植物研究雑誌 J. J. Jpn. Bo t. 80:282-302(2005)

A Taxonomic Revision of L. Section Omalocarpus DC. sensu sensu lato (): Part 1

Svetlana Svetlana N. ZIMAN a ，Friedrich EHRENDORFER b， Yuichi KADOTA¥ Carl S. KEENER d， Olga N. TSARENKO a， Elena BULAKH a，Bryan E. DUTTON e

aN. aN. G. Kholodny Institute of Botany ，National Academy of Sciences ， Tereshchenkivska Tereshchenkivska 2，Kiev ， 01601 UKRAINE; E-mail: E-mail: [email protected] bInstitute bInstitute of Botany of the Vienna University ，Rennweg 14 ，Vienna ， 1030 AUSTRIA; CDepartment CDepartment of Botany ，National Science Museum ，Tokyo Amakubo Amakubo 4-1-1 ，Tsukuba ， 305-0005 JAPAN; dThe dThe Pennsylvania State University ，208 Mueller Laboratory ， University University Park ，Pennsylvania ， 16802 U. S. A.; eDepartment eDepartment of Biology ，Western Oregon State University ， Monmouth ，Oregon ，97361 U. S. A. (Received (Received on March 15 ，2005)

The of Anemone section Omalocarpus (Ranunculaceae) ， with members distributed distributed throughout the Northem Hemisphere and especially in East Asia ， was re- evaluated evaluated on the basis of a critical mo 中hological analysis of extensive herbarium mate- rials rials and living populations. A conspectus ，key ，mo 中hometric tables ， and figures illustrating illustrating taxonomic characters 訂 e presented. Three series and 11 species ，in c1 uding

eight eight subspecies and 12 varieties of A. narcissiflora and four varieties of A. demissa 紅 e recognized recognized and discussed with respect to their relationships. This paper will be divided

into into two parts. In Part I the results of the analysis the A. narcissiflora complex are re 田 ported. ported.

Key words: Anemone narcissiflora ，Anemone sec t. Omalocarpus D C.， comparative mor- phology ， taxonomy.

Because of its considerable diversity of the Anemone which ， for the A. species ，Anemone L. is one of the most diffi- narcissiflora complex ， included 10 species ， cult cult genera in the Ranunculaceae. The A. two subspecies and six varieties. However ， narcissiflora narcissiflora complex 陀 presented by A. this 佐eatment was too brief and did not in- narcissiflora narcissiflora and related taxa is taxonomi- clude data for the whole sec t. Omalocarpus. cally cally most complicated within Ane- Consequently in this paper we present a criti- mone. The complex is widely disjunctive and cal morphological analysis of extensive her- occur occur within the Northen Hemisphere while barium material of all the foregoing taxa other other species occur mainly in East Asia. with respect to their taxonomy ，distribution In In the previous papers (Ziman et al. 1994 ， and relationships. 1995 ，1997 ，etc.) we examined a part of the A. A. narcissiflora complex ， mainly in limits of Literature review the the former Soviet Union. Recently ， in the Anemone narcissiflora L. (error narcissi- “ Flora of China ，"羽 Tang ，Ziman and Dutton folia) was described by Linnaeus in 1753 (2001) (2001) published a taxonomic treatment of together with two allied species ， A. sibirica October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 283

L. L. and A. fasciculata L. ， from Eurasia. scence distinction. He included A. narcissi- Willdnow (1797) later described A. flora and A. demissa in the first series ， and A. umbellata ， and Adams (1 805) described A. polyanthes ， A. elongata and A. tetrasepal αin speciosa speciosa (both from the Caucasus). the second one. De Candolle (1 817 ， 1824) recognized all In a comprehensive herbarium-based three three species within the A. narcissiflora study ， Juzepchuk (1 937) re-examined A. complex complex which he classified in sec t. narcissiflora and allied taxa collected within Omalocarpus Omalocarpus DC. on the basis of their com- the former Soviet Union. The A. narcissi- pressed ， suborbicular ， glabrous achenes ， flora complex was classified in se r. scapose scapose stems ， and umbelliferous inflore- Narcissiflorae Juz. (nomen nudum). scences. scences. Juzepchuk recognized A. sibirica ， A. During During the nineteenth century several spe- fasciculata and A. specios 仏 and he described cies cies allied to A. narcissiflora were described ten new species (A. shrenkiana Juz. ， A. [A. [A. ch ηisantha C. A. Mey. from the biarmiensis Juz. and others) in sec t. Caucasus ， A. αconitifolia Turcz. from Omalocarpus (= subgen. Homaloc 紅 pus Siberia ， and A. elongata D. Don ， A. DC.). Consequently Juzepchuk believed A. polyanthes polyanthes D. Don ， A. tetrasepala Royle and narcissiflora s. st r. did not occur within the A. A. demissa Hoo k. f. & Thoms. from the territory of the former Soviet Union ， but was Himalayas Himalayas (East Asia)]. replaced there with a series of vicariant The first sketch of sec t. Omalocarpus was races. prepared prepared by Hooker (1 872) ， and he included Within the flora of only A. in in it five species (A. narcissiflora ， A. narcissiflora was recognized (Zapalowicz elongata ， A. demissa ， A. polyanthes and A. 1908 ，Hayek 1927 ， Pawlowski 1956 ， Chater tetrasepala) ，despite the fact that the author 1973 ，Pignatti 1988 ，etc.). of of the first monograph on Anemone ，Pritzel According to Sharma et al. (1 993) ， the (1841) ， regarded sec t. Omalocarpus (= flora of India included four species of sec t. Homalocarpus) Homalocarpus) as consisting of only A. Omalocarpus (A. demissa ， A. elongata ， A. narcissiflora narcissiflora and A. polyanthes. Pritzel had polyanthes and A. tetrasepala). However ， A. included included A. elongata and A. tetrasepala in narcissiflora was regarded as absent there. sec t. Anemonospermos ， and A. sibirica in Hara and Williams (1979) recognized A. sec t. Anemonanthea. demissa ， A. elongata and A. polyanthes in Early Early in the twentieth century ， Finet and the flora of Nepal ， but they also included in Gagnepain Gagnepain (1904) viewed A. narcissiflora in this flora two new species ，A. β/'scopurpurea a broad sense and regarded A. demissa and H. Hara (1973) and A. smithiana Lauener & A. A. polyanthes as its varieties. They did not Panigrahi (1975) ，a conclusion also reached accept accept de Candolle' s circumscription of by Chaudkhary (1988). Anemone and included A. elongata and A. In the flora of Pakistan ， three endemic tetrasepala tetrasepala in the so-called seventh section ， species were described recently by Qureshi their their “eighth section" consisting of A. and Chaudhri [(1 988): A. mirajabii R. A. dichotoma ， A. gelida ， A. narcissiflora ， A. Qureshi & M. N. Chaudhri ， A. lowariensis R. rivularis rivularis and A. begoniifolia. A. Qureshi & M. N. Chaudhri and A. In In the next monograph on Anemone ， abdurrehmanii R. A. Qureshi & M. N. Ulbrich Ulbrich (1 905 ， 1906) recognized two series Chaudhri]. All three species are very close to within within sec t. Omalocarpus (= Homalocarpus) ， A. polyanthes. However ， Riedl and Nasir viz. ， ser. Involucratae Ulbr. and ser. (1 991) ，who treated Anemone for the “ Flora Involucellatae Involucellatae Ulb r.， on the basis of inflore- of Pakistan ，" did not accept these species ， 284 植物研究雑誌第80 巻第5号 平成17 年 10 月 and and in their opinion ， only A. polyanthes and in the “ Flora of China." A. A. tetrasepala from sec t. Omalocarpus occur In treating Anemone for the flora of Japan ， in in Pakistan. Ohwi (1 953) in cI uded both A. nαrcissiflora In In the flora of Iran and adjacent territories ， and A. sikokiana Makino in sec t. Rechinger Rechinger and Riedl (1 992) recognized A. Omalocarpus. protracta protracta (instead of A. narcissiflora) ， A. In the flora of ， Britton poly α nthes and A. tetrasepal α， but these (1891) recognized A. narcissiflora to be dis- authors authors believed that A. low αriensis and A. tributed disjunctively in Alaska and the abdurrehmanii abdurrehmanii also occur there. They in- Rocky Mountains. Shortly thereafter ， cluded cluded all these species in sect. Anemonan- from the Rocky Mountains were described as thea thea (not Omalocarpus). A. zephyra A. Nelson (Nelson 1906). Only In In the flora of China ，Wang (1 974 ， 1980) Rydberg (1 917) accepted this species. Later included included eight species in sec t. Omalocarpus authors (e. g. ， Calder and Taylor 1968 ， (= (= Homalocarpus) [A. crinita ， A. schrenkian α Weber 1976 ，Dorn 1977 ， Scott 1994 ，etc.) (instead (instead of A. narciss 伊ora) ， A. demissa ， A. did not recognize A. zephyr αas a distinct imbricata ，and four endemics (A. cathayensis specles. Kitag. ， A. schantungensis Hand.-Mazz. ， A. The A. narcissiflora complex was already robusta robusta W. T. Wang and A. taipaiensis W. described earlier as the genus Homalocarpus T. T. Wang)]. Quite recently Wang described Schur (Schur 1866) ， but this segrega- generic three three na 町 ow endemics from China (A. tion was not supported by later authors. tengchongensis tengchongensis W. T. Wang ， A. multi- Recently Holub (1 973) again proposed to lobulata lobulata W. T. Wang & L. Q. Li and A. sep 訂 ate sec t. Omalocarpus from Anemone as nutantiflora nutantiflora W. T. Wang & L. Q. Li (Wang a separate genus (Omalocarpus). and and Li 1986 ，Wang 1996). According to the In his summaηof additions to the flora of descriptions ，all these plants are veηclose to the former Soviet Union ， Cherepanov (1 981 ， narcissiflora. A. narcissiflora. 1996) followed Holub (1 973) ， and he ac- Recently ， in the modern version of the cepted almost all the taxa of A. narcissiflora “ Flora of China" (2001) ，Wang ，Ziman and sensu Juzepchu k. In his treatment of Dutton Dutton (2001) published a taxonomic treat- Anemone ， Starodubtsev (1 989 ， 1991) also ment of the genus Anemone ，and in it A. the followed Holub ， and accordingly recognized narcissiflora narcissiflora complex includes ten species ， sec t. Omalocarpus as the genus Anemonast ・ two subspecies and six varieties. In it A. rum Holub which in cI uded three sections multilobulata multilobulata and A. tengchongensis were re- (Anemonastrum ， Elongata Starodub. and duced duced to synonyms of A. nαrcissiflora ， and Imbricata Starodub.) consisting of 12 species A. A. nutantiflora was excluded because of the occurring in the former Soviet Union. lack lack of flowers. On the basis of a complex molecular- In In this paper we present a critical morpho- biological and morphological analysis ，Hoot logical logical analysis of extensive herbarium ma- et al. (1994) recognized sec t. Omalocarpus terials terials including a discussion of the sensu Tamura (1967) which in cI uded two relationships relationships of the taxa of the foregoing sec- subsections ， Omalocarpus and Himalayicae. tion. tion. Our analysis includes comparative data Within sec t. Omalocarpus Hoot et al. (1994) on basal and involucral leaves ， flowers and recognized seven species of the A. achenes achenes of most taxa accompanied by se- narciss{ 月ora complex (A. narcissiflora ， A. lected lected illustrations. Detailed morphological laxa ， A. sibirica ， A. crinita ， A. fasciculata ， A. descriptions descriptions of the foregoing taxa are ex- biarmiensis and A. zephyra) and five species cluded cluded because we recently published them from the flora of East Asia (A. demiss α， A. October October 2005 Journal of Japanese Botany Vo 1. 80 No. 5 285 polyanthes ， A. tetrasepala ， A. smithiana and 島laterials and Methods A. ルSCOpUl 仰 re α). These authors also noted Our treatment is based on both herbarium that that all these taxa (except A. demissa and A. and living materials. The herbarium material polyanthes) polyanthes) may be subspecies of A. involved about 4000 specimens contained in narcissiflora. narcissiflora. 32 collections (AA ，BC ，BCC ，BKL ，BM ， Recently Recently Ziman et al. (1 997) published a BRNU ，CLUJ ，E ，GH ，K ，KRA ，KRA 恥1， taxonomic taxonomic revision of the main part of the A. KW ，KW ，LW ，LWS ，LE ，MARSSJ ，MHA ， narcissiflora narcissiflora complex (within the limits of NY ，PAC ，PRG ，SAV ，SLO ，TAD ，TASH ， the the former Soviet Union and Central TBI ，TNS ，US ，V AB ，VLA ，W ，WU - abbre- Europe). Europe). This study was based on an exami- viations following Holmgren and Holmgren nation nation of 42 natural populations and ample 1998). From these specimens ， as well as herbarium herbarium materials in addition to critical from our specimens collected in Central data data on ontomorphogeny and petiolar anat- Europe ， the former Soviet U nion ， and in omy derived from living materials. The new North America (mainly in 1980-1997) ， both scheme scheme of infraspecific structure of the A. f1 0wer and fruit samples (about 350) were narcissiflora narcissiflora complex was proposed ， in studied. Standard anatomical techniques which which 12 species were reduced to subspe- were employed including light and scanning cies ，varieties and synonyms of A. narcissi- microscopy (Ziman et al. 1997) which were flora flora sensu stricto. used especially for the f1 0wer and fruit analysis. analysis. In addition to the herbarium mate- Circumscription Circumscription and Definition of Sect. rial ，we examined about 1000 adult plants Omalocarpus and 500 plants representing the early stages In In our treatment we follow mainly Tamura of growth collected in more than 40 natural (1995) (1995) by accepting that the A. narcissiflora populations of the A. narcis 叫flora complex. complex complex is properly classified in Anemone The results of the study of these plants were sec t. Omalocarpus (not “Homalocarpus"). partly analyzed in Ziman et al. (1997). According According to Tamura ，all plants within this section section have erect or ascendent short ，thick Results and Discussion rhizomes ， rosulate 3-5-sected or ternate Character evaluation basal basal leaves with basallY vaginate petioles ， As a result of our study ，we produced mainly mainly fasciculate scapes and umbelliferous Tables 1 and 2 which include average values

cymes ，a 5-8-1eaved perianth ，dilated fila 司 and limits of 23 morphome 甘ic characters of ments ， and sessile compressed ， broadly leaves ，stems ，f1 0wers and achenes. winged winged glabrous achenes in globose or The taxa of the section differ in the num- ovoidal ovoidal heads. In addition to these charac- ber and the length of stems. However ，we ters ，we believe the unbranched rhizomes ， noted a tendency of small plants to have axillary axillary scapes or stems (monopodial main more stems. The taxa differ also in size of shoot) ，sessile involucral leaves ，subglobose basal leaves and blades. However ， the basal achenes achenes with short ，more or less bent styles and involucr 叫 leaf number ， and pedicel and and 3-colpate pollen grains (Huyn 1964 ， length are too variable and appear to be cor- 1970 ，Savitski 1982) are also important char- related with the age of the plants and the acters acters in common to taxa of this section. overall localities where these plants grow. Moreover ，we reg 紅 d the perianth as tepals Most taxa are characterized by 5-7 tepals ， rather rather than sepals. but A. cathayensis ， and A. polyanthes have 5-10 5-10 tepals and A. elong αtα ， A. shikoki αnα， A. A. smithiana ， A. taipaiensis 4-5 tepals. Tepal 286 286 植物研究雑誌第80 巻第5号 平成17 年 10 月

Table 1. Averages and ranges (in parentheses) of flowers and fruits in the species of Anemone sec t. Ornalocarpus

NT LT WT NTV NTVA LST LAB WAB LAS i2 i3

A. A. cathayensis 7 15 8 1. 9 4 2 5 5.5 4.5 1.1 1. 2 5.0 (5-10) (5-10) (1 0-18) (5-10) (3-5) (1 -3) (4-5) (5-6) (4-5) (1-1. 2) A. A. demissa 7 14 8 1. 7 5 3 4 6.5 5.5 1. 9 1. 2 3 .4 (5-7) (5-7) (8-18) (4- 12) (3-7) (1 -3) (3-5) (6- 7) (4-6) (1. 8-2) A. A. elongata 5 16 9 1. 8 3.5 0 .2 4 8 6 1. 3 1. 2 6.1 (5) (5) (1 2-20) (6-12) (3 -4) (0- 1) (3-5) (6-9) (4-7) (1-1. 75) A. β lscopurpurea 4.5 8 3 2.6 4 2 4 (4- 5) (6-10) (2-5) (3-5) (1 -3) (3-5) A. A. narcissiflora 6 15 9 1. 7 5 2 4 6 5 1. 2 1. 2 5.0 (5-7) (5-7) (1 2-18) (6-10) (3-7) (0-3) (3-5) (5-8) (4-6) (1-1. 4) A. A. polyanthes 7 20 12 1. 4 4 3 5 8 7 1. 7 1. 2 6.6 (5-10) (5-10) (15-25) (8-15) (3-5) (1 -5) (4- 6) (6 ー10) (5-8) (1. 6- 1. 8) A. A. robusta 6.5 18 9 2.0 4 2 4 6.5 4.5 1.1 1. 5 6.0 (6-7) (6-7) (1 5-25) (7-10) (3-5) (1 -3) (3-5) (6-7) (4-5) (1ー1. 2) A. A. shikokiana 4.5 8 5 1. 6 3.5 。 3.5 5.5 3.5 0.7 1. 5 3.5 (4-5) (4-5) (7-10) (4- 6) (3-5) (0) (3 -4) (5-6) (3 -4) (0.6-1) A. A. smithiana 5 20 12 1. 6 6 2 5 7 4.5 1. 5 1. 5 4.6 (5) (5) (1 5-25) (10- 15) (5-7) (1 -3) (3-6) (6- 8) (4-5) (1 -2)

A. A. tα， ip αiensis 5 8 5.5 1. 4 5 4 4 6.5 3.2 1. 3 2.0 5.0 (5) (5) (7-10) (5-6) (3-7) (3-5) (3-5) (6-7) (3 -4) (1-1. 5) A. A. tetrasep αiα 4.2 17 10 1. 7 6 4(3-5) 5 10 7 1. 8 1. 4 5.2 (4-5) (4-5) (1 2-20) (6-12) (5-7) (4-6) (8-12) (5-8) (1. 5-2)

Abbreviations: Abbreviations: NT (number of tepals) ，LT (l ength of tepals ，mm) ，WT (width of tepals ，mm) ，i1 (LTIWT ratio) ，NTV (number of basal basal tepal veins) ，NTVA (number of tepal vein anastomoses) ， LSt (l ength of stamens ，mm) ，LAB (l ength of achene bodies ，mm) ， WAB (width of achene bodies ，mm) ，LAB (l ength of achene styles ，mm) ， i2 (LAB 川 TAB ratio) ，i3 (LAB/LAS ratio).

Table 2. Averages and ranges (in parentheses) of the vegetative organs in the species of Anemone sec t. Ornalocarpus

NS LS NBL LPBL LBBL WBBL 2 LIL LP NF

A. A. cathayensis 4 30 6 14 5 6 2.9 0.9 6 4 4 (2-6) (2-6) (1 5-4 0) (4-8) (6-25) (3-6) (4-9) (4-7) (2-5) (2-5) A. A. demissa 4 30 8 18 6 5 2.5 1. 2 2.5 4 5 (2-5) (2-5) (1 5-50) (5-12) (5-30) (3-8) (4-6) (2-3) (2-5) (3-7) A. A. elongata 1. 5 60 10 25 10 9 2.5 l.1 5 5 12 (1 -2) (4 0- 80) (5-15) (1 5-30) (8-15) (6-12) (5-14) (3-15) (7-15) A. A. fuscopurpure α 4 10 6 7 2.5 3 2.8 0.8 2.5 4.5 3 (2-5) (2-5) (5-15) (5-7) (5-10) (2-3) (2 -4) (2-3) (4-5) (2-5) A. A. narcissiflora 1. 2 25 5 12 5 6 2 .4 0.8 3 4 4 (1 -2) (10-4 0) (4-7) (7-20) (3-7) (4-8) (2-6) (2-8) (1-7) A. A. polyanthes 1. 2 40 5 25 8 10 2.5 0.8 4 4 10 (1 -2) (25-60) (4-7) (1 0-30) (5-10) (5-12) (3-5) (3-7) (7 -15) A. A. robusta 1. 2 80 5 20 6 8 3.3 0.3 4 7 7 (1 -2) (70-120) (4-7) (1 5-30) (4- 7) (4-9) (3-5) (5-10) (5-10) A. A. shikokiana 2 35 5 20 7 10 2.8 0.7 3.5 3.5 4 (1 -3) (25-50) (4-8) (1 5-25) (5-10) (5-12) (3 -4) (3 -4) (3-8) A. A. smithiana 2 35 7 20 6 8 2.5 0.7 4 5 5 (1 -3) (2 0-4 5) (4- 9) (1 0-25) (4- 8) (6-12) (3-5) (3-6) (3-10) A. ωipaiensis 3 30 9 12 6 5.5 2.0 1.1 3 5 3 (2 -4) (1 5-4 0) (5-12) (5-15) (3-8) (4-7) (2 -4) (3 -6) (1 -5) A. A. tetrasepala 1. 2 45 10 25 12 15 2.1 0.8 6 8 8 (1 -2) (30-60) (5-15) (1 5-30) (8-15) (1 0-20) (4-8) (4- 10) (7 -15)

Abbreviations: Abbreviations: NS (number of stems) ，LS (l of ength stems ，cm) ，NBL (number of basalleaves) ，LPBL (l ength of petioles of basal leaves ，cm) ，LBBL (l ength of blades of basalleaves ，cm) ，WBBL (width of blades of basalleaves ，cm) ，1 (LPBL /L BBL ratio) ， 2 (LBBLIWBBL ratio) ，LIL (l ength of involucral leaves ，cm) ，LP (l ength of pedicels ，cm) ，NF (number of flowers on a stem). October October 2005 Joumal of Japanese Botany Vo l. 80 No. 5 287 size size generally ranges from 8-18 x 4-12 mm ， example ， the shape of tepals (obovate ，ellip- but but in A. robusta ， A. polyanthes and A. tic or oblong-elliptic ， with rounded tips). smithiana smithiana tepals are larger (1 5-25 x 8-15 Moreover ， the colour of the perianth within mm) ， and in A. taipaiensis and A. shikokiana the m 吋ority of taxa varies considerably ， and they they are smaller (7-10 x 4-6 mm). Tepal plants with flowers having several colours lengt h/ tepal width ratios are mainly 1.6-1.9 ， occur commonly within the same population. but but in A. polyanthes and A. taipaiensis this Polychromatous flowers were also frequently ratio ratio is 1.4， and in A. robusta it is 2.0. noted ， e. g. ，whitish-blue ， orange-brown ， Mo 中hometric values of stamens and ca 中els deep blue-violet ，blackish-red ， white with are are similar in all species of this section. violet tinges ，etc. ，all within the same plan t. Within Within most Ulxa ， the achenes are mainly Styles of carpels and achenes are mainly co- similar similar (achene body size: 5-8 x 3-6 mm; nical-cylindrical with straight or slightly achene achene styles: 1. 0- 1. 4 mm) ， but there are curved apices. The basal leaves ，involucral few taxa (e. g. ， A. polyanthes ， A. tetra- bracts ， stems and pedicels are all quite vari- sepala) sepala) with rather large achene bodies (6- able with respect to their pubescence. 12 12 x 5-8 mm) and long styles (1 .5-2.0 mm). After an examination of the qualitative Achene body lengt h/ width ratios and length characters ，we noted that tepal colour varies of of achene body/length of achene style often considerably in many taxa ， although white vary. vary. An index of 1. 1- 1. 3 (in most taxa) re- flowers predominate (viz. ， in A. narcissiflora ， flects flects a subspherical achene shape ， but an A. elongata ， A. polyanthes ， A. tetrasepala ， index index of 1.4 -1.8 reflects an ellipsoidal shape. etc.). Blue flowers occur in A. demissa ， and The second index is mainly 5-6.5 because of red flowers in A. fuscopurpurea and A. the the rather short styles ， but there are several smithiana. taxa taxa having a ratio of 3.2-3.5. Tamura (1 995) noted the dilated filaments After After an examination of about 80 charac- in sec t. Omalocarpus ， but our research ters ，we narrowed the list to 26 critical showed only a tendency for dilated fila- mo 中hometric characters and 19 critical ments. In most species the filaments are only qualitative qualitative characters essential for revising slightly broadened ， and only in several taxa the the taxonomy of the section. (e. g. ， in A. smithiana) they are distinctly di- With respect to quantitative characters ， the lated. following following are regarded as' essential: length We analyzed tepal pubescence and vena- and and width of basal leaf blades and petioles tion variation (including anastomosing veins) due due to a tendency to change from orbicular- which have become useful characters in ovate ovate (wider than longer) blades with long working out the infraspecific taxonomy of narrow narrow petioles to oblong (l onger than the A. narcissiflora complex (Ziman et al. wider) wider) blades with short broad or nearly ab- 1997). sent sent petioles. If all petiolules of these leaves Several sections of Anemone have a di- become longer ， they transform into the twice mo 叩hic perianth and staminodes ， although 3-sected 3-sected ones ， and if only a petiolule of the previous taxonomists had overlooked these central central segment becomes longer ， the basal characters. According to our data ， in sec t. leaves leaves transform into pinnatifid ones. Within Omalocarpus dimorphic tepals occur in A. the the quantitative characters of achenes ， the polyanthes and A. demissa ， and staminodes achene achene body and style length are considered in A. demissa. the the essential ones. Finally ， the majority of the plants of this There There are many similar or variable charac- section are characterized by densely ters ters within the qualitative characters; for pubescent basal and involucral leaves ， stems 288 植物研究雑誌第80 巻第5号 平成17 年10 月

Table Table 3. Critical taxonomic and phylogenetic characters in Anemone sec t. Omalocarpus

Character Character state 。

Basal Basal leaf blade division 3-10bed or 3-parted 3国 sected or pinnatifid Basal Basal leaf blade shape wider than long longer than wide Stem length 20 cm or more less than 20 cm Inflorescences Inflorescences compound branched simple umbelliferous or l-flowered Involucral Involucral leaves larger than basal leaves smaller than basal leaves or or equal to them Flower Flower number 3 or more less than 3 Tepal Tepal number 5-7 4-5 or 6-10 Tepal Tepal length more than 10 mm 10 mm or less than 10 mm Tepal Tepal length/width ratio less than 1. 7 1.7 or more Perianth Perianth monomorphic sometimes dimorphic Tepal Tepal anastomosing veins present absent Staminodes Staminodes absent some tI mes present Tepal Tepal pubescence pubescent glabrous Filament Filament shape linear or slightly dilated distinctly dilated Style Style shape straight or slightly bent curved and pressed Achene pubescence pubescent glabrous and and pedicels. Pubescent tepals ， however ， grains ， achenes with curved styles ，com- occur occur in half of the taxa (e. g. ， in A. pressed achene bodies ，wrinkled ， folded or demissa ， A. taipaiensis ， A. robusta and tuberculate achene surface ， and base chro- others) ， although glabrous achenes predomi- mosome number of x = 7. nate. nate. On the other hand ， plants of se r. After After our examination of the significance lnvolucellatae are characterized by com- of of the foregoing characters ，we selected 16 pound branched inflorescences ， presence of critical critical characters for our taxonomic and large involucral leaves and small bracteoles ， phylogenetic phylogenetic analyses (Table 3). This list ir ト 3-parted or 3田 lobed basal leaf blades ，ellip- cludes cludes variations in the shape of basal and soid pollen grains ， and achenes having involucral involucral leaf blades ，tepals ，filaments ， and straight or bent styles ， wrinkled achene sur- achene achene styles ， stem and tepal size ， flower face ， and x = 7 or 8. and and tepal number ， pubescence of tepals and achenes ， and achene surface. As a result of Taxonomic treatment our our study we noted the essential distinctions ln this part we describe characteristics of between between series lnvolucratae and each species of sect Omalocarpus ， with lnvolucellatae lnvolucellatae Ulb r. (1906) on the basis of taxonomic notes. We exclude from our de- inflorescence inflorescence differences ，although we added scriptions the characters in common to all several several additional details. taxa ，viz. ，short ，vertical ， unbranched rhi- According According to our data ， within se r. zomes ，axillary reproductive shoots ，basally lnvolucratae lnvolucratae simple umbelliferous cymes vaginate petioles of basal leaves ， cordate predominate ， with occasional compound bases of their blades ， sessile involucral inflorescences inflorescences or solitary flowers. The most leaves ，abaxially pubescent tepals ，spherical characteristic characteristic features of these plants are the achene heads ， compressed achene bodies ， 3-sected 3-sected basal leaf blades with distinct three-colpate pollen grains ， and chromosome petiolules ， spherical or ellipsoid pollen numbers. October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 289

Conspectus of the subdivisions in Sect. Se r. 3. Fuscopurpurea (V. F. Tarasevicz Omolocarpus & R. P. Chaudhary) S. Ziman ，Kadota & Anemone L. ， Sp. Pl. 538 (1 753). Type: A. Bu1akh ，sta t. nov. Type: A. ルscopurpurea H. coron αri αL. Hara. Sec t. Omalocarpus DC. ，Sys t. Na t. 1: 212 Anemone sec t. Fuscopurpure αV. F. (1817). (1817). Type: A. narcissiflora L. Tarasevicz & R. P. Chaudhary in Bo t. Zhurn. Sec t. Petasodes Spreng. ，Sys t. 2: 1963 72: 895 (1 987). (1 825). Inflorescences simp1e-umbelliferous; basa1 Sec t. Homalocarpos DC. in Linnaea 14: 1eaf b1ades 3-parted ， without petio1u1es; 124 (1 841). achenes unknown; basic chromosome num- Homalocarpus Schur ， Enum. Pl. ber unknown. Transsi1v. Transsi1v. 2 (1 866). 11. Anemone ルscopurpurea H. Hara Subsec t. Omalocarpus (DC.) Tamura in Sci. Sci. Rep. Os ak: a Univ. 16: 21 (1 967). Key to the species of Anemonastrum Ho1ub in Fo1ia Geobo t. Anemone section Omalocarpus Phytotax. Phytotax. 8: 165 (1 973). 1a. Inflorescences main1y simp1e-umbellifer- Ser. Ser. 1. Involucratae U1b r. in Bo t. Jahrb. ous or flowers solitary; basa1 1eaf b1ades 3- 37: 37: 209 (1 906). Type: A. narcissiflora L. sected; tepa1s main1y 6-15 mm 1ong ，fi1a- Inflorescences Inflorescences main1y simp1e-umbellifero- ments linear; achenes with curved sty1es

us us or flowers solitary; basa1 1eaf b1ades 3- pressed to achene bodies …...・ H ・..…...・ H ・..… 2 sected ， with distinct petio1u1es; pollen grains 1 b. Inflorescences compound ， branched; main1y spherica1; achenes with curved sty1es basa1 1eaf b1ades 3・parted or 3-1obed; tepa1s pressed pressed to achene bodies; achene surface 10-25 mm 1ong ， fi1aments slightly di1ated;

wrink1ed ， fo1ded or tubercu1ate; basic chro- achenes with straight or bent sty1es ...・ H ・...8 mosome numbers x = 7. 2a. Tepa1s 15-20 mm 1ong ， glabrous; stems 1. 1. Anemone narcissiflora L. 70-120 cm 1ong; basa1 1eaf b1ades orbicu1ar-

2. 2. Anemone demiss αHoo k. f. & Thoms. ovate ， subg1abrous ...・ H ・.....・ H ・.. 3. A. robusta

3. 3. Anemone robust αW.， T. Wang 2b. Tepa1s smaller ，more or 1ess pubescent; 4. 4. Anemone taipaiensis W. T. Wang stems 1ower; basa11eaf b1ades various shapes 5. 5. Anemone smithiana Lauener & ..3

Panigrahi Panigrahi 3a. Tepa1s 4-5 ，pubescent …...・ H ・..…...・ H ・...4 6. 6. Anemone catayensis Kitag. 3b. Tepa1s 5-10 ， pubescent or glabrous …6 4a. 4a. Basa1 1eaf b1ades ovate ，3-7 x 4-8 cm; Ser. Ser. 2. Involucellatae U1br. in Bo t. Jahrb. tepa1s white ， with 3-5 anastomosing veins ，

37: 37: 209 (1 906). Type: A. polyanthes D. Don 7-10 x 5-6 mm ....・ H ・....・ H ・. 4. A. taipaiensis Inflorescences Inflorescences compound ， branched; basa1 4b. Basa1 1eaf b1ades rounded ，wider than 1eaf 1eaf b1ades 3-parted or 3-1obed ， without 1ong; tepa1s red (pink) ， with 1-3 anasto-

petio1u1es; petio1u1es; pollen grains ellipsoid; achenes mosing veins and of various size .…....・ H ・..5 with with straight or bent sty1es; achene surface 5a. Basa1 1eaf b1ades pentagona1 ，pi1ose ，4- wrink1ed; wrink1ed; basic chromosome number x = 7 8x 6-12 cm; tepa1s ovate to elliptic ，15-

or or 8. 25 x 10-15 mm ....・ H ・....・ H ・... 5. A. smithiana 7. 7. Anemone polyanthes D. Don 5b. Basa1 1eaf b1ades rhombic ，glabrous ，2- 8. 8. Anemone tetrasepala Roy1e 3x 2 -4 cm; tepa1s 1inear-1anceo1ate ，6-10 x

9. 9. Anemone elongata D. Don 2-5 mm ....・ H ・..…....・ H ・. 11. A. fuscopurpurea 10. 10. Anemone shikokiana Makino 6a. Basa1 1eaf b1ades orbicu1ar; invo1ucra1 1eaves 1eaves 1arger than basa1 1eaves; tepa1s 6-10 ， 290 植物研究雑誌第80 巻第5号 平成17 年 10 月 pink pink or white ，glabrous …・ 6. A. cathayensis nated here): “Habitat in Alpibus ，Austriae ， 6b. 6b. Basal leaf blades not orbicular; Helvetiae ，Sibiriae". N.710.19 (LINN !). involucral involucral leaves smaller than basal leaves; Basal leaves 4-9; petioles 5-30 cm long ，

tepals tepals 5-7 ， of various color ， pubescent or villous or pubescent; blades 3-sected ，pen 同

glabrous glabrous …...・ H ・.....・ H ・H ・H ・..……....・ H ・-… H ・H ・...7 tagonal ，suborbicular or orbicular-ovate ，3- 7a. 7a. Stems mainly solitary ，erect; basal leaf 7x 4-12 cm ，sparsely pubescent ，villous or blades blades wider than long; tepals always subglabrous ，ciliate; segments petiolulate or

monomorphic ， white or yellowish ， with subsessile; central segments 3-parted to 3 田 anastomosing anastomosing veins or without them ，pubes- cleft ，rhombic-ovate or broadly rhombic ，ul- cent cent or glabrous; achenes with styles 0.8-1.2 timate lobules ovate to linear; lateral

mm long ....・ H ・....・ H ・....・ H ・. 1. A. narcissiflora segments unequa lI y 2-3 ・parted ， oblique:- 7b. 7b. Stems 3-5 ，ascending; basal leaf blades flabellate. Scapes 1-2 ， 10-50 cm long ， longer longer than wide ，tepals sometimes dimor- vi lI ous or sparsely puberulent; cymes phic ， mainly blue or pink ， with anastomosing 1-5 (-7)-flowered. lnvolucral leaves 3 or veins ，pubescent; achenes with styles 1.8-2.2 more; blades 3-p 紅白d to 3-cleft ， broadly

mm long ....・ H ・...・ H ・...・ H ・.....・ H ・.. 2. A. demiss α rhombic to na 汀 owly obovate ，2-6 cm long ， 8a. 8a. Basal leaf blades 3-1obed ，coriaceous; mainly pubescen t. Pedicels 2-5 ( -8) cm long ， tepals tepals 4， 12-20 x 6-12 mm ， with 3-5 pubescen t. Tepals 5-6(-7) ， white or yel- anastomosing anastomosing veins ，glabrous; achene bodies lowish (rarely pinkish) ，elliptic-obovate ， with

8-12 mm long ，styles 1. 5-2 mm long ....・ H ・- rounded narrowed apices and bases ， mainly …. 7. A. tetrasep α la monomorphic (rarely dimorphic ， e. g. ， in 8b. 8b. Basal leaf blades 3-parted ，herbaceous; some populations in Japan) ， 12-18 x 6-10 tepals tepals 5-10 ， of various sizes ， with or without mm ，sparsely puberulent or glabrous; basal anastomosing anastomosing veins ，puberulent or glabrous; veins 3-7 ， anastomosing veins 3-5 or absen t. achene achene bodies and styles smaller than above Stamens 2-5 mm long; filaments linear; an- ..9 ..9 thers oblong. Ovaries subspherical-ovoid ， 9a. 9a. Tepals 5-10 ， reflexed ， with 7-9 compressed ， mainly glabrous ， having distinct anastomosing anastomosing veins ， sparsely puberulent; lateral wings; styles short ，bent ， 0.5-1.2 mm achenes achenes sparsely puberulent ， with curved long (Figs. 1a-h). Achene bodies obovate ，

styles styles ...・ H ・.....・ H ・H ・H ・..…...・ H ・. 8. A. polyanthes 5-8 x 4-6 mm ，glabrous (rarely with solitary 9b. 9b. Tepals 5，straight ， without anastomosing hairs) ， wings 0.8- 1. 4 mm wide; styles veins ， glabrous or puberulent; achenes hooked ， 0.8-1 .4 mm long (Figs. 2a-h).

glabrous ， with substraight styles ….....・ H ・. 10 Achene surface folded ，tuberculate or r紅 ely 10a. 10a. Tepals 12-20 x 6-12 mm ， basally wrinkled (Figs. 3a-h). cuneate ，apically rounded; achene bodies 6- Distribution and habita t: Europe ， Central 9 mm long ， with wings 1.2-1.3 mm wide Asia ， Far East ， Eastern China ， North and and styles 1-1.5 mm long …. 9. A. elongata America (U. S. A. and NW Canada) ， al t. 10b. 10b. Tepals 10-12 x 4-6 mm ， basally 600-3000 m ， in mountain meadows and rounded ，apically acute; achene bodies 5-6 edges of forests. mm long ， with wings 0.5-0.6 mm wide and Notes: Linnaeus (1753: 542) originally de- styles styles 0.6-1 mm long …… 10. A. shikokiana scribed A. narcissiflora as plants having umbellate umbellate inflorescences and ovate com- 1. 1. Anemone narcissiflora L. ， Sp. Pl. 1: pressed glabrous fruits. We found within the 542 (1 753) - Anemonastrum narcissiflorum LINN collection of types four specimens of (L.) (L.) Holub in Folia Geobo t. Phytotax. A. narcissiflora ， and we reg 訂 d the first of (Praha) (Praha) 8: 165 (1 973). Lectotype (desig- these specimens ， having the note “ A. October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 291 multiflora multiflora (N.710.31) ， as the Linnean nized two subspecies of A. narcissiflora ， holotype. holotype. It is a with two stems 22 and subspp. narcissiflora and biarmiensis (Juz.) 25 25 cm long ， four basal leaves on petioles 4- Jalas. But Hermann (1 956) ， Zimmerman 6 cm long ， and 3-sected blades without (1 975) ，Pignatti (1 988) and others did not petiolules ， four involucral leaves and 4-5 recognize any subspecies and varieties of A. flowers flowers with 5 or 6 white tepals 17-18 x 8- narcissiflora in their regional floras of 9mm. Central and South Europe. Anemone narcissiflora has long been re- Based on an examination of about 1500 garded garded as a variable species ， since de herbarium specimens in his treatment of the Candolle Candolle (1 817 ， 1824) describing five varie- A. narcissiflora complex ，Shipchinski (1 912) ties: ties: va r. fasciculata DC. with fasciculate proposed to recognize 10 varieties and five peduncles ， occurring in Cappadocia; va r. forms in the flora of Eurasia. He accepted monanth αDC. with solitary flowers ， occurs four varieties of A. narcissiflora which were in in mountains of Europe ， Asia and N orth previously described by other authors (viz. ， America; America; var. villosissima DC. with villous vars. laxa Ulbr. ， chrysantha Ulbr. ，linea- stems stems and subsessile flowers ， occurs in the riloba Regel & Radde and turkestanica Aleutians Aleutians to Unalaska; va r. pedicellaris DC. Lipski). He also described six additional va- with with long pedicels ， and var. frigida DC. with rieties (vars. linneana Schipcz. ， alaschanica glabrous glabrous basal leaves has linear ultimate lob- Schipcz. ， caucasica Schipcz. ， uralensis ules ules with no geographic range given. Schipcz. ，yuldussica Schipcz. and pekinensis Ulbrich Ulbrich (1906) recognized within A. Schipcz.) ， as well as five forms (f. protracta narcissiflora narcissiflora two subspecies ，two varieties Schipcz. ，f. contracta Schipcz. ，f. glabra and and one form: subspp. typica Ulb r. and Schipcz. ， f. villosissima Schipcz. ， and f. chrysantha chrysantha Ulbr. ，vars. genuina Ulb r. and superba Schipcz.). protracta protracta Ulbr. ， and f. laxa Ulbr. Within “ Flora URSS" Juzepchuk (1 937) Zapalowicz Zapalowicz (1 908) described three varie- elevated four previously described varieties ties ties and four forms of A. narcissiflora in the and forms of A. narcissiflora to species flora flora of Poland (including a part of W. level: A. laxa (Ulb r.) Juz. ， A. protracta Ukraine Ukraine to Galicia): v訂 s. podolica Zapa l.， (Ulb r.) Juz. ，A. νillosissima (DC.) Juz. ， A. zawratensis zawratensis Zapa l. and pienina Zapa l.， f. sachalinensis (Miyabe & Miyake) Juz. In ad- humifusa humifusa Zapa l.， f. latisepala Zapa l.， f. dition ， Juzepchuk described six species monantha Zapa l. and f. macrantha Zapa l. within this complex: A. bi α rmiensis Juz. ， A. Notwithstanding Notwithstanding these early treatments ， brevipenduculata Juz. ， A. calva Juz. ， A. subsequent subsequent taxonomists have not been uni- crinita Juz. ， A. impexa Juz. and A. form form in recognizing the variability within A. schrenkiana Juz. narcissiflora. narcissiflora. For example ，Hayek (1927) ac- It is remarkable that in the flora of Turkey ， cepted cepted three varieties in the flora of the Davis et al. (1965) accepted A. narcissiflora Balkans Balkans (vars. monanth αDC. ，fascicul α ta as including two subspecies: subspp. DC. and elatior (Schur) Beck). More narcissiflora and willdenowii. They regarded recently ， Pawlowski (1 956) proposed that subsp. narcissiflora as co 町 esponding to A. plants plants referable to A. narcissiflora in the fasciculata L. and subsp. willdenowii to A. mountains mountains of Central Europe be recognized umbellata Willd. as as subsp. europaea Paw l.， but only Futak After revising the Eastern Asian taxa of (1 983) accepted this subspecies in the “Flora the A. narcissiflora complex ，Tamura (1 958) of of Slovakia." Within the flora of Europe ， recognized four of Juzepchuk' s species at the both both Tutin (1 964) and Chater (1 973) recog- variety level: vars. villosissima DC. ，sibiric α 292 292 植物研究雑誌第80 巻第5号 平成17 年10 月

(L.) (L.) Tamura ， crinita (Juz.) Tamura and Nelson) Dutton ，Keener & Ford]. brevipenduculata brevipenduculata (Juz.) Tamura). In addi- The generally accepted diagnostic charac- tion ， within the flora of Japan and Sakhalin ， ters within the A. narcissiflora complex in- Tamura described two varieties of A. clude tepal number ， size and colour ， achene narcissiflora: narcissiflora: vars. nipponica Tamura and size ， flowering stem variation (several- or japonomonantha Tamura. Moreover ， within one-flowered) ，stem and pedicel length ， leaf the the flora of Japan Ohwi (Kitagawa) (1 982) shape ，and plant pubescence. According to accepted accepted A. narcissiflora var. sachalinensis our recently published results (Ziman et al. Miyabe & Miyake. 1997) ，most of these characters are too vari- Wang (1980) accepted A. narcissiflora as able even within the same natural popula- an an aggregate species having four varieties tions to be of any significant diagnostic use. (vars. (vars. protracta Ulbr. ，crinita (Juz.) Tamura ， The results of our study of ontomo 中ho- sibiric α(L.) Tamura and turkestanica geny within 19 populations of eight races of Schipcz.). Schipcz.). According to Wang and Li (1 986) the A. narcissiflora complex (Ziman et al. and Wang (1 996) ， three new endemics in the 1997) have shown that everywhere the gen- flora flora of China are very close to A. eral scheme of ontomorphogeny exists ，and narcissiflor α. Thus ， A. multilobulata differs all populations 紅 e very similar in terms of from A. narcissiflora in having imbricate peculiarities of their life cycle. segments segments of basalleaf blades and five yellow Within the characters of anatomical struc- tepals ， A. nutantiflora with nutant pedicels ， ture of basal leaf petioles ，additional diag- and A. tengchongensis with papery blades of nostic characters include the a町 angement of basal basal leaves and larger tepals (18-20 x 8-10 vascular bundles ，number of large bundles ， mm). However ，all essential characters are in and sometimes peculiarities of collenchyma. common with A. narcissiflora ，and besides ， We note within the A. narcissiflora complex data data on the flowers of A. nutantiflora and the distinctions in size and number of tepals achenes achenes of A. tengchongensis are absen t. and their venation (especially anastomosing Therefore ，we have no basis for recognizing veins). any of these taxa as species ，and we did not All plants of the A. narcissiflora complex accept accept them in the modern version of the possess similar achene characters including “ Flora of China" (Wang ，Ziman and Dutton the morphological peculiarities mentioned 2001). 2001). above ， and the anatomical s住ucture of As a result of a critical examination of the pericarp and testa. However ， there are certain flora flora of Alaska and a ‘jacent territories ，Hul- distinctions in the size of achene bodies and ten ten (1 944) regarded A. narcissiflora as hav- their wings ， the microsculpture of the achene ing ing four subspecies: subspp. sibirica (L.) surface ， the number of layers of the endoca 中 Hulten ，villosissima (DC.) Hulten ，interior and the thickness of the endoca 中 and testa Hulten Hulten and alaskana Hulten ，which differed (their indices). mainly mainly by the shape of the basal leaf blades ， These peculiarities of the generative and and the number and colour of flowers . In the vegetative organs have been of considerable annotated annotated list of the N orth American Flora use in separating the infraspecific taxa (sub- Kartesz Kartesz (1994) cited all of them ， but in the species and varieties) of the A. narcissiflora treatment treatment of Anemone in “ Flora of N orth complex. As a result of our examination ，we America" Dutton et al. (1997) recognized have concluded that the A. nαrcissiflora three three of the above taxa [as vars. monantha complex is one aggregate species disjunc- DC. (instead of subspp. sibirica and tively distributed in Eurasia and North alaskana) ，villosissima DC. and zephyra (A. America that includes eight subspecies and October October 2005 Joumal of Japanese Botany Vo l. 80 No. 5 293

varieties. 12 varieties. one endocarp layer; achene surface folded;

Caucasus Caucasus .....・ H ・.....・ H ・... b. subsp. fasciculata Conspectus Conspectus of Anemone narcissiflora 4b. Petiolules of the central segment of basal 1a. 1a. A. narcissiflora L. subsp. narcissi- leaf blades mainly longer than those of the flora. flora. Vars. narcissiflora ， podolica Zapa l.， lateral segments; achenes with (1-) 2-3 en- crimea crimea Ziman & Fedoronchuk docarp layers; achene surface wrinkled- b. 1b. A. narcissiflora L. subsp. fasciculata folded; Central Asia …… c. subsp. protracta (L.) (L.) Ziman & Fedoronchu k. Vars. 5a. Tepals 5-7 ，8-15 x 6-10 mm; Siberia ... fasciculata fasciculata (L.) DC. ，subun{ β ora C. A. Mey. ， .. d. subsp. crinita willdenowii willdenowii Boiss. 5b. Tepals 5-9 ， of various sizes; Far East and

1c. 1c. A. narcissiflora L. subsp. protracta Alaska ....・ H ・-……...... ・ H ・-… H ・H ・.....・ H ・.6 (Ulbr.) (Ulbr.) Ziman & Fedoronchuk. Vars. 6a. Plants villous; tepals 15-25 x 10-15 mm; protrac ωUlb r.， contracta (Ulbr.) Scipcz. achene surface folded-tuberculate; Kuriles ，

d. 1d. A. narcissiflor αL. subsp. crinita (Juz.) Aleutes ，partly Sakhalin and J apan …H ・H ・-

Kitag. Kitag. Vars. crinita (Juz.) Tamura ， …...・ H ・.. e. subsp. villosissim α lineariloba lineariloba Regel & Radde ， uralensis 6b. Plants more or less scarcely puberulent; Schipcz. ，sachalinensis Miyabe & Miyake tepals 10-15 x 8-12 mm; achene surface

1e. 1e. A. narcissiflora L. subsp.ν illosissima without tubercules ・・ H ・H ・-…...・ H ・....・ H ・....・ H ・...7 (DC.) (DC.) Hulten 7a. Achene with wings 1- 1. 5 mm wide and 1f. 1f. A. narcissiflora L. subsp. alaskan α styles 0.9- 1. 1 mm long; achene surface Hulten Hulten folded; Alaska and Yukon .…・・ Ig. Ig. A. narcissiflora L. subsp. zephyra (A. .. f. subsp. alaskana Nelson) Nelson) B. E. Dutton 7b. Achenes with wings 0.5-1.2 mm wide 1h. 1h. A. narcissiflora L. subsp. nipponica and styles 1. 4- 1. 6 mm long; achene surface (Tamura) (Tamura) Kadota wrinkled- folded; J apan … h. subsp. nipponica

Key to infraspecific taxa of 1a. A. narcissiflora L. subsp. narcissi- Anemone narciss すlora flora 1a. 1a. Tepals pilose ， with anastomosing veins; Tepals pilose ， 5-7 ， 12-20 x 7-15 mm ， achene achene surface folded (sometimes tepal anastomosing veins mainly 3-7.

tuberculate- or wrinkled-folded) ....・ H ・...・ H ・.2 Achene bodies 6-7 x 4-5 mm ， with one en- b. 1b. Tepals glabrous ， anastomosing veins ab- doca 叩 layer; achene surface folded. sent sent (rarely solitary); achene surface 孔10untains of Europe.

tuberculate tuberculate (rarely folded) ....・ H ・-… H ・H ・...・ ...5 1. var. narcissiflora 2a. 2a. Tepals 5-7 ，12-20 x 7-15 mm; achene A. narcissiflora subsp. typica Ulb r. in Bo t.

bodies bodies 6-7 x 4- 5mm .…...・ H ・....・ H ・...・ H ・...・ H ・.3 Jahrb. 37: 273 (1 906). 2b. 2b. Tepals 5-6 ， 10-15 x 6-8 mm; achene A. narcissiflora var. linneana Shipcz. in

bodies bodies 5-6 x3 -4 .5mm …H ・H ・....・ H ・...・ H ・...・ ...4 Tr. Bo t. Sada Jurjev Univ. 95 (1 912). 3a. 3a. Anastomosing veins mainly 3-7; achene A. narcissiflora va r. pienina Zapa l.，

surface surface folded ， with folds p紅 allel to achene Consp. Fl. Galic. Cri t. 2: 231 (1908).

axis; axis; Europe ...・ H ・..…. a. subsp. narcissiflora A. narcissiflora va r. zawratensis Zapa l.， 3b. 3b. Anastomosing veins 1-3; achene surface Consp. Fl. Galic. Cri t. 2: 231 (1908). tuberculate-folded; tuberculate-folded; U. S. A. ，Rocky Moun- A. narcissiflora subsp. europ αωPaw l.， Fl.

tains tains ...・ H ・.....・ H ・.....・ H ・H ・H ・... g. subsp. zephyra Tat r. 1: 281 (1 956). 4a. 4a. Petiolules of all basalleaf blade segments Plants are characterized by predominately distinct ，more or less similar; achenes with straight basal leaf lobes ， 3-7 basal tepal 294 植物研究雑誌第80 巻第5号 平成 17 年 10 月 veins ， with 1-3 anastomosing veins ，vari- of basal 1eaf blades distinc t. A 町 angement of ous1y ous1y shaped outline of petio1ar cross sec- petio1ar vascular bund1es radia1 or transi- tion ， and predominate1y dorsiventra1 tiona l. Tepa1s pilose ， with numerous a町 angement of petio1ar vascu1ar bund1es. anastomosing veins. Achenes with one 1ayer Dist r.: C. ， E. and S. Europe (Spain ， of endoca 叩， achene surface main1y fo1ded ， France ，Ita1y ，Bu1garia ，Germany ，Austria ， the folds situated both a10ng and across Switzerland ， Rumania ， Czech ， Slovakia ， achene axis. Po1and ，and Ukraine). Dit r.: Caucasus ( ， Georgia ， Armenia) Armenia) and Turkey. 2. 2. var. podolica Zapa l.， Consp. Fl. Ga1ic. 4. var. fasciculata (L.) DC. ，Prod r. 1: 212 Cri t. 2: 231 (1908). Type: UKRAINE: (1 817).

Voh1ynj Voh1ynj (former Zas1av1 County) ， Ska10zub A. fasciculata var. longipedunculat αL.M.， (LE !). Kem.-Na t.， Fl. Gruzia 3: 239 (1 975).

A. A. narcissiflora f. laxa U1b r. in Bo t. Jahrb. P1ants several 四 flowered ，tall ， with dense1y 37: 37: 273 (1 906) - A. narcissiflora var. laxa pubescent ， white tepa1s 18-25 x 12-18 mm. (U1b r.) Shipcz. in T r. Bo t. Sada Jurjev Univ. Large vascular petio1e bundles 7-11. Achene 95 95 (1 912) - A. laxa (U1b r.) Juz. ， F l. URSS surface folded. 7: 7: 209 (1937). Distr.: Caucasus (Russia ， Georgia) ， in P1ants P1ants differ from those of va r. margins of forests and suba1pine meadows ， narciss 伊ora by predominate1y falcate basa1 a1 t. 1800-2500 m. 1eaf 1eaf 10bes ，7-9 tepa1 veins ，5-9 anastomosing 5. var. subuniflora C. A. Mey. ， Verz. veins ，rounded-pentagona1 outline of petio1ar Ko1en. (1 849). Type: unknown. cross cross section ， and radia1 a町 angement of A. speciosa Adams ，Beit r. N atu r. Weber petio1ar petio1ar vascu1ar bund1es. Mountains of C. Mih r. 1 (1 805). Europe ， Subcarpathia and Podo1je. A. chrysopetala Rup r.， Fl. Cauc. 12 (1 869). 3. 3. va r. crimea Ziman & Fedoronchuk ， A. narcissiflora subsp. chrysantha U1b r. in Taxon. Taxon. Evo l. Anemone nαrcissiflora com- Bo t. Jahrb. 37: 273 (1906). p1ex p1ex 34 (1 997). Type: UKRAINE: Crimea ， A. narcissiflora L. va r. caucasica Shipcz. Nikitskaya Nikitskaya Yaila ， Kr asnyi Kamenj ， in T r. Bo t. Sada Jurjev Univ. 95 (1 912). 14.8.1990 14.8.1990 Ziman (KW !) A. chrysantha Grossh. ， F l. Cauc. 2: 105 P1ants P1ants are simi1ar to va r. podolica ， but diι (1 939). fer fer by the rounded outline of the petio1ar A. narcissiflora L. subsp. fasciculata (L.) cross cross section ， 1arger number of 1arge 1eaf Ziman & Fedoronchuk va r. speciosa petio1e petio1e vascu1ar bund1es (5-9 and 9-11) ，and (Adams) Ziman & Fedoronchuk Taxon. reduced reduced collenchyma. Evo l. Anemone narcissiflora comp1ex 34 Dit r.: Ukraine ， Mountains of Crimea (1 997). (Nikitskaja (Nikitskaja Yai1a). Plants with solitary flowers and yellow scarcely scarcely pubescent tepa1s. Large vascu1ar b. 1b. A. narcissiflora L. subsp. fasciculata petio1e bund1es 3-5. Achene surface fo1ded. (L.) (L.) Ziman & Fedoronchuk ， Taxon. Evo l. Distr.: Caucasus (Russia ， Georgia) ， in Anemone narcissiflora complex 34 (1997) - subalpine and a1pine meadows ， a1 t. 2500-

A. A. fasciculat αL.， ， Sp. Pl. 542 (1 753). 3500m. Lectotype Lectotype (designated here): “ Habitat in 6. va r. willdenowii Boiss. ， F l. Orien t. 1: Oriente". Oriente". N.710.20 (LINN !). 14 (1 867) - A. narcissiflora subsp. willdeno- Petiolu1es Petiolu1es of centra1 and 1atera1 segments wii (Boiss.) Davis ，No t. Roy. Bo t. Gard. October October 2005 Journal of JapanesD Botany Vo l. 80 No. 5 295

Edinb. Edinb. 26: 175 (1 965) - A. narcissiflora basal leaf blades shortly petiolulate. subsp. subsp. willdenowii (Boiss.) Greuter & Burdet Dist r.: Mountains of C. Asia (Kazakhstan in in Willdenowia 19: 43 (1989) ， nom. 叩supe 町rfl. and W. China). Type: Type: Paratype 広: ARMENIA: Alagoe 白z， Dali-Chai 1d. A. narcissiflora L. subsp. crinita River ，Khadra-Mugum-Jurt ，18.7.1832 ，Bush (Juz.) Kitag. ， Lineam. Fl. Manch.: 213 (BM !). (1 939) - A. crinita Juz. ， F l. URSS 7: 209 A. A. umbellata Willd. ， Sp. Pl. 2: 1284 (1 937). Lectotype (designated here): “Habi- (1 799). tat in Sibiria. Gmelin". N.710.18 (LINN !). A. A. impexa Juz. ， F l. URSS 7: 209 (1 937). A. sibirica L. ， Sp. Pl. 538 (1 753) - A. Plants Plants differ from those of va r. speciosa sibirica L. ， Sp. Pl. 541 (1 753) - A. mainly mainly by their linear lobules of basal and narcissiflora L. subsp. sibirica (L.) Hulten ， involucral involucral leaves ， few-flowered stems and Fl. Aleu t. Is l. Alaska 4: 734 (1944) - A. tuberculate- folded achene surface. narcissiflor αsubsp. sibirica (L.) A. Love & Distr.: Distr.: Transcaucasia (Armenia) ，Turkey ， D. L りve in Bo t. Notiser 128: 511 (1 975) ， in in rocky slopes ， al t. 2400-3100 m. nom. superfl. - A. narcissiflora L. var. sibirica sibirica (L.) Tamura in Acta phytotax. c. 1c. A. narcissiflora L. subsp. protracta Geobo t. 17: 115 (1 958). Type: RUSSIA: (Ulb r.) Ziman & Fedoronchuk Taxon. Evo l. Altai (LE !). Anemone narcissiflora complex 34 (1 997). Tepals 5-7 ， 8-15 x 6-10 mm ， glabrous Petiolules Petiolules of the central segment of basal (rarely subglabrous) ， anastomosing veins ab- leaf leaf blades mainly longer than those of the sent (rarely solitary). Achene surface tuber- lateral lateral segments. Tepals pilose ， with few culate (except in plants from Sakhalin). anastomosing anastomosing veins. Achene surface wrin- Dist r.: Russia (Siberia ， Far East ，Kam- kled-folded kled-folded (small folds situated irregularly). chatka and Sakhalin) ，China ， Mongolia and Dist r.: Tadzhikistan. Kazakhstan and W. N. Korea. China. China. 9. var. CI ・inita (Juz.) Tamura in Acta 7. 7. va r. protracta Ulbr. in Bo t. Jahrb. 37: Phytotax. Geobo t. 17: 115 (1 958). 273 273 (1 906) - A. protracta (Ulb r.) Juz. ， F l. A. crinita Juz. var. alpina A. V. Polozhii URSS 7: 209 (1 937). Type: unknown. in Sys t. Zame t. Mate r. Herb. Toms k. Univ. A. A. narcissiflora L. var. turkestanica Lipski 85: 6 (1 974).

in in Tr. Bo t. Sada Ju りev. Univ. 13: 101 Plants with several 同 flowered cymes and (1 912). shortly petiolulate basal leaf blades. Tepals Tepals densely pilose; basal leaf blades A 町 angement of petiolar vascular bundles ra- long-petiolulate. long-petiolulate. dia l. Achene surfaced tuberculate. 8. 8. var. contracta (Ulbr.) Shipcz. in Tr. Dist r.: Russia (Siberia and Far East) ， Bo t. Sada Ju りev Univ. 95 (1 912) - A. China and Mongolia. narcissiflora narcissiflora L. f. contracta Ulbr. in Bo t. 10. va r. Ii neariloba Regel & Radde in Jahrb. Jahrb. 37: 273 (1 906). Lectotype (designat- Acta Hor t. Petropo l. 8 (1884). Type: ed ed here): KAZAKHSTAN: Dzhungar Ala unknown. Tau (LE !). A. αconitifoli αTurcz. in Bull. Soc. Na t.

A. A. nαr 陀ci おs叫flora αL. va r. yuldussic αShipcz. Mosc. 61 (1 840). Lectotype (designated in in Tr. Bo t. Sada Jurjev Univ. 95 (1 912). here): “Inter Jacutian et Ochotian ， ad florae A. A. schrenkian αJuz. ， F l. URSS 7: 209 Aldan" ，1835 ， Turczaninov (BM !). (1 937). A. brevipedunculata Juz. ， F l. URSS 7: 209 Tepals Tepals scarcely pubescent or subglabrose; (1 937) - A. narcissiflora var. bre νipeduncu- 296 296 植物研究雑誌第80 巻第5号 平成17 年10 月

lata lata (Juz.) Tamura in Acta Phytotax. Geobo t. Lopatina). 17: 17: 115 (1 958). Lectotype (designated here): Dist r.: Russia (Sakhalin) and Japan RUSSIA: Far East ，Ussuri ， Tyutikhe River (Hokkaido and northen Honshu). valley valley (LE!). A. A. calva Juz. ， F l. URSS 7: 209 (1 937) - A. 1e. A. narcissiflora L. subsp. villosissima narcissiflora narcissiflora subsp. cal να(Juz.) A. L りve & (DC.) Hulten ， F l. Aleu t. Is l. Alaska 4: 734 D. D. L りve in Bo t. Notiser 128: 511 (1 975). (1 944) - A. narcissiflora var.ν illosissima Lectotype Lectotype (designated here): RUSSIA: Far DC. ， Prodr. Sys t. 212 (1 817) - A. villosis- East ， mountains between Yakutsk and sima (DC.) Juz. ， F l. URSS 7: 209 (1 937) -A. Okhotskoe Okhotskoe (LE !). narcissiflora L. subsp. villosissimum (DC.) A. A. narcissiflora L. va r. sibirica (L.) A.L りve & D. Love in Bo t. Notiser 128: 511 Tamura in Acta Phytotax. Geobo t. 17: 115 (1 975) ，nom. superfl. Type: U. S. A.: the (1 958) ， p. p. exc l. type. Aleutians ， Unalaska Island (GE). Plants Plants with l-few-flowered cymes and Plants villous. Tepals glabrous [sometimes sessile sessile primary segments of basal leaves. pubescent - e. g. ， in plants from S. Aleutes ， Leaf Leaf ultimate lobules ovate to narrowly Kagamil ，2.7.1944 ， Miller 1045 (US)] ，5-9 ， ovate. ovate. Arrangement of petiolar vascular bun- 15-25 x 10-15 mm ， without anastomosing dles dles radia l. Achene surface tuberculate. veins (rarely solitary); achenes with wings Distr.: Distr.: Russia (Siberia ， Far East) and N. 1-2 mm wide and styles 1-2 mm long; Korea. Korea. achene surface folded-tuberculate (small 11. 11. var. uralensis Shipcz. in Tr. Bo t. Sada folds situated irregularly ， but also Ju りev Univ. 95 (1 912). Lectotype (desig- tubercules). nated nated here): RUSSIA: Ural ， M t. Yurma (for- Dist r.: Russia (Kuriles and partly mer Perm Province) (LE !). Sakhalin) ， U. S. A. (Aleutians) and Japan. A. A. biarmiensis Juz. ， F l. URSS 7: 209 (1 937). lf. A. narcissiflora L. subsp. alaskana A. A. narcissiflora subsp. biarmiensis (J uz.) Hulten ， F l. Aleu t. Is l. Alaska 4: 734 (1 944). J. J. Jalas in Ann. Bo t. Fenn. 25: 297 (1 988). Type: unknown. Plants Plants with several-flowered cymes and A. nαrcissiflora subsp. sibirica (L.) Hul- petiolulate petiolulate basal leaves. A 町 angement of ten ， F l. Aleu t. Is l. Alaska 4: 734 (1 944) ，p. p. petiolar petiolar vascular bundles transitiona l. exc l. type. Achene surface tuberculate. A. narcissiflora subsp. interior Hulten ， F l. Dist r.: Russia (Ural). Aleu t. Is l. Alaska 4: 734 (1 944). 12. 12. var. sachalinensis Miyabe & Miyake ， Plants more or less scarcely puberulent; l. Fl. Saghalin 10 (1 915) - A. sach αlinensis tepals 5-9 ， glabrous ， 10-15 x 8-12 mm ， (Miyabe (Miyabe & Miyake) Juz. ， F l. URSS 7: 209 without anastomosing veins; achenes with (1 937) - A. narcissiflora subsp. sachalinen- wings 1- 1. 5 mm wide and styles 0.9- 1. 1 mm sis sis (Miyabe & Miyake) Ziman & long; achene surface folded. Fedoronchuk ， Taxon. Evo l. Anemone Dist r.: U. S. A. (Alaska) and Canada (Yu- narcissiflora narcissiflora complex 34 (1997). Type: un- kon). known. Plants Plants with few-flowered cymes and Ig. A. narcissiflora L. subsp. zephyra petiolulate petiolulate basal leaf blades. Arrangement of (A. Nelson) A. L りve ， D. L りve & Kapoor ， petiolar petiolar vascular bundles dorsiventra l. Arctic Alp. Res. 3: 149 (1 971) - A. zephyra Achene surface folded (e. g. ， M t. Chekhova ， A. Nelson in Bo t. Gaz. 42: 51 (1 906) - A.

Shmidt Peninsula) or tuberculate (e. g. ，M t. narcis sifio rαsubsp. zephyr α(A.， Nelson) W. October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 297

A. Weber in Phytologia 41: 486 (1 979) ， 8.1990 ， Kobiv (KW); Nesamovite Lake ，30.7.1987 ， nom. superfl. - A. narcissiflora var. zephyra Ziman (KW); M t. Turkul ，27.7.1987 ， Fedoronchuk (KW); Mt. Brescul ，24.7.1987 ， Fedoronchuk (KW); (A. (A. Nelson) B. E. Dutton & Keener in Svidovets Svidovets Range ， M t. Bliznitsa ， 17.6.1986 ，Ziman Phytologia 77: 85 (1 994). Type: unknown. (KW; 28.6 .1 987 ，Ziman & Dutton (KW); Marmarosh Described from C. Rocky Mountains. Range ， M t. Pip Ivan Marmaroshski ，17.6.1986 ，Ziman Tepals 5-7 ， pilose ， 12-20 x 7-15 mm ， (KW); Chivchino-Grynjavy Range ， M t. Chivchin ， with few anastomosing veins; achene bodies 27.7.1989 ，Ziman (KW). 6-7 x 4-5 mm. achene surface folded- SLOVAKIA: Belanskie Tatry ， Velicka Dolina ， 20.7.1892 ，Wiany (LE); Lysanki ， 1913 (KRAL KRA); tuberculate. tuberculate. t. M t. B 吋ach ，7.1939 ， Deyl (BRA); 16.7.1967 ，Vasak Dist r.: U. S. A. ，Rocky Mountains (Mon- (BRA); Tristarska Dolina ，14.7 .1 946 ， Futak (BRA); tana ，Wyoming ，Colorado). Havran ， 13.8.1959 ， Klasterski (BRA); Zadne Medodoly ，Zdiarska Viedla ，27.6 .1 961 ， Dostal (BRA); lh. lh. A. narcissiflora L. subsp. nipponica Belanska Kopa ， 26.6.1963 ， Dostal (BRA); Zdiar ， Vidla ， 22.6.1969 ， Sojak (BRA); Coprske Sedlo ， (Tamura) Kadota in K. Iwats. & al.， F l. Jap. 1. 8.1990 ，Ziman & Siposova (KW); Velka Fatra ， IIa: IIa: 66 (2005) - A. narcissiflor αva， r. nippo ・ Kryzna ， 21. 5.1 923 ， Nabelek (BRA) ， 27.6.1987 ， nica nica Tamura in Acta Phytotax. Geobo t. 17: Marhold (KW); Mala Fatra ， Chleb ， 1943 ， Futak 115 115 (1 958). Type: JAPAN: Honsu ，Prov. (BRA); Zapadny Tatry ， Lyptovske Hole ， Cervene Iwashiro ， M t. Iide ， 1910 (KYO !). Vrchy ， 14.7.1954 ， Sourek (BRA); Nyzkie Tatry ， Dumbier ，1. 8.1991 ， Stepanek (KW). A. A. narcissiflora v訂 jα~ponomonantha CZECH: Sudety ， Hruby Jesenik ， Peterstein ， Tamura in Acta Phytotax. Geobo t. 17: 115 12.7.1920 ， Swardl (PRG); Velka Kotlina ，26.6.1960 ， (1 958). Dostal; 30.5 .1 969 ， Deyl (PRG); Vysoki Jeseniky ， A. A. narciss 伊ora var. frigida DC. ， va r. Velka Kotlina ，27.7.1932 ， Veber (PRG); Krknose ， monantha DC. et va r. pedicellaris DC. ， Snezne Jamy ，1. 7.1925 ， Dostal (PRG); Vesselgruh ， Prodr. Prodr. 1: 212 (1 817). 26.8.1929 ，Wihan (PRG); Cerlov u zanidku ，1930 ， Tratler Tratler (PRG); Brunnberg ， 7.7.1931 ， Japp (PRG); A. A. narcissiflora L. va r. al α schanica Zabske Zabske louca ， 1460 m ， 11. 6.1954 ， Dostal (PRG); Schipcz. et var. pekinensis Shipcz. in Tr. Ripska Stena ， 1450 m ， 19.6.1954 ， Dostal (PRG); Bo t. Sada Jurjev Univ. 95 (1 912). Kr einen Teich ， 11. 6.1958 ， Baenitz (PRG); Velka Plants Plants more or less scarcely puberulen t. Kotelna Jama ， 12.7.1960 ， Shiffner (PRG); Ulohe Tepals 5-9 ， 10-15 x 8-12 mm ， glabrous ， Hleive ， 4.7.1977 Uborg (PRG); Gesenka ，4.6.1986 ， without anastomosing veins (rarely solitary). Shiffner (PRG); M t. Rykhory ， 18.7 .1 990 ，Ziman & Zaverukha Zaverukha (KW). Achenes with wings 0.5-1.2 mm wide and ROMANIA: Transsylvania ，Retezat ，乱 1t. Borescu ， styles styles 1.4 -1.6 mm long; achene surface 12.7.1924 ， Nyarady (CLU 1); Retezat ，Petrosani Dist r.， wrinkled-folded (small folds situated irregu- M t. Piu1a ， 22.7.1956 ， Cauros; 16.7 .1 971 ， Coldea larly). larly). (CLUJ); Rodnenses Mts. ，Bistrita-Nasaul ，Korongyis ， In several populations dimorphic tepals 7.8.1925 ， Borza (CLUJ); Nagy-Korongyis ，10.7.1942 ， Antonius Antonius & Nyarady (CLUJ); Muntenia Distr. ， with two whorls were noted (outer sepals Prahova ，2 1. 7.1925 ， Borza (CLUJ); Brasov Dist r.， larger larger and puberulent ，inner ones smaller and M t. Bucegi ， 4.8.1930 ， Nyarady (CLUJ); Caraiman ， glabrous). glabrous). Valea Jepilor ， 25.6 .1 939 ， Constantinescu (CLUJ); Dist r.: Japan (central Honshu). Bihor ， Stana de Vale ，Bohodei ，27.8.1939 ， Borza (CLUJ); (CLUJ); Banatus ，Severin Dist r.， M t. Tarcu ，25.7.1942 ， Borza Borza (CLUJ); Mts. Fagarasului ， M t. Clcapatrei ， List List of Representative Specimens Examined 11. 8.1973 ， Voik (CLUJ); 乱1t. Piatra ， Kr ayul ， subsp. subsp. narcissiflora 25.6.1998 ，Ziman ， Tsarenko (KW). va r. narcissiflora POLAND: Beskidy ， Babia Gora ， Kosciolki ， UKRAINE: E. Ca 中athians ， Chernogora Range ， 30.6.1932 ， Szafer 118 (LE). M t. Petros ， 14.7 .1 986 ， Kobiv (KW); M t. Goverla ， AUSTRIA: Alps ，加1t. Schneeberg ， 26.5 .l 868 ， 1. 7.1948 ， Bylik (KW); M t. Pip Ivan Chernogorski ， Woloszak (KRA); 6.8.1991 ，Ziman & Ehrendorfer 298 298 植物研究雑誌第80 巻第5号 平成17 年 10 月

(KW); Gauswicsen ，27.6.1869 ， Felkel (KRA); M t. RUSSIA: N Caucasus ，Teberda ，M t. Malaya Khatipara ， Oetscher ， 6.8.1990 ， Ehrendorfer (KW); M t. 15.6.1978 ， Zivan (KW); Dombai ，M t. Musa-Achtara ， Untersberg ，6.8.1991 ，Ziman & Ehrendorfer (KW). 1. 7.1989 ，Ziman & Fedoronchuk (KW); Amanauz SWITZERLAND: 乱1ontreux ， 24.5.1906 ， Ravine ，1. 7.1 989 ，Fedoronchuk (KW). Syreishchikow Syreishchikow (LE). FRANC E: Pyrenees ， Pic de Gers ，22.7 .1 861 ， Ball (W); va r. subun if1 0ra 7.7 .1 958 ， Mailland (GH); Muravy ， 11. 7.1958 ， GEORGIA: C. Caucasus ， Kazbegi ， Orzveri ， Jasiewich Jasiewich (KRA); Haute-Savoie ， Col de Confins ， 24.6.1924 ，Rzewanowski (TBI); Gergeti ，10.6.1962 ， 1O ~7.1963 ， Bersier (CLUJ). Khutzishwili (TBI); Sabertse ，3 1. 7.1 974 ， Davitanidze SPAIN: C. Pyrenees ， 15.7.1924 ， Guatr (BARSS); (TBI); Svaneti ， Ridge Bakh ，M t. Khodzhalj ，25.7 .1 935 ， Espot ，Estanyets ，7.7.1934 ，Quer (BARSS); Bagyer 丸 Dolukhanov & Khradze (TBI); Shkedi ， 9.7.1978 ， 6.7.1973 ，Vigo (BARSS); Conflent ， Coll de Portos ， Tskhenshur & Chelidze (TBI); Abakura Pass ， sobre sobre Noedes ，28.7.1980 ，Romo (BARSS); E Pyrenees ， 24.7 .1 935 ， Kharadze (TBI). Du Coll de Portes au Coronat ，15.7.1898 ，Sennen RUSSIA: N. Caucasus ，Mamissonski Pass ，27.6.1897 ， (BARSS); Valle de Arau ， Val d' Arches ，6.7.1962 ， Desulavi (TB I); 20.7.1967 ，Kharadze & Gviniadze Rinos-Marhues Rinos-Marhues (V AL); Lerida ， Espot ， 21. 7.1983 ， (TBI); 21. 7.1967 ， Kharadze (TBI); 25.6.1970 ， Mateo (VAL); Vall de Crau ， lowland rola la font del Avazneli & Kharadze (TBI); Balkaria ，SuuιAuz ， Moixew ，27.7.1984 ， Sofiano (BARSS); Lerida ， Alta 16.6 .1 927 ，Bush (TBI); Dombai ，M t. Musa-Achtara ， Ribagorta ，23.7.1996 ，Aran (VAL). 4.7.1989 ，Ziman & Fedoronchuk (KW); M t. Ptysh ， ITALY: Prov di Udine ， Cavallo M t.， 15.6.1951 ， 2.7.1989 ，Ziman & Fedoronchuk (KW). Grichuitti Grichuitti 569 (NY); Mataiur M t.， 15.6.1965 ， Grichuitti Grichuitti (GH); Maritime Alps ， Piemonte ， S var. willdenowii Limonetto ，29.6 .1 982 ， Burizi & Krendl (WU). RUSSIA: W Caucasus ， Caucasia Reserve ，M t. Abago ， BULGARIA: Vitosha Mts. ， Stribmy ， 16.6.1955 ， 3.6.1979 ， Zaikonnikova (LE). Vyhodzevski Vyhodzevski (CLUJ); 13.9.1972 ， Kozhuharov 809 ARMENIA: Sarykamysh ，24.6.1910 ， Roll (KW); M t. (LE); (LE); 13.6.1976 ，Kozhuharov & Petrova (BCC). Ararat ，1912 ， Sosnowski (TBI). SLOVENIA: Kamniske Alpe ，28.6.1965 ， Martincic TURKEY: Karss ， Sub-Charkh ， 1912 ， Sosnowski 57417 (LE); Gorenjsko ， Kokrsko Sedlo ，6.6.1965 ， (TBI); Yalnizcandag ， near Ardanuc ， Davis & Hedge Martincic Martincic (CLUJ). 29672 (K); Coruh (Artvin) ，Ardanuc ，Kordevan dag at ALBANIA: Scardus ， Lynbitm ， 15.7.1890 ， Dorfler Kutulyayla ，29.6.1957 ， Davis & Hedge 30350 (K); 1187 ，2776 (W). Pontus ，Dist r. Trabzon ， Pontic Range ， S. of Rizeh ， 4.7.1960 ， Furse & Synge 897 (K). va r. podolica UKRAINE: Dolina Dniestru ， Okriany w Tlumackiem ， subsp. protracta 30.5.1921 ， Wilczinski (KRA); Gologory ，Lysa Gora ， var. protracta 30.5.1923 ， Wilczinski (KRA); 25.6.1987 ，Ziman & TADJIKISTAN: Pamir Al 吋， Gissar ，Varzob ，1968 ， Kobiv (KW); Opolje ， Kasova Gora ， 20.5.1929 ， Kamelin (TAD); Karateginski Rudge ，1948 ，Popov Kozlowski Kozlowski (KRA); Ivano-Frankivski Reg. ， Chertova (T AD); Zeravshan Ridge ，Kazan- Temir ，1952 ， Nikitin Gor ，23.6.1987 ，Ziman & Malinovski (KW). (TAD); Surkho Ridge ，1947 ，Popov (TAD); Hobu- Rabat Rabat Pass ，23.6 .1 979 ，Ziman (KW); Anzob Pass ， var. var. CrI mea 17.7.1954 ， Kaletkina (TAD); 20.6.1979 ，五 man (KW); UKRAINE: Crimea ， Nikitsk 司a Yaila ， Kr asny 31.6.1990 ，Ziman & Fedoronchuk (KW). Kameny ，6.8.1989 ，Ziman (KW); 14.8.1990 ，Ziman & KAZAKHSTAN: Tian Shan ，Alma-Ata Reserve ， M t. Ehrendorfer Ehrendorfer (KW). Talgar ，19.7.1990 ，Ziman & Fedoronchuk (KW). subsp. subsp. fasciculata va r. contracta var. var. fasciculata KAZAKHSTAN: Semipalatinski Reg. ， Ustj- GEORGIA: Central Caucasus ， Kazbegi ， Dzhuta ， Kamenogorsk ， Ush-Kutej ， M t. Sarym-Sakty ， 10.7.1936 ， Gagegiladze (TBI); 28.5.1978 ， Ziman 30.6.1909 ， Reznichenko (LE); Z 司san ，Narym Ridge ， (KW); Trialeti ， M t. Shuano ， 4.7.1967 ， Khintibidze Okpekty ，25.6.1914 ， Shaposhnikov (AA); Uch-Bulak ， (TBI); (TBI); Bakuriani ，1.6.1978 ，Ziman & Mardaleishvili 20.7.1914 ， Saposhnikov (LE); Djungar Alatau ，Alma- (KW); Svaneti ， Abakuri Pass ，24.7.1935 ，Kharadze Ata Ravine ，23.6.1928 ，Goloskokov (AA); Djildy- (TBI); (TBI); Bakuriani ，Tsykhidzhvari ，28.6.1988 ，Ziman & Karagaj ， 13.7.1934 ， Rubtsov (AA); M t. Mataj ， Fedoronchuk (KW). Ishmurdun ， 20.6.1956 ， Goloskokov (LE); Kok-Su ， October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 299

2.6.1959 ，Goloskokov (LE); Taldy-Kurgan ，Kara- Tala ， va r. uralensis 28.7.1928 ， Shipchinski (LE); Saur Ridge ，M t. Tass ， RUSSIA: Ural ，Il menski Ridge ，26.5.1927 ， Tulina 18.7.1930 ， Goncharov (LE); Tarbagatai Ridge ， (LE); Pereva りskoe Plateau ， M t. Shemur ，23.7.1972 ， Sarganak Sarganak Pass ，7.7.1948 ， Stepanova (AA); Kungei Igoshina (LE); Sereb りanski Ridge ，26.7.1925 ， Igoshina Ridge ， Karagaily Ravine ，6.7.1960 ， Roldugin (AA). (LE); M t. Kameny ，Kharyuzny ，5.07.1939 ， Igoshina (LE); (LE); Miasski Ridge ，Il meny ，2 1. 7.1 951 ， Juzepchuk subsp. subsp. crinita (LE). var. var. crinita RUSSIA: Tuva ，S 司an ， Soj-Tajga Plateau ，29.6.1986 ， va r. sachalinensis Turuta Turuta (KW). RUSSIA: Sakhalin ， Zaozernoe ， M t. Nisijama ， MONGOLIA: River Zamtea ，6.06.1894 ， Clements 31. 7 .1932 ，Take (VLA); Poronajski Dist r.， Kotikovo ， (LE); (LE); Khalkha ， Iro River ，8.7.1906 ， Novitski (LE); 12.7 .1 933 ， Khosino (VLA); Makarovski Dist r.， Ustj- Ulan-Bator-Khoto ，Urga ，5.8 .1 927 ，Ikonnikov-Galitski Pugachevo ， M t. Khassidake ， 25.7.1935 ， Sirasaka (LE); (LE); M t. Kentei ，24.7.1928 ，Ikonnikov-Galitski (LE). (VLA); Dolinski Dist r.， Dudino ，30.6.1936 ， Takakhasi CHINA: Hopei ， Hsiao wutaishan ，7.1934 ，W. T. Wang (VLA); Kr asnogorsk ，23.06.1949 ，Kravchenko (VLA); 61723 (GH); Wan-ping ，4.7.1955 ，Yang 1545 (E); E Bojnitsa Cape ， 24.7.1968 ， Egorova (VLA); M t. Kansu ，Moule ，Ischagora ，1885 ， Potanin (LE). Chekhova ， 24.7.1966 ， Probatova (VLA); M t. Vodorazdelnaia ， 17.7 .1 988 ， Vyshin (VLA); M t. va r. lineariloba Balagan ，24.7.1988 ， Vyshin (VLA); 乱1t. Lopatina ， RUSSIA: Primorje Reg. ， M t. Snezhnaja ， 15.7.1936 27.7.1988 ， Vyshin (VLA). Shishkin Shishkin (LE); Stolbovoj Mts. ，17.8.1936 ， Kolesnikov JAPAN: Hokkaido ，Rebun Is 1.， Wen-nai ，2.7.1953 ， (VLA); 11.8.1994 ，Ziman (KW); Ternej Ridge ， M t. Furuse (TNS); Hokkaido ， Prov. Ishikari ，. M t. Glukhomanka ，5.7.1964 ， Stepanova (VLA); 15.8.1968 ， Taisetsusan ，19.7.1941 ，Okuyama (TNS); Hokkaido ， Kolesnikov Kolesnikov (VLA); 6.8.1984 ， Kharkevich (VLA); Prov. Hidaka ， M t. Poroshiridake ， 1600 m ，25. 7 .1971 ， Pozharsk ，Bikin ，25.6.1981 ， Vyshin (VLA); Ternei ， Koyama & a1. 3781 (TNS); Hokkaido ， Prov. Hidaka ， Opasnaya River ， 5.8 .1 982 ， Oleinikov (VLA); M t. Chirorodake ， 1700 m ， 31. 7.1 986 ，Kadota & Dalnegorsk ， Tulapinski Pass ，9.9.1956 ， Voroshilov Midorikawa 13311 (TNS); Tohoku ， Akita Pref. ，M t. (VLA); 8.6.1961 ， Kurentsova (VLA); Kamenka ， Chokaisan ，6.8.1929 ，Okuyama 2246(a) (TNS)/ 18.6.1974 ， Rechan (VLA); Rudnaya Pristany ， 19.6.1975 ， Volynets (VLA); Zazerkalnoe Lake ， subsp. villosissima 22.5.1981 ， Shemetova (VLA); Kr asnorechensk ， RUSSIA: Kuri1es ，Rubeteu ，28.0 1. 1884 ，Miyabe (LE); M t. Medvezhya ， 17.6.1983 ， Gularjants (VLA); 5.8. ， M t. Vetrenaya ， 11. 8.1978 ， Barkalov 1994 ，Ziman & Gularjants (KW); Ussuri ， Katen ， (VLA); Ozernyi Cape ，28.6.1979 ， Barkalov (VLA); 22.7 .1 958 ， Kurentsova (VLA); Chuguev ， Shelikhova Bay ，5.8.1979 ， Barkalov (VLA); Zelenaya ， M t. Oblachn 司a，15.8.1961 ， Gorovoi (VLA); 15.7.1969 ， 11. 8.1978 ， Barkalov (VLA); 7.7.1979 ， Barkalov Voroshilov Voroshilov (VLA); 15.6 .1 983 ， Vyshin (VLA); (VLA); M t. Dozhdevaya ，5.8.1979 ，Vyshin (VLA); 24.9.1983 ， Vyshin (VLA); Seacost of Japanese Sea ， ，Kataoka ，30.7.1948 ，Vorobyev (VLA); 1972 ， Preobrazhenie ，1. 8.1 994 ，五 man (KW). Khabarovski Serdyuk (VLA); ， Parusnaya Bay ，13.8.1982 ， Reg. ， Anyui ， Tardoki- Yana ， 25.8.1956 ， Rozenberg Barkalov (VLA); Jodnyi Cape ， 14.8.1989 ，Vyshin (VLA); 20.7.1989 ， Kharkewich (VLA); Chu-Anui (VLA); Bogatyrj Ridge ，Stokap ， ，24.8.1988 ， Pass ，10.7.1983 ，Kharkewich (VLA); Daur ，10.8.1983 ， Barkalov (VLA); M t. Kamennik ，17.8.1979 ， Barkalov Vyshin Vyshin (VLA); Tumnin ， Bolshoi Jan Ridge ， (VLA); ， 11. 7.1 959 ， Pobedimova (VLA); M t. Komandnaya ， 21. 10.1984 ， Vyshin (VLA); ， Boljshoe Lake ，9.7.1978 ， Barkalov (VLA); Magadan Reg. ，Oksy Ravine ，9.9 .1 976 ，Khokhryakov Moneron ，30.5.1974 ， Stepanova (VLA); M t. Daynan ， (VLA); Okhotsk ， Ajano-Mai ， Nelkan ， 7.7 .1 978 ， 24.7.1981 ， Kimoto (VLA); ，Nemo Bay ， Kharl 王evich VLA); 3.6 .1 979 ， Kharkewich (VLA); 11. 7.1981 ， Barkalov (VLA). Chukotka ， Vankarem ， 7.7.1989 ， (LE); U. S. A.: Aleutians ， Unalaska ， Dutch Harbor ， Buryatiya. Buryatiya. Sajan ，Barum-Kharaduz ，25.8.1994 ， Turuta 17.6.1899 ， Coville & Kearney 1722 (GH); 8.7.1899 ， (KW). Coville & Kearney 1782 (GH); 25.5.1932 ，Eyerdam MONGOLIA: Hangai ， Uzin-Tabi ， 17.7.1926 ， 787 (K); 31. 5.1 932 ，Eyerdam 809 (K); KagamiI ， Prokhanov Prokhanov (LE); Egin-gola Ravine ， Urtuin-daba ， 2.7 .1 949 ， Miller 1045 (US); Attu ，ODonnel Valley ， Tsagan-ur ，22.7.1972 ，Grubov (LE). 23.6.1944 ， Morenhouse (GH); Massacre Bay ， 28.5.1945 ，Van Schaack 336 (K). 300 300 植物研究雑誌第80 巻第5号 平成17 年10 月 subsp. subsp. alaskana Pref. ，M t. Shiroumadake ，3.8.1938 ， Hayashi (TNS); u. u. S. A.: Alaska ，Kodiak ，25.7.1904 ，Pipir 4470 (K); Prov. Sagura ， Shizuoka Pref. ， M t. Arakawadake ， Beringh Beringh Straigt ，Diomede Island ，14.8 .1 926 ，Porsild & 23.8.1950 ， Furuse (GH); M t. Arakawadake ，29.7 .1 951 ， Porsild Porsild 1692 (GH); Kantishna Distr. ， 21. 7.1939 ， Muramatsu 3363 (TNS); M t. Arakawadake ，1. 7.1958 ， Nelson Nelson (NY); Wild Hoa ，23.7.1939 ， Nelson (GH); Mts. Furuse (GH); Prov. Shinano ， M t. Komagatake ， Roberts ，Juneau ，15.9.1939 ， Nelson & Nelson 4437 Nogaike ， 2800 m ， 2.8.1959 ， Tamura 7627 (E); (GH); (GH); Nutzotin Mts. ， near mi 1345 ，12.8.1944 ，Raup Yamanashi Pref. ， Prov. Kai ， between the summits of 12797 12797 (GH); Ancorage ，Chugach Mts. ，14.7.1947 ， Neil M t. Kitadake and Kitadake-goya ， Ashiyasu-mura ， 21207 21207 (NY); Mts. NE of Blocks Cabin ， head of Big 20.8.1965 ，Tamura & Hotta 3627 (E); Yamanashi River ，10.7 .1 950 ， Drury 4065 (GH); Falls Creek ， Kenai Pre f.， Mts. Yatsugatake ， M t. Akadake ， 15.8 .1 958 ， Lake ， 21. 7.1951 ， Calder 6149 (US); Boundary Okawa (TNS). Roadhouse ， near Yukon territory ，9.8.1951 ，Scamman 6290 (GH); Point Hope ， 160 mi NW of Kotzebue ， References 22.8.1951 ，Scamman 6396 (GH); Cathedrae Bluffs Adams M. F. 1885. Decades Quinque Novarum Creek ， 27.7.1957 ， Spetzman 857 (US); Moresey Specierum Plantarum Caucaso et Iberiae Com. Island ，M t. de la Touche ，Fairfax Inlet ， Tasu Sound ， Mussin-Puschkin observavit et definitiobus atque 16.8.1957 ， Calder & Taylor (US); Alaska Range ， descriptionibus illustrativ M.F.Adams. Beitr. Maclaren Maclaren glacier ， 1100 m ，13.8.1958 ，Shacklette (US); Natu r. Weber et Mih r. Vo l.l. Brooks Brooks Range ， John River ， Anaktwik Pass ，700 m ， Baumberger H. 1970. Chromosomenzahlbestimmun- 22.7.1959 ， Campbell (US); 85 mi of Fairbanks ， ges und Karyotypanalysen bei den Gattungen 19.6.1964 ， Harms 2738 (GH); Chugach St. Peak ， Anemone ， Hepatica und Pulsatilla. Be r. Schweiz. Eklutka Eklutka Valley ，Bold Ridge ，14.08 .1 984 (U.S.). Bo t. Ges. 80: 17-96. CANADA: British Columbia ，Yukon territory ，4 mi Britton N. L. 189 1. The American species ofthe genus SE of Ptarmigan Heart ，13.7.1948 ，Raup et al. 13595 Anemone L. and the genera which have been re- (GH); (GH); Moresby Island ，Lake Takakia ， 10 mile S. of ferred to it. Ann. New Y ork Acad. Sci. 6: 215- Moresby Logging Camp ，25.7.1964 ， Calder & Taylor 232. 36285 36285 (K); Lower Lapie ，Range Crossing ，26.6.1994 ， Bruhl P. 1896. Descriptions of new and rare Indian Porsild Porsild & Breitung (N.Y.); Mackenzie Dist r.， plants. Ann. Roy. Bo t. Gard. Calc. 5: 76-8 1. 25.7.1949 ，Cody & McCanse 3137 (GH). Calder J. A. and Taylor R. L. 1968. Anemone L. Fl ora of of the Queen Charlotte Islangs. Monogr. N 4. Part subsp. subsp. zephyra 1: 339-34 1. Systematics of the Vascular Plants. U. U. S. A.: Colorado ，Rocky Mts. ，W of Soryo Peak ， Research Branch. Canada Department of 12500 12500 ft.， 1868 ，Vasey (E); Long Peak ，4200 m ，1868 ， Agricultture. Vasey (NY); Larimer Co. ，Cameron Pass ，3400 m ， Chaudhary R. P. 1988. A Synopsis of the genus 8.1 894 ， Osterhout (E); 2 1. 7.1951 ， 6149 (GH); Anemone (Ranunculaceae) in Nepal and the adja- 18.8.1997 ，Ziman & Dutton (KW); Beaver Greek cent Regions of the Himalaya. Bo t. 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S. S. N. ジーマンヘ F. エーレンドルファー"， C.

S. キーナーc，門田裕ーヘ O. N. ツアレンコ"， E. ブラークa. B. E. ダットンペイチリンソウ属ハ 日本産植物の学名はハクサンイチゲが A. クサンイチゲ節[広義] (キンポウゲ科)の分類 narcissiflora L. subsp. nipponica (Tamura) Kadota ， 学的再検討(1) エゾノハクサンイチゲがA. narciss 抑ora subsp. イチリンソウ属ハクサンイチゲ節[広義] (キ crinita (J uz.) Kitag. va r. sachalinensis Miyabe & ンポウゲ科)の分類学的再検討を行った.その結 Miyake となる. 果， 3 列と 11 種が認められ，それらの検索表を付 ("ウクライナ・ M.G. コロドゥニィ植物学研究所，

した.本論文ではその第一部として，ハクサンイ b ウィーン大学植物学研究所，

チゲ複合体の解析結果を報告した.ハクサンイチ C ペンシルバニア州立大学，

ゲ複合体を Anemone narcissiflora L. 1 種にまとめ， d 国立科学博物館植物研究部，

8 亜種12 変種を認めた. A. narcissiflora は北半球 e西オレゴン州立大学) に広く分布するが，ヒマラヤ地域には分布しない.