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JJB 079 43 71.Pdf 官官庁志位・@ノ物刷mBMbFUol雑エコロイ人 ギ心ゆ 'v ， ιc d 同，，J A、‘，， 一 斗 A Taxonomic Revision of Anemone L. Subgenus Anemonanthea (DC.) (DC.) J uz. sensu lato (Ranunculaceae) 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 leaf development ， types of rhizomes ， shoot structure ， shape and and size of basal and involucralleaves ，tepal 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 ， taxonomy. 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 leaves ， 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 genus 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 flower 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 plants 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 flowers with 5 to 20 tepals ， 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 rhizome 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.
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