow involucralleaf petioles ，leaf- pentagonal ，4-7 x 5-7 cm ，abaxially sparsely let petiolules 3-5 mm long ， and 5 elliptic puberulent; puberulent; petiolules 3-5 mm long; central pilose tepals 10-18 mm long with 5-9 leaflets leaflets mainly undivided ，rhombic-obovate; anastomoses. We confirm the opinion of bases bases cuneate; margins crenate to se 町 ate; Starodubtsev (1991) conceming the alliance apices apices broadly acute to obtuse; lateral seg- of A. udensis and A. umbrosa. However ， the ments ments similar to central ones ， but oblique- latter differs from the former by its 5-8 elliptic elliptic (Fig. lE). Pedicels 5-10 cm long ， glabrous tepals having solitary vein puberulen t. Tepals 5，obovate-elliptic ， with anastomoses and much shorter leaflet wide wide bases and na 町 ow apices ，white ，10- petiolules. Nevertheless ，we believe that A.

18 18 x 5-13 mm ，appressed 司 pilose (sometimes udensis is closest to A. trifolia ， having in subglabrous); subglabrous); basal veins 5-7 ， vein common with it such characters as the undi- anastomoses anastomoses 5-9. Stamens 4- 6 mm long; vided basal and involucral leaflets ， filaments filaments filiform; anthers ellipsoid ，connec- involucral leaf petioles 1-3 cm long ， tives tives wide. Ovaries oblong-ovoid ， 1-2 mm monomo 叩hic tepals ， and achene styles 1.5- long ，densely covered with hairs 0.1-0.2 mm 2.5 mm long. long; long; styles straight or curved ， 1-1.5 mm long; long; stigmas linear (Fig. 2F). Achene bodies 7. Anemone trifolia L. ， Sp. Pl. 540 basally basally slightly narrowed ，ovoid ， slightly (1 753). compressed ， with narrow ribs ， 2-3 x 1. 5 TYPES: “Habitat in Gallia" ，N 710.24 mm ， densely puberulent (hairs 0.1-0 .2 mm (LINN!); Anemone 2， Herb. Clifford long); long); styles s紅白 ght to curved ，1. 5-2 mm (l ectotype-BM!). long ，glabrous; stigmas linear (Fig. 3C). Anemone trifolia L. subsp. trifolia. Chromosome number: n = 24 (St 紅 0- Anemonanthea trifolia (L.) Niew l. in dubtsev dubtsev 1991). Amer. Mid l. Na t. 3: 174 (1914). Distribution: Distribution: NE Asia: Russia (Far East) ， (L.) Holub in Folia China China (J ilin ，Liaoning) ，N Korea; occurring Geobo t. Phytotax. Praha 8: 166 (1 973). in in forests or shady slopes ， al t. 20 0- 500 m. Subsp. albida (Mariz) Ulbr. in Bo t. Jahrb. Specimens Specimens examined: RUSSIA; Primorje Reg.: ， 37: 220 (1 906). Ocean Station ，29 .4 .1956 ， Voroshilov 8193 (MHA); TYPE: PORTUGAL: “Serra do Gerez: Suputinka Suputinka River ，Mountain-Forest Station ，17.6.1959 ， Leonte ，Caldas ，" Henriq & Tait (Herb. ?). February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 61

Anemone albid αMariz， in Bo l. SOC. Bro t. 1970 ， Favarger 1961) ， 16 (Langlet 1932). 4: 4: 101 (1886). Distribution: Europe (Finland ， Austria ， Anemonoides albida (Mariz) Holub in France ，Spain ，N Portugal ，It aly ，Croatia ， Folia Folia Geobo t. Phytotax. Praha 9: 166 (1 973). Bosnia & Herzegovina ，Slovenia ，Hungary ， Subsp. Subsp. brevidentata Ubaldi & Puppi in Romania); occurring inforests and forest Candollea Candollea 44: 145 (1 989). margins ， al t. 100-1800 m. TYPE: ITALY: “ Borzonasca (Genova) ， Specimens examined: AUSTRIA; Carinthia ， prope iuxta iuxta viam ad Brizzolara in silva Ostryae urbem Klagenfurt ，1821 ， Jabornegg (KW); Kl agenfurt ， 1825 ， Schult (KW); Carinthia ，Villach ，Dobratsch ， C訂 pinifoliaea" ，2.5.1979 ， Ubaldi (holotype ・- 10.9 .1 990 ，Skwortsow (MHA); ITALY; Apennines ， BOLO). 1811 ，Gaudin (MHA); Mts.Andepus et Safira Croc ， Rhizomes long-branched ，horizontal to as- 1854 ，de Pavillon (KW); Prov. Udine ， Valli del cending ，2 -4 mm in diameter. Basal leaves Natisone ，1 1. 5.1980 ， Chiapella (MHA).

2-3 ，scale 聞 like ，and solitary ， with distinct The author' s description included the pres- blades; blades; petioles 8-25 (-30) cm long ，sparsely ence of rhizomes ， temate basal leaves with pilose; pilose; blades temate ，3-5 (-9) x 4-6 cm ， ovate undivided se 町 ate leaflets ，and one- adaxiall adaxiall y sp 紅 sely pilose along veins ， flowered stems. Pritzel (1841) noted the abaxially abaxially pilose throughout; petiolules 3-5 nodulose rhizomes ，solitary basalleaves with mm long; centralleaflets undivided ， ovate to subsessile segments ， involucral leaves on elliptic; elliptic; bases cuneate; margins se 町 ate to distinct petioles ，and glabrous or pubescent dentate; dentate; apices acuminate; lateral leaflets C紅 pels. In their 位eatments of Anemone for similar similar to central ones ， but oblique. Scapes the “ Flora Europaea" both Tutin (1964) and 12-25 12-25 (-35) cm long ， glabrous or sp 征 sely Chater (1 973) regarded A. trifolia as allied to

pilose; pilose; cymes 1-2- flowered. Involucral leaf A. nemorosa ，and Chater mentioned its 3目 petioles petioles 1-3 cm x 1-2 mm ， glabrous or P紅白d (not temate) basal leaves without any sparsely sparsely puberulent; blades temate ，similar to further divisions (which are frequently ab- those those in basal leaves ， but larger ，4-8 x 5-10 sent). cm ，pilose; petiolules 1-2 mm long; central This species has some major distinctions leaflets leaflets undivided ， rhombic to elliptic; lateral from A. nemorosa (shape of basal and leaflets leaflets similar to central ones ， but oblique involucral leaf blade leaflets ， n紅 row (Fig. (Fig. lF). Pedicels 3-8 (-10) cm long ， involucral leaf petioles ， monomorphic pilose. pilose. Tepals 5-8 ，elliptic ， with wide bases perianth ， achenes with na 町 ow ribs ，and lin- and acute apices ， white or bluish ，12-20 x 8- ear stigmas). However ， the same characters 12 12 mm ， glabrous; basal veins 3-5 ， vein render it close to A. udensis ， although A. anastomoses anastomoses solitary. Stamens 4- 8 mm long; trifolia distinctly differs from A. udensis by filaments filaments apically dilated; anthers oblong- its few-flowered cymes ，5-8 glabrous tepals ellipsoid ，connectives wide. Ovaries oblong- with wide bases and apices ， having solitary ovoid ， basally narrowed ， 0.7-2 mm long ， vem anastomoses. densely densely covered with hairs 0.1-0.2 mm long; Anemone trifolia is known as a variable styles styles straight or slightly curved ， 0.7-1 mm taxon with several subspecies or varieties. long; long; stigmas linear (Fig. 2G). Achene bod- We accept three subspecies of A. trifolia: ies ies ovoid-ellipsoid ，slightly compressed ， with subspp. trifolia ，α lbida and brevidentata. narrow narrow ribs ， 3-4 x 1.5-2 mm ， densely puberulent puberulent (hairs 0.1 -0 .2 mm long); styles Key to the subspecies

apically apically curved ， 1-1.5 mm long ，glabrous; la. Tepals white; achene heads nodding... ・H ・ stigmas stigmas linear (Fig. 3D). … b. subsp.α lbida Chromosome number: n = 8 (Baumberger 1b. Tepals mainly bluish; achene heads erect 62 62 植物研究雑誌第79 巻第1号 平成16 年2月

...... 2 ...... 2 branced ， 1-1.5 mm in diameter. Basalleaves 2a. 2a. Tepals 15-20 mm long; scapes 20-25 cm 2-3 ，scale-like ， and solitary ， with distinct long; long; leaf blade teeth distinc t...... blades; petioles 5-15 cm long ，puberulent or

-…...・ H ・...・ H ・H ・H ・.a. subsp. tl φ lia subglabrous; blades 3-sected ，pentagonal ，2- 2b. 2b. Tepals 10-15 mm long; scapes 15-20 cm 4x 2 -4 cm ， appressed-puberulent or long; long; leaf blade teeth minute … glabrous; petiolules 1-2 mm long; central …...・ c. subsp. brevidentata segments 3-1obed ， elongate-ovate; bases cordate cordate to cuneate; margins se 汀 ulate; apices a. 7 a. Anemone trifolia L. subsp. trifolia. acuminate to obtuse; lateral segments une- Plants Plants are characterized by all features of qually 2-1obed ， ovate to orbicular; bases nar- species species as well as its erect achene heads ，dis- rowly cuneate; m 紅 gins crenate to dentate. tributed tributed throughout its range except the Scapes 8-20 (-30) cm long ，subglabrous or Iberian Iberian Peninsula. pilose; cymes 1- flowered. Involucral leaf 7b. 7b. Anemone trifolia L. subsp.α lbida petioles 1-2 (-3) cm x 1-2 mm ，glabrous or (M 紅 iz) Ulbr. pilose; blades 子sected ，similar to those in Plants Plants are characterized by white flowers basalleaves ，pentagonal to triangular ，2 -4 x and and nodding achene heads; occu 町ing in NW 3-5 cm ，puberulent; segments subsessile; Spain Spain and Portuga l. central segments 3-1obed ， rhombic; lateral 7c. 7c. Anemone trifoli αL. subsp. breviden- segments unequally 2-1obed ， obovate to el- tata tata Ubaldi & Puppi. liptic (Fig. 1G). Pedicels 3.5-6 cm long ， Plants Plants differ from those of subsp. trifolia puberulent or villous. Tepals 5，elliptic ， with by the smaller size of all parts ，including the wide bases and apices ，white ，7-14 x 4-10 flowers ， and in particular ， the shorter leaf mm ，sparsely puberulent; basal veins 5-6 ， blade blade teeth (ca. 1 mm deep); occurring in vein anastomoses absent (rarely solitary). Apennines. Apennines. Stamens 4- 6 mm long; filaments filiform; anthers anthers ellipsoid; connectives na 町ow. 8. 8. Anemone umbrosa C.A.Mey. in Staminodes 2.5 -4 mm long sometimes pre- Ledeb. ， F l. Al t. 2: 361 (1 830). sen t. Ovaries basally narrowed ， oblong- TYPE: “ In sylvis umbrosissimis ad ovoid ， 1-1.2 mm long ， densely covered with rivulum rivulum Grammatucha prope Riddersk hairs 0.1-0.3 mm long; styles straight or Sibiriae Sibiriae altaicae ぺLedebour 795 (holotype- slightly curved ， 0.8-1 mm long; stigmas LE!). LE!). slightly dilated (Fig. 2H). Achene bodies Anemone koraiensis Nakai in Bo t. Mag. ovoid-ellipsoid ，2 -4 x 1.5-2 mm ， densely Tokyo 33: 7 (1 919). TYPE: “ Corea media ， in puberulent (hairs 0.2-0.3 mm long); styles campis campis Senpo ，" Ishidaya 3089 (holotype- TI!). straight or slightly curved ， 0.7-1 mm long ， Anemone umbrosa C.A.Mey. subsp. glabrous; stigmas linear. extremiorientalis extremiorientalis Starod. in Bo t. Zhum. 67: Chromosome number: n = 8 (Starodubtsev 353 353 (1982). TYPE: “ Oriens Extremus ， 1982). Primorje Primorje Meridionalis ， secus fluvium Li- Distribution: NE Asia: Russia (Far Eas t) ， Fudin ，" 21.5.1869 ，Maximovicz (holotype- China (He i1 ongjiang ，Jilin ，Liaoning) ， Korea; LE!). LE!). occurring in forests and shadow slopes ， al t. Anemonoides umbrosa (C.A.Mey.) Holub 200-500 m. in in Folia Geobo t. Phytotax. 8: 166 (1973). Specimens examined: RUSSIA; Primo りe Reg. ， Anemonoides extremiorientalis (Starod.) Nadezhdinski Distr. ， close Baranovskaja ，3.6.1974 ， Starod. ，Vetrenytsy 162 (1 991). Kh arkevicz (MHA); ne 紅 Vladivostok ，17 .4 .1976 ， Rhizomes horizontal or ascending ，long- Nechaeva (MHA); Te りukhe Distr. ， near Tetjukhe ， February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 63

17.6.1964 ， Chizhevska (MHA); Khasanski Dist r.， Koidzumi [l ectotype (designated by Hara in Kedrovaja Kedrovaja Padj Reserve ， 18 .1 0.1977 ， Makarov 1976)-TI]. (MHA); Temej Distr. ，Kunami ，19.6.1985 ， Shaulska Anemone sciaphila M.Popov in Bo t. (MHA); KOREA; Manchuria ， Prov. Cham-gyon ， Mate r. Herb. Komarov Bo t. Ins t. Ak ad. M t. Musanga ，4.6.1897 ，Komarov (LE). This This species was described as close to A. Nauk URSS 14: 141 (1 951). TYPE: nemorosa on the basis of its ternate basal and RUSSIA: Sachalin ，18.5.1948 ，Popov (holot involucral involucral leaves (distinct petioles in A. ype-LE!). nemorosa) ， and solitary flowers with 5 white Anemone umbrosa C.A.Mey. subsp. pubescent pubescent tepals. Juzepchuk (1 937) noted its sciaphila (M.Popov) Starod. in Bo t. Zhurn. deeply deeply bipartite lateral leaflets (resemble 67: 354 (1 982). palmatifid) palmatifid) and short thickened carpel stig- Anemonoides soyensis (Boiss.) Holub in Folia Folia Geobo t. Phytotax. Praha 11: 81 (1 976). mas. mas. Recently ， Starodubtsev (1 991) sepa 圃 rated rated A. extremiorientalis from A. umbrosa Anemonoides sciaphila (M.Popov) Starod. on the basis of its basal and involucral leaf in Vetrenytsy 162 (1 991). blade blade distinctions ， and he classified both spe- Rhizomes horizontal or ascending ，long ， cies cies in the genus Anemonoides. cylindric ， 1-2 mm in diameter. Basalleaves Anemone umbros αdiffers from A. 2-3 ，scale-like (3-5 mm long) ， and solitary ， with with distinct blades; petioles 10-25 cm long ， nemorosa by its n紅 row involucral leaf peti 旬 。les and several essential flower characters glabrous; blades 3-sected ，pentagonal ，2 -4 x ， (e.g. ， monomo 中hic perianth ， pubescent 2-4 cm adaxially sparsely pilose; petiolules tepals tepals without anastomoses). It is rather 1-2 mm long; central segments toothed ， close close to A. soyensis ， having in common with ovate to rhombic; bases cuneate; margins the the latter leaflet petiolules 1-2 mm long ，1- crenate-dentate; apices acute to obtuse; lat- flowered flowered scapes and glabrous tepals lacking eral segments similar to central ones ， but vein vein anastomoses. Because of the variability basally oblique. Scapes 10-20 cm long ， of of the shape of the leaf blades (the diagnostic subglabrous; cymes l-flowered. Involucral character) ， we are not recognizing A. leaf petioles 1-2 cm x 1-2 mm ，glabrous; extremiorientalis. extremiorientalis. blades 3-sected ，wide-elliptic ，2 -4 x 2-3 cm ， glabrous glabrous or pilose along margins; petiolules 9. 9. Anemone soyensis Boiss. in Bull. 1-2 mm long; central segments elliptic to Herb. Herb. Boiss. 7: 590 (1 899). oblanceolate; bases narrowly cuneate; mar- TYPES: JAPAN: Hokkaido ， Prov. Kitami ， gms crenate to se 町 ate; apices acuminate; lat- Cape Soya ，20.6 .1 891 ， Faurie 7212 (l ecto- eral segments similar to central ones (Fig. type ，designated by Hara in 1976-P); Soya lH). Pedicels 2-5 (-7) cm long ，pilose. Division ，Wakkanai ，19.5.1926 ， Saito 1747 Tepals 5-7 ，elliptic-obovate ， with wide bases (paratype-BM!). (paratype-BM!). and apices ，white ， 9-12 x 4- 8 mm ，glabrous; Anemone debilis Fisch. ex Turcz. var. basal veins 3-5 ， vein anastomoses absent soyensis soyensis (Boiss.) Makino in Bo t. Mag. (sometimes solitary). Stamens 3-7 mm long; Tokyo 19: 86 (1 905). filaments filiform; anthers oblong-ellipsoid ， connectlves connectlves n訂 row. Ovaries narrow-ovoid ， Anemone umbrosa C.A.Mey. v訂 . yezo ・ ensis ensis Miyabe ex Matsum. ， Shokubutsu Meii 1-2 mm long ， densely covered with hairs rev. rev. ed. 23 (1 895) ，nom. nud. 0.01 mm long; styles slightly curved ，0.5-0.8 Anemone yezoensis Koidz. in Bo t. Mag. mm long; stigmas linear (Fig. 21). Achene Tokyo 31: 138 (1 917). TYPE: JAPAN ， bodies ovoid-ellipsoid ， 4- 5x 1-2 mm ， Hokkaido ， Prov. Ishikari ， M t. Daisetsu sparsely puberulent (hairs less than 0.1 mm long); long); styles curved to recurved ， ca. 1 mm 64 64 植物研究雑誌第79 巻第l号 平成16 年2月 long ，glabrous; stigmas linea r. ex Turcz.) Huth in Bull. Herb. Boiss. 7: 1074 Chromosome number: n = 16 (Starodubt- (1 897). TYPE: JAPAN: Yesso ， Kiu-Siu und sev sev 1982). Riu-Kiu-Inseln aub Rebunshiri und Riishiri. Distribution: Distribution: NE Asia: Russia (Sakhalin) ， Anemone caerulea DC. subsp. gracilis (F. Japan Japan (Hokkaido); occurring in forests ， al t. Schmidt) Ulbr. var. gracilis Ulbr. in Bo t. 100-1500m. Jahrb. 37: 218 (1 906). Specimens Specimens examined: RUSSIA; Sakhalin ， Anemone caerulea DC. subsp. gracilis 18.5.1948 ，Popov (LE); Dolinski Distr. ， near Dolinsk Ulb r. v紅 . linearis (Schltd.) Ulbr. ， 1. c. 15.5.1949 ，Tolmachev (MHA); 2.5.1953 ，Gyzha & (1 906). Motorina Motorina (KW); 6.7 .1 980 ， Starodubtsev (MHA); Anemone gracilis F.Schmidt var. debilis Korsakovski Korsakovski Dis 江， near Novikova ，8.6.1962 ， Egorova 75 75 (MHA); Mt. Majorova ，1 1. 6.1966 ， Egorova 3353 (Fisch. ex Turcz.) Koidz. in Bo t. Mag. (MHA); Starodubskoe ， 6.5.1968 ， Egorova & Tokyo 31: 139 (1 917). RUSSIA: Kamts- Alekseeva Alekseeva (LE); Okhotskoe ，17.5.1981 ，Mazurenko & chatka ，Fischer (l ectotype-LE!). Khokhriakov Khokhriakov (MHA); Evstafievo ， 5.6.1981 ， Anemone gracilis F. Schmidt v紅 . linearis Mazurenko & Kh okhriakov (MHA). CHINA; (Schld t. ex Ulbr.) Koidz. ，1. c. (1 917). Lyaonin ，Benjsi ，Sezsiachun ，19.5.1950 ，Noda (LE). Anemone debilis JAPAN; Hokkaido ， M t. Ashibetsu ， Furano-shi ， Fisch. ex Turcz. var. 13.6.1978 ， Mimoro & Tsugaru 4896 (MHA); linearis (Schld t. ex Ulbr.) Makino & Vagishiri ， Haboro-cho ， Tomamae-gun ， 5.6.1979 ， Nemoto ， F 1. Jap. ed. 1: 961 (1 925). Mimoro 2025 (MHA); Kitami Prov. ，Utanobori ，29 .4. Anemone debilis Fisch. ex Turcz. var. 1972 ，Yamanoi 108 (TNS). gracilis gracilis (Schltd t.) H.H 紅 a in Bo t. Mag. The status of A. soyensis was debatable for Tokyo 49: 6 (1 935). a long time ， and was v訂 iously regarded as A. Anemonoides debilis (Fisch. ex Turscz.) debilis debilis var. soyensis or A. umbrosa subsp. Holub in Folia Geobo t. Phytotax. Praha 8: sciaphila. sciaphila. Moreover ，similar plants were de- 166 (1 973). scribed scribed twice as A. yezoensis (from Japan) Rhizomes long ，horizontal or ascending ， and and as A. sciaphila (from Sakhalin). 1. 5-3 mm in diameter. Basal leaves 2-3 ， In In our opinion ， A. soyensis is allied to A. scale-like (5-8 mm long) ， and solitary ， with umbrosa and differs from the latter mainly distinct blades; petioles 3-8 (-15) cm long ， by shape of central segments of the basal glabrous; blades ternate ，pentagonal ， 2-3 x leaves leaves (3-10bed rhombic in the former and 2-4 cm ，adaxially pilose; petiolules 2-5 mm se 町 ate elongate-ovate in the latter) ， and long; central leaflets narrow-ovate to larger larger glabrous tepals ， but smaller achenes. lanceolate-elliptic; bases cuneate; margins crenate crenate to co 訂 sely toothed; apices acute or 10. 10. Anemone debilis Fisch. ex Turcz. in obtuse; lateral segments similar to central Bull. Bull. Soc. Na t. Mosc. 27: 274 (1 854). leaflets ， but smaller and sligh t1 y oblique. TYPE: RUSSIA ， Kamtchatka ， Tigil ， Scapes 5-15 (-20) cm ，pilose; cymes ト Rieder Rieder (l ectotype-LE!). flowered. Involucral leaf petioles 1-2 cm x Anemone ranunculoides L. var. gracilis 1-2 mm ， basally sligh t1 y vaginate ， F.Schmidt F.Schmidt in Linnaea 6: 574 (1 831). mainlysubglabrous; blades temate ， 2-5 x 2- Anemone linearis Schltd. ex Ledeb. ， F 1. 3 cm ， glabrous or rarely sparsely pilose Ross. Ross. 1: 14 (1 841). along veins; petiolules 2-5 mm long; central Anemone caerulea DC. var. gracilis leaflets linear-lanceolate; bases narrowly (Schltd.) (Schltd.) Ledeb. ，1. c. (1841). cuneate; margins entire to serrulate or Anemone gracilis F. Schmidt in Mem. toothed; apices acuminate; lateral leaflets Acad. Acad. Sc i. St. -Petersb. 7: 102 (1 868). similar to central ones (Fig. 11). Pedicels 1- Anemone caerulea DC. var. debilis (Fisch. 2 cm long ，pilose. Tepals 5-6 ，oblong- February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 65 elliptic ， with na 町 ow bases and rounded api- conceming A. debilis. However ， Juzepchuk ces ， white or greenish abaxially ， 5-7 x 2-5 (1937) ， Ohwi (1957) and Starodubtsev mm ， glabrous; basal veins 3-5 ， vein (1991) noted the short thick branches of its anastomoses anastomoses absen t. Stamens 2.5-5 mm rhizomes ，linear-lanceolate leaf segments ， long; long; filaments filiform; anthers ellipsoid ， solitary flowers with 5 small white glabrous connectives connectives wide. Ovaries ovoid ， 0.5-1 mm tepals ， and narrowly ovate ， small (ca. 3 mm long ， densely covered with hairs 0.1 mm long) pubescent achene bodies. long; long; styles straight or curved ，0. 4-0 .7 mm Anemone debilis has several characters long; long; stigmas linear (Fig. 2J). Achene bodies (e.g. ，greenish-white glabrous tepals without ovoid-ellipsoid ， slightly compressed ， with vein anastomoses) which are absent in A. n紅 row ribs ， 2-4 x 1-1.5 mm ， densely caerulea ， but present in A. soyensis. In addi- puberulent puberulent (hairs less than 0.1 mm long); tion ， both of these taxa have similar achenes. styles styles straight to curved ， 0.5-1.2 mm long ， Therefore ，we confirm the opinion of Dutton glabrous; glabrous; stigmas linear (Fig. 3E). (1 996) conceming the close relationship of Chromosome number: n = 8 (Kurita 1961 ， these species ， together with A. umbrosa. Starodubtsev Starodubtsev 199 1). According to our data ， A. debilis differs from Dstribution: Dstribution: NE Asia: Russia (Far East ， A. soyensis mainly by its smaller tepals (4- Sakhalin ， Kamchatka ， Kuriles) ， J apan 8 mm long only in the former and 10-15 mm (Hokkaido ， Honshu); occurring in coniferous in the latter) ， colour (white-greenish and forests forests and open slopes ， in mountains. white) and shape (narrow and wide bases). Specimens Specimens examined: RUSSIA; Primorje Reg. ， Nikolaevsk Nikolaevsk near Amur ，18.5.1903 ， Shestunov (LE); Ser. 2. Altaicae (Starod.) Ziman ， Kadota Tatarski Tatarski Strait ， De-Castri Bay ，23.5 .1 912 ，Derbek & Bulakh. (LE); (LE); Sovetskaja Bay ， near sea ，3 1. 5.1 916 ，Kr ylov (E); 11. Anemone altaica Fisch. ex C.A.Mey. S Sakhalin ， vicinity South Sakhalinsk ， Kanuma ， l. l. t. (1 5.10.1948 ，Popov (LE); Firsovo ne 訂 sea ，13.6.1959 ， in Ledeb. ，F Al 2: 362 830). Pobedimova & Konovalova 298 (LE); S Kamchatka ， TYPES: “ In montibus altaicus" ， Schangin Petropavlovsk ， 17.6.1921 ， Hulten (LE); Kuriles ， [(herb. apud Ledebour) holotype-LE!]; “ In Kunashir ， near Urvitovo ，6.6.1963 ， Egorova 899 (LE); montibus montibus Kurtschum haud r征 a" ，Meyer; “Ad Magadanski Magadanski Reg. ，Kh asynski Dist r.， ne 訂 Snezhnaja ， portum Petro- Pauli Kamtschatkae" ， A. de Dolina ，2 1. 6.1975 ，Kh okhriakov (MHA); Kamchatka Peninsula ， Elizovo ， 25.7.1969 ， Dvorakovskaya Chamisso (paratypes-LE!). (MHA); Sakhalin: near V i11 age Te 中enie ，25.6 .1 965 ， Anemone nemorosa L. subsp. altaica Egorova Egorova (MHA); near Juzhno-Sakhalinsk ，16.6 .1 980 ， (Fisch. ex Ledeb.) Korsh. in Acta Hort. Starodubtsev Starodubtsev & Nedoluzhko (MHA); Tymovski Dist r.， Petropo l. 12: 29 (1 892). Kirovskoe ，25.6.1980 ， Starodubtsev & Nedoluzhko Anemonoides altaica (Fisch. ex C.A.Mey.) (MHA); near V i11 age Aleksandrovka ， 26.6.1980 ， Holub in Folia Geobo t. Phytotax. Praha 8: Starodubtsev Starodubtsev 5521 (MHA). JAPAN; Hokkaido ， Prov. Tokachi ， Oikimanai Lake ，28.5.1975 ， Furuse 8628 (K); 166 (1 973). t. M t. Ashibetsu ，Furano-shi ， 13.6.1978 ，Mimoro & Rhizomes horizontal stolon-like ，ca. 2 mm Tsugaru Tsugaru 4897 (MHA); Honshu ，Nikko ， Tochigi Pre f.， in diameter ， and oblique short nodulose ，5- 7.1924 ， Momiko (LE); Manza Pass ， 1800 m ， 8 mm in diameter. Basal leaves 2-3 ，scale- 19.5.1963 ， Kanai (LE); Nagano Pref. ， M t. Ogura ， like (4- 6 mm long) ， and solitary ， with dis- Ki taaiki ・mura ，26.5.1976 ， Furuse 11066 (K); Shizuoka Pref. ， M t. Tekaridake ， al t. 2500 m ，15.6.1979 ， Kadota tinct blades; petioles 4-8 cm long ， 7068 (TNS). subglabrous; blades temate ，2.5 -4 x 3-6 cm ， Anemone debilis was firstly described as a glabrous; petiolules 5-10 mm long; central p紅 t of several different species(e.g. ， A. leaflets 3-10bed ，pentagonal; bases cuneate; gracilis ， A. caerulaea). Moreower ，there was m 紅 gins crenate to denticulate; apices considerable considerable taxonomic confusion acuminate or obtuse; lateral leaflets similar 66 66 植物研究雑誌第79 巻第1号 平成16 年2月 to to central leaflets ， but bilobate or bicleft ， (MHA); Novosibirski Reg. ， Toguchinski Dist r.， slightly slightly oblique ， and the ultimate lobules ob- Kotorovo ，7.6 .1 974 ，Lashchinski & Volkova (MHA). tuse. tuse. Scapes 10-20 (-30) cm long ， BASHKIRIA; 8 km N of Ufa ，M t. Sutoloka ，2.5 .1 942 ， Kotov Kotov (KW); 11. 5.1942 ， Kotov (KW); 18.5.1942 ， subglabrous; subglabrous; cymes l-flowered. Involucral Kotov (KW). RUMANIA; Transsylvania ， Muresului leaf leaf petioles 1-3 cm x 3-5 mm ，f1 at and intre Toplita si Deda ，27.6.2000 ， Oroian & Ziman winged ，sparsely puberulent or subglabrous; (KW). CHINA; W Hupeh ，7 .4 .1900 ， Wilson 53 (K). blades blades temate ， larger than those in basal Anemone aZtaic αinitially was separated leaves ， 3-6 x 4-8 cm ，puberulent; petiolules from A. nemoros αon the basis of the distinc- 2-5 2-5 mm long; centrallea f1 ets 3-1obed ，rhom- tions in its rhizome and leaf shape ， and tepal bic bic to ovate; bases cuneate; margins i町 egu- number and shape. Ulbrich (1 906) ，however ， larly larly dentate or se 町 ate; apices acuminate; regarded A. nemorosa ，A. αZtaica ， and A. lateralleaflets lateralleaflets smaller ，oblique ，ultimate lob- umbrosa as “species collectiva". ules ules obtuse (Fig. lJ). Pedicels 3-5 cm long ， Anemone alt α ica is close to A. nemorosa puberulen t. Tepals 8-12 ， oblong to on the basis of several characters (dilated narrrowly narrrowly elliptic ， with rounded bases and involucral petioles ，3-parted or 3-1obed cen- apices ，white ， bluish or reddish-violet ，di- tral lea f1 ets ，solitary f1 owers ， and dimorphic morphic ，glabrous ， in two circles: outer glabrous tepals with 3-5 vein anastomoses) ， tepals tepals 20-25 x 8-12 mm ， basal veins 5-9 ， but the distinctions between these taxa 紅 e vein vein anastomoses 3-5; inner tepals 10-12 x essential ， and they include the types of rhi- 3-4 mm ， without anastomoses. Stamens 4- zomes (dimorphic in A. αZtaica) ， perianth 8 mm long; filaments filiform; anthers ellip- (many-leaved) and achenes (smaller but hav-

soid ， connectives wide. Ovaries na 町 ow 田 ing longer styles). ovoid ， basally narrowed ， 1-1.5 mm long ， Anemone aZtaica is distributed mainly in densely densely covered with hairs 0.1-0.3 mm long; NE Asia. Recently Oroian (1997) discovered styles styles straight ，0. 4- 0.7 mm long; stigmas a unique population in C Europe (Romania ， subcapitate subcapitate (Fig. 2K). Achene bodies basally Transsilvania). In 2000 we re-examined slightly slightly narrowed ， ovoid ， slightly com- plants of this population together with pressed ， with narrow ribs ，4- 5x 1.5-2 ffiffi ， Orpian and we agree that plants of this popu- densely densely puberulent (hairs 0.1-0.3 mm long); lation are indeed A. aZtaica (not A. nemorosa). styles styles curved to recurved ， 0.5-1 ffiffi long ， basally basally puberulent; stigmas slightly dilated 12. Anemone pseudoa It aica H.Hara in J. (Fig. (Fig. 3F). Jpn. Bo t. 15: 767 (1 939). Chromosome number: n = 8 (Baumberger TYPES: JAPAN: “Hakodate (Hokkaido) ， 1970) ， 16 (Chupov 1975) ， 32 (Lavrenko and iter secundum ぺ1861 ，Maximowicz (holo- Serditov Serditov 1986). type-GH!; isotypes- BM! ， NY!). Distribution: Distribution: Europe (Romania) ，NE Asia: Anemone h αkodatensis Nakai in Bull. Na t. Russia Russia (Altai) ， China (Henan ，Hubei ，Shanxi ， Sci. Mus. (33): 6 (1 953). TYPE: JAPAN: Xinjiang); Xinjiang); occu 町 ing in forests and under Hondo bo r. & Hokkaido aust r.， Prov. Uzen ， bushes ， also by streams ， al t. 1200-1800 m. 30 .4 .1908 ，Ogunimoto (l ectotype-TNS!). Specimens Specimens examined: RUSSIA; Turukhanski Reg. ， Anemone αZtaica Fisch. ex Ledeb. f. Podkamennaya Tunguska ，6.1888 ，Kηvchikova (KW); aZbiflora Yamam. & Tsukam. in F1. Pechorski Pechorski Reg. ，1881 ，Ivanitski (MHA); Tomski Reg. ， Hakodate 37 (1 932). near near Tomsk ，9 .5 .1903 ， Nekrasov (KW); Stepanovka ， Anemone pseudo αhα ic αH.H: 紅 a var. 28.5.1923 ， Sergievskaya (KW); Kutasheva Village ， 20.5 .1 933 ，Konusova & Yakubova (KW); Ushayka pseudoaltaica. River ， 18.5 .1 948 ， Nogina & Mikhailova (KW); Anemone pseudo αZtaica H.H; 紅 a v 紅. Kemerov Reg. ， near river Tesh ，20.5.1948 ，Kuminova gracilis (H.Hara) H.Ohba in J. Jpn. Bo t. 69: February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 67

116 116 (1 994). TYPE: JAPAN: “Honsyu Distribution: NE Asia: Japan (Hokkaido ， (Honshu). (Honshu). Prov. Sagami ， Hakone" Honshu); occurring mainly in forests ， al t. 30 .4 .1939 ，Hisauti" (holotype-TI!). 100-1200m. Anemone pseudoaltaica H.Hara var. Specimens examined: JAPAN; Hokkaido ，Hakodate ， katonis katonis H.Ohba ，1. c. (1 994). TYPE: JAPAN: 1861 ，Maximowicz (K); Insula Jesso ， circa Hakodate ， “Honshu Prov. ，Yamagata Praef. ， Tobishima 27 .4 .1861 ， Albrecht (LE); Prov. Shiribeshi ， Okushiri 1.， Is 1.， 2.5.1983 ， Takahashi 3822 (MHA); Prov. Oshima ， Island 4.1993 ，H.Ohba (holotype-TI!). ぺ Nanae ，Onuma Park ，alt. 150m ，10.5.2003 ，Igarashi Anemonoides pseudoaltaica (H.Hara) (TNS); Honshu ， Miyagi Pref. ，Onoda-cho ， Naruse Holub in Folia Geobo t. Phytotax. Praha 8: River ，9.5.1983 ， Ohashi & al. 11770 (MHA); Prov. 166 166 (1 973). Oshima ，Hakodate-shi ， M t. Hakodateyama ，4.5.1985 ， Rhizomes Rhizomes long horizontal or ascending ，2- Takahashi 5362 (MHA); Hondo ，Echio ，Arakawa- 3 mm in diameter ， and short nodulose ， 5-6 machi ，1. 4.1956 ， Togashi 1283 (K); Tohoku ， Prov. Rikuzen ， M t. Daito ， 22 .4 .1977 ， Takahashi (K); mm in diameter. Basalleaves 2-3 ，scale-like Tominami ， Prov. Uzen ， Murayama-city ， 4.5.1977 ， (3 -4 mm long) ， and solitary ， with distinct Takahashi (K); Fukushima Pref. ，ShIrakawa-no-seki- blades; blades; petio1es 5-15 cm 10ng ，glabrous; ato ，26 .4 .1978 ，Mimoro & Tsugaru 4659 (MHA); blades blades temate ， pentagona1 ， 2-3 x 2-3 cm ， Fukushima ，ne 訂 Shinkassi-onsen ，Mafune ，Nishigo- glabrous glabrous or sparsely pilose along veins; mura ，27 .4.1 978 ，Mimoro & al. 4717 (MHA); Ishikawa Pref. ，M t. Saruyama ，27.3.1987 ， Deguchi & al. 6575 petiolules petiolules 2-5 mm 10ng; cen 町al 1eaflets (MHA). deeply deeply 3-10bed ， ovate to rhombic; bases nar- This species was described from the flora rowly rowly cuneate; margins se 汀 ate and incised; of J apan as similar to A. αltaica ， but differing apices apices acuminate or acute; latera1 1eaflets in the shape of basal leaf blades and tepal wide-ovate ， deeply bilobed or bipartite. colou r. According to our data ， A. Scapes Scapes 10-20 (-30) cm long ， glabrous~ pseudoaltaica is close to A. altaica by its cymes 1-flowered. lnvolucral leaf petioles basal leaf blades but differs mainly by its 10-20 10-20 x 3-5 mm ， basally dilated ， monomo 中hic perianth ，basally dilated fila- subglabrous; subglabrous; blades temate ，similar to those ments ，basally rounded ovaries and achenes in in basal 1eaves ， but larger ， 3-6 x 3-5 cm ， without lateral ribs ， sublinear stigmas ， glabrous; glabrous; petiolules 3-5 (-10) mm long (Fig. shorter achene sty1es ， but 10nger achene 1K). 1K). Pedice1s 2.5-5 cm 10ng ，pilose. Tepals hairs. 8-12 ，lanceolate ， with rounded bases and In 1953 Nakai described A. hakodatensis apices ，reddish-purple or rarely whitish ，10- as a species close to A. altaica ， but differing 30 30 x 5-10 (-20) mm ， monomorphic ， mainly by its tepal colour ， and later ，Ohwi glabrous; glabrous; basal veins 3-5 ， vein anastomoses (1 984) ， the author of treatment on Anemone 3-5. 3-5. Stamens 5-7 (-10) mm 10ng; fi1aments for the “ Flora of Japan" Ohwi (1 984) did not basa l1 y dilated; anthers ellipsoid ，connectives recognize this taxon. wide. wide. Ovaries ovoid ，ca. 2 mm long ，densely We examined herbarium material of A. covered covered with hairs 0.3-0.5 mm long; styles hakodatensis and believe that its essential S甘泊 ght ， ca. 1 mm long ， stigmas sublinear characters (rhizomes of two types ，dilated (Fig. (Fig. 2L). Achene bodies ovoid-ellipsoid ， and winged involucral leaf petioles up to 30 l1 basa y narrowed ， 3.8-5 x 1.5-2 mm ， mm 1ong ，involucral leaf b1ades larger than sp 訂 sely puberulent (hairs 0.2 -0 .3 mm long); those in basal leaves ， and 8-12 mono- styles styles curved or recurved ，0.7- 1. 5 mm long ， morphic tepals) are simi1ar to A. pseudo- glabrous glabrous or basally sparsely puberulent; stig- altaic α; therefore ， we do not accept mas sublinear (Fig. 3G). A. hakodatensis as a species. Chromosome number: n = 16 ，24 (Kurita Anemone pseudoaU α ica has been regarded 1956 ， Nishikawa 1985). 68 68 植物研究雑誌第79 巻第1号 平成16 年2月 as as a variable taxon ， and within it three varie- Anemone maximowiczii Juz. in Kom. ， F l. ties ties (e.g. ， v紅 s. pseudoaltaica ， gracilis URSS 7: ?? (1 937). (H.Hara) (H.Hara) H.Ohba and katonis H.Ohba) ， and Anemone amagisanensis Honda in Bo t. four four forms (e.g. ，f. hanagasa Inag 北 i ，f. Mag. Tokyo 49: 1 (1 934). pleniflora pleniflora Inagaki ，f. tosshozana Inagaki and Anemone juzepczukii Starod. in Bo t. f. prol 俳ra Kadota) were described (l nagaki Zhum. 68: 1016 (1 983). TYPE: RUSSIA: and and Nakano 1964 ， Ohba 1994 ， Kadota “ Oriens ex 住emus ， insula Sachalin ， 1998). 1998). After an examination of the herbar- promontorium meridionale jugi Susun 吋， ium material and the pertinent literature ，we vicinitis opp. Juzhnoδachalinsk ， prope pag. recognize recognize the three varieties of A. Kanuma" ，7.5.1948 ，Popov (holotype-LE!). pseudoaltaic α. Anemone raddeana Regel var. raddeana. Anemone raddeana Regel var. integra Key to the varieties Huth in Bull. Herb. Boiss. 5: 1072 (1 897). la. la. Scapes 25-30 cm long; tepals 25-30 mm TYPE: JAPAN: “ Prope Hakodate. It er long...... ・H ・H ・H ・.....・ H ・.....・ H ・..… c. var. katonis secundum ぺ1861 ， Maximowicz (syntype- lb. lb. Scapes 5-20 cm long; tepals 10-25 mm GH!). long. ・・・・…・...... ・…・・・・・・…・・・・・・… ....2 Anemone raddeana subsp. villosa Ulbr. in 2a. 2a. Scapes 15-20 cm long; tepals 15-25 mm Bo t. Jahrb. 37: 221 (1 906). TYPE: “So long.... ・H ・.....・ H ・-…...・ H ・.a. vぽ . pseudoaltaica nur auf dem Festlande; auf dem Schan- 2b. 2b. Scapes 5-15 cm long; tepals 10-15 mm Alingebirge und dessen Auslaufern ，z. B. bei long...... ・H ・.....・ H ・H ・H ・.....・ H ・.. ..b. var. gracilis Bai Viktoria" ，1860 ， Nadimoff (syntype). Anemone raddeana Regelsubsp. glabra 12a. 12a. Anemone pseudoaltaica var. pseudo- Ulb r.， l. c. 221 (1 906). altaica. altaica. Anemonoides raddeana (Regel) Holub in This This variety is characterized by all fea- Folia Geobo t. Phytotax. Praha 8: 166 (1 973). tures tures of A. pseudoaltaica. Plants occur in Anemonoides raddeana v紅 . lacerata Y. L. Hokkaido Hokkaido and Honshu (Hyogo Pref. and Xu in Bull. Bo t. Res. Forest Univ. 13: 121 northward) ， and the above forms occur in the (1 993). TYPE: CHINA: 4.4 .1991 ，Y.L.Xu t. M t. Tosshozan of Hokkaido and in Aomori 888 /1 456814 (holotype-PE!). Pref. Pref. of Honshu. Rhizomes long horizontal or ascending ，2- 12b. 12b. Anemone pseudoaltaica var. gracilis 3 mm in diameter ， and short nodulose ， 4-8

(H.Hara) (H.Hara) H.Ohba in J. Jpn. Bo t. 69: 116 mm in diamete r. Basalleaves 2-3 ，scale 四 like (1 994). (1 0-20 mm long) ， and solitary ， with distinct Plants Plants differ from those in var. blades (develop close to scapes); petioles 5- pseudoaltaica pseudoaltaica by shorter scapes (5-15 cm) ， 20 cm long ，glabrous or sparsely puberulent; and and smaller f1 0wers (tepals 10-15 mm long). blades temate ， 2-5 x2 -4 cm ，pentagonal or 12c. 12c. Anemone pseudoaltaica var. katonis suborbicular ，subglabrous or sparsely pubes- H.Ohba in J. Jpn. Bo t. 69: 116 (1 994). cent; petiolules 3-5 mm long; centrallea f1 ets Plants Plants differ from those in both other va- medially 3-1obed or deeply toothed only ， rieties rieties by longer scapes (25-30 cm long) and obovate-elliptic; bases cuneate to truncate; larger larger f1 0wers (tepals 25-30 mm long). margins coarsely crenate to se 町 ate; apices obtuse; obtuse; laterallea f1 ets similar to central ones ， 13. 13. Anemone raddeana Regel in Bull. but oblique. Scapes 10-30 cm long ， Soc. Soc. Na t. Mosc. 34: 16 (1861). subglabrous; cymes l- f1 owered. Involucral TYPE: “ Mandshuria ad f1. Amur montes leaf petioles 5-12 x 1-2 mm ， glabrous; Burejae" ，4 .1 858 ，Radde (holotype-LE!). blades 3-sected ，2 -4 x 2-5 cm ，rhombic- February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 69 pentagonal ，sericeous; petiolules 1-2 mm Shirakawa-no-seki-ato ，26 .4 .1978 ， Mirnoro & Tsugaru long; long; all segments narrowly ovate ，apically 4658 (MHA); Saitarna Pre f.， Chichibu-gun ，M t. Buko ， deeply deeply but obtusely toothed (Fig. lL). 1917 ，島 1akino 33964 (LE); Shiga Pref. Inukarni-gun ， M t. Oikedake ， al t. 900 rn ，29 .4 .1963 ， Kodarna 9964 Pedicels Pedicels cm long sp 訂 sely puberulent or 1-3 1-3 ， (TNS). subglabrous. subglabrous. Tepals 9-15 ，linear-lanceolate ， This species was described from the Far with with rounded bases and apices ，white-tinged East as plants with short rhizomes and large purple purple or white ，12-20 x 4-5 mm ，glabrous; scales ，solitary basal leaves with long peti- basal basal veins 3-5 ， vein anastomoses 1-3. oles and ternate blades ，and solitary flowers Stamens 8-15 mm long; filaments basally with 10-15 sepals. The specific status of this and apically slightly dilate; anthers ellipsoid ， species was never in doubt ， but its relation- connectives connectives na 町 ow. Ovaries ovoid ， basally ships remained uncertain. narrowed narrowed (stalks ca. 0.1 mm long) ， ca. 2 mm Anemone radde αna stands within sec t. long ， densely covered With hairs 0.1 mm Anemonanthea close to the A. αhα ica- long; long; styles s町aight ，ca. 1 mm long; stigmata A. pseudoaltaica subgroup ， having in com- slightly slightly dilated (Fig. 2M). Achene bodies mon several essential characters (rhizomes ovoid ovoid to ellipsoid ，slightly compressed ， with of two types ，8-12 glabrous tepals with sev- narrow narrow ribs ， 3-4 x 1.5-2 mm ， densely eral vein anastomoses ，solitary flowers ，and puberulent puberulent (hairs 0.2 -0 .3 mm long); styles ovoid carpels and achenes with distinct curved ， 1-1.5 mm long ， glabrous; stigmata styles). There are ，however ， several impor- sublinear. sublinear. tant distinctions ， mainly in leaf shape (in A. Chromosome number: n = 16 (Kurita raddean αthe basal and involucral leaflets 1956 ， Baumberger 1970 ， Starodubtsev are are toothed or medially 子lobed ， the 1991). 1991). involucral leaf petioles much shorter and Distribution: Distribution: Russia (Far East) ， China narrower ， the flowers have more numerous ， (Heilongjiang ， Jilin ， Shandong ， Zh 吋iang) ， longer but narrower tepals having fewer vein Japan Japan (Hokkaido ，Honshu ，Shikoku) ， Korea; anastomoses ，and dilated filaments). occu 町 ing in forests and meadows ， al t. 100- Juzepchuk (1 937) regarded the allied ， but 1000 m. glabrous ， plants from Japan as A. Specirnens Specirnens exarnined: RUSSIA; Ussuri Reg. ， Ussuri maximowiczii. Another allied species ， A. Reserve ，ne 訂 M 司ski Kljuch ，14.5.1972 ， Kostenko ， (VLA); 2.5.1974 ， Kharkewicz & Buch (MHA); juzepczukii was described by Starodubtsev Arnurski Arnurski Reg. ，Arkharinski Dist r.， Kundur ，4.6.1981 ， (1983) from Sakhalin on the basis of its less Starodubtsev Starodubtsev & Nedoluzhko (MHA ，VLA); Prirnorje divided involucralleaf segments. In addition ， Reg. ， village Novoshivskoe ， 14 .4.1 913 ， Gurzenko several infraspecific .taxa of A. raddeana (LE); (LE); Russkyi Island ，22 .4 .1921 ， Desulavi 302 (LE); were described (viz. ， subspp. villosa Ulbr. ， Upper p訂 t of Suputinka Valley ， 20 .4 .1932 ， and glabra Ulbr.; v紅 s. integra Huth and Skibinskaya Skibinskaya (LE); Kh asanski Distr. ，village Prirnorski ， 9.5.1980 ， Starodubtsev (VLA); Kh asanski Distr. ， lacerata Y.L.Xu). Kedrovaja Kedrovaja Padj ， 10.9.1980 ， Starodubtsev (MHA); We agree with Ohwi (1 984) in not recog- Sakhalin ， near Kh olrnsk ，Uentornari Valley ，19 .4 .1948 ， nizing of A. maximowiczii in the flora of Popov (LE); near Tyrnovski ，12.5.1968 ， Egorova & Japan ，and we do not support any recognition Sharornova Sharornova (LE); 5.5.1972 ， Kostenko (VLA); 14.5 of a specific status for A. juzepchukii ， but in 1972 ， Kostenko (VLA); ne 訂 Dolinsk ，6.7.1980 ， Starodubtsev Starodubtsev (VLA); 24.6.1982 ，Starodubtsev (VLA). following Wang & al. (2001) ，we accept va- JAPAN; Hokkaido ， Insula Jesso ， circa Hakodate ， rieties raddeana and lacerata. In this 町eat- 1861 ， Albrecht (LE); Prov. Th uri ，Tornakornai-shi ， ment ， va r. lacerata differs from var. Misawa ，27 .4 .1984 ， Takahashi 4889 (MHA); Prov. raddeana by its villous basal leaf petioles ， Hidaka ，Mitsuishi-gun ，Mitsuishi-rnachi ，20.5.1984 ， broadly rhombic basal and involucral leaf Takahashi Takahashi 4904 (MHA); Honshu ， Fukushirna Pref. ， 70 植物研究雑誌第79 巻第1号 平成16 年2月 central central segment blades ， and their many den- Kumazawa M. 1936. Pollen grains morphology in tate tate apices. Moreover ，v 訂 . lacerata differs Ranunculaceae ， Lardizabalaceae and Berberi- also also from vぽ . raddeana in that it is a local daceae. Jap. J. Bo t. 8: 19 -4 6. endemic endemic within China (N Zhejiang). Kurita M. 1955. Cytological studies in Ranunculaceae 111. 111. The karyotypes of seven species in Delphinium ， Lycoctonum and Aconitum. Bo t. Mag. Tokyo 68: References References 248-251 (in Japanese). Baumberger Baumberger H. 1970. Chromosomenzahlbestimmun- 一一一 1956. Cytological studies in Ranunculaceae IV. ges ges und K 訂 yotypanalysen bei den Gattungen Further notes on karyotypes of De 伊hinium and its Anemone ，Hepatic αund Pulsatilla. Be r. Schweiz. allied genera. 1. Jpn. Bo t. 31: 192-195 (in Bo t. Ges. 80: 17-96. Japanese). Candolle Candolle A. P. de 1817. Regni Vegetabilis Systema Linnaeus K. 1753. Species Plantarum. Vo l.1. Naturale. Naturale. Vo l. 1. Paris. Stockholm. 一一一 1824. Prodromus Systematis Naturalis Regni Spach H. 1839. Anemone L. Historie Naturelle des Vegetab i1i s. Vo l. 2. Paris. Vegetaux Phanerogames. XV: 242-253. Paris. Chater Chater A. O. 1973. Anemone L. Fl ora Europaea ，ed. 2. Miller P. 1754. Gardemer's Dictionary. London. 1: 1: 262-264. Cambridge University Press ， Ohba H. 1994. A new variety of Anemone pseudo- Cambridge. Cambridge. altaica H. Hara (Notulae ad plantas Japoniae 5). J. Ehrendorffer Ehrendorffer F. and Samuel R. 200 1. Contribution to a Jpn. Bo t. 69: 116-118 (in Japanese). molecular molecular phylogeny and systematics of Anemone Ohwi J. 1957. Anemone L. Fl ora of Japan: 44 4-4 46. and and related genera (Ranunculaceae-Anemoninae). Smithsonian Institution ， Washington D. C. Acta Acta Phytotax. Sin. 39: 293-309. Oroian S. 1997. Vestigiile unei populatii de Anemone

Femald M. L. 1928. Contribution from the Gray nemorosa L. ssp.α lt α， ica (Fisch.) Korsh. in defileul Herbarium Herbarium of Harvard University. 82. The North Muresului intre Toplita si Deda. Ocro t. Na t. Med. American American Species of Anemone Section Inco 吋ura t. 41: 77. Anemonanthea. Anemonanthea. Rhodora 30: 180-188. Prantl K. 1888. Beitrage zur Mo 叩hologie und Finet Finet A. and Gagnepain F. 1904. Contributions a la Systematik der Ranunculaceen. Bo t. Jahrb. Sys t. 9: flore flore de l' Asie Orientale d'apres l' herbier du mu- 225-272. seum de Paris. IV. Anemone. Bull. Soc. Bo t. Fr. Pritzel E. 1842. Anemonarum revisio. Li nnaea 15: 51: 51: 56-76. 561-698. Gray S. F. 182 1. Anemonanthea D C. Na t. Arr. Bri t. Pl. Savitski V. 1982. Morphology ， Evolution and 2: 2: 72 4- 725. Classification of Pollen Grains within the Family Holub J. 1973. New names of Phanerogamae. 2-3. Ranunculaceae. 124 pp. Kiev ，Naukova Dumka (in Folia Folia Geobo t. Phytotax. Praha 8: 155-179. Russian). Holmgren P. K.， Holmgren N. H. ， Bamette L. C. 1990. Starodubtsev V. N. 1989. New taxa of the subtribe Index Index Herbariorum. 1. The of Herbaria the W orld. Anemoninae (Ranunculaceae). Bo t. Zhum. 74: Ed.8. Ed.8. Regnum Vegetabile. 134 4- 1346 (in Russian). Hoot S. ， Rezniceka A. A.， Palmer J. D. 1994. 一一一 199 1. Vetrenytsy: Systematics and Evolution. Phylogenetic Phylogenetic relationships in Anemone 197 pp. Leningrad ，Nauka (in Russian). (Ranunculaceae) (Ranunculaceae) based on mo 叩hology and Tamura M. 1967. Morphology ， ecology and phylogeny chloroplast chloroplast DN A. Sys t. Bo t. 19: 169-200. of the Ranunculaceae - 7. Sci. Rep. Osaka Univ. Huynh K.ーL. 1970. Le pollen du genre Anemone et du 16: 21 -4 3. genre genre Hepatica (Ranunculaceae) et leur taxonomie. 一一一 1991. A new classification of the family Pollen Pollen et Spores 12: 329-364. Ranunculaceae 2. Acta Phytotax. Geobo t. 42: 177- Inagaki Inagaki K. and Nakano T. 1964. [Title??]. 1. Hokkai 187. Gakuen Univ. ，sec t. 11 B ， 15: 25. 一一一 1995. Anemone L. Die naturlichen J anczewski E. 1892. Etudes mo 中hologiques sur le Pflanzenfamilien. Band 17: 324-349. Berlin ， genre genre Anemone L. Rev. Gen. Bo t. 4: 241-258. Duncker ，Humblo t. Juzepchuk Juzepchuk S. V. 1937. Anemone L. In: Komarov V. L. 一一一， Mizumoto Y. and Kubota H. 1977. Observation (ed.) ， Flora URSS 7: 236-283. Moscow ，Nauka (in on seedlings of the Ranunculaceae. J. Jpn. Bo t. 52: Russian). Russian). 293-304. Kadota Kadota Y. 1998. A new form of Anemone Ulbrich E. 1906. Ub er die systematische Gliederung pseudoaltaica pseudoaltaica H.Hara from Aomori ，Japan. J. Jpn. und geographische Verbreitung der Gattung Bo t. 73: 289. Anemone L. Bo t. Jahrb. Sys t. 37: 171-256. February February 2004 Journal of Japanese Botany Vo l. 79 No. 1 71

Wang W. -T. 1974. Notulae de Ranunculaceis 一一一， Ziman S. N. ， Dutton B. E. 200 1. Anemone L. Sinensibus. Sinensibus. Acta Phytotax. Sin. 12: 170-180. Flora of China 6: 307-328. Science Press ， Beijing 一一一 1980. Subtribe Anemoninae Spach. Flora & Mossouri Botanical Garden Press ，S t. Lous. Reipub l. Popu l. Sin. 28: 1-72. Beijing (in Ziman S. 1985. Mo 中hology and Phylogeny of the Chinese). Chinese). Family Ranunculaceae. 248 pp. Kiev (in Russian).

S. ジーマンヘc. S. キーナ -1 門田裕一C ，E. ブ

ラークヘ O. ツァレンコヘB. E. ダットン d .イチリ ンソウ属 Anemonanthea BE:.属[広義] (キンポウ イカイチゲ， A. tr( ザfoli 臼αL.， A. um ηlbrosa αC.A ゲ科)の分類学的再検討 I リイチゲゲ、 ， A. soyensis Boiss. エゾイチゲ， A. debilis キンポウゲ科イチリンソウ属のうちで広義の Fisch. ex Turcz. ヒメイチゲ A. altaica Fisch. ex Anemonanthea 亜属の分類学的研究を行った.そ C. A. Mey. ，キクザキイチゲ A. pseudoaltaica H.Hara ， の結果，広義の Anemonanthea 亜属に， 7 つの新 アズマイチゲ A. raddeana Regel の13 種について報 属内分類群を含む， 2 亜属， 5 節， 11 列を認める 告した. ことカ f できた. (a ウクライナ・ M.G. コロドゥニイ植物学研究所，

ノ〈ート I では ， Anemone nemorosa L.， A. bペンシルパニア州立大学， amurensis amurensis (Korsh.) Kom. ウラホロイチゲ(ヤチイ c国立科学博物館植物研究部，

チゲ)， A. caerulea L.， A. uralensis D C.， A. d西オレゴン州立大学) ranunculoides ranunculoides L.， A. udensis Trautv. & C. A. Mey. ノt