植物研究雑誌 Originals J. Jpn. Bot. 83: 127–155 (2008)

Modern View on the of the L. Sensu Stricto ()

Svetlana N. ZIMANa, Elena V. BULAKHa, Yuichi KADOTAb and Carl S. KEENERc

aN.G. Kholodny Institute of Botany, National Academy of Sciences, 2, Tereshchenkivska street, Kiev, 01601 UKRAINE; bDepartment of Botany, National Museum of Nature and Science, 4–1–1, Amakubo, Tsukuba, 305-0005 JAPAN; c208, Mueller Laboratory, Pennsylvania State University, University Park, Pennsylvania, 16802 U.S.A. (Received on 17 November, 2007)

As a result of the re-examination of the Anemone genus taxonomy, we accept this genus as including 15 subgenera, 23 sections, 4 subsections, 23 series and 118 . Within Anemone,6subgenera, 2 sections and 16 series were proposed originally by us, and we re-examined the taxonomic state of ca. 50 species and intraspecific taxa. Besides, we elaborated the annotated conspectus of the genus Anemone and the key for determina- tion of its species, series, subsections, sections and subgenera on the basis of the analysis of about 70 characters of the achenes, , , above-ground and underground shoots and roots. (Continued from J. Jpn. Bot. 81: 193–224, 2006)

Key words: Anemone L., comparative morphological analysis, taxonomic revision.

Literature survey first systematic treatment of this genus. He The genus Anemone L. is one of the larg- regarded as independent genus and est genera within the family Ranunculaceae. proposed to place Anemone and Hepatica The status of this genus, the number of its into the tribe Anemoneae DC., together with species, its division into sections and other Knowltonia Salisb., Thalictrum L., Hydrastis intergeneric taxa, as well as their taxonomic L. and Hamadryas Comm. & H. Lév., etc. In and evolutionary relationships are long- his Anemone system, there were six groups, debated. and only four of them included species of Beginning from Linnaeus (1753), who Anemone sensu proprio (Anemonanthea DC., described the genus Anemone, there have Anemonospermos DC., Omalocarpus DC., been different opinions on its delimitation and Pulsatilloides DC.), while two groups and size. One year after Linnaeus, Miller (Pulsatilla DC. and Preonanthus DC.) em- (1754) separated from Anemone the genera braced species currently placed in Pulsatilla. Pulsatilla Mill., Hepatica Mill. and Gray (1821) elevated the section Mill., and shortly afterwards Anemonanthea to generic level (Anemo- Adanson (1763) described the allied genus nanthea S. F. Gray), although exactly this Oriba Adans. group of species was already described as Jussieu (1789) recognized the genus the genus Anemonoides (Miller 1754). Anemone in the Linnaean sense, while Rafinesque (1825) recognized the North Candolle (1817, 1824) was the author of the American species (A. caroliniana and others)

—127— 128 植物研究雑誌 第83巻第3号平成20年6月 as the genus Hepatica Raf., while Gay 17 series (viz., Himalayicae Ulbr., Begonii- (1845) described the genus Barneoudia folia Ulbr., etc.). As a whole, Ulbrich revised C. Gay from as close to in his monograph 76 species of Anemone Anemone. Meanwhile, Schur (1866) re- distributed worldwide. garded A. narcissiflora and allied species Ulbrich’s Anemone system was generally as the genus Homalocarpus Schur, accepted for many years, and its profound Schlechtendal (1856) segregated the South examination was absent, except for the par- American species allied to Pulsatilla as the ticular proposals, mainly within the so called monotypic genus Oreithales Schlecht., and “local floras”. Thus Juzepchuk (1937) gave some forty years later the same taxon was several additions for species occurring erroneously described as the genus Capethia within the borders of the former Soviet Britt. (Britton 1892). Union and he regarded all Anemone sec- Sprengel (1825) accepted generic state of tions sensu Ulbrich as subgenera (viz., Anemone and Pulsatilla only, Spach (1839) Anemonanthea DC., Homalocarpus DC., who described subtribe Anemoninae recog- Pulsatilloides DC., Anemonidium Spach, nized both Pulsatilla and Hepatica as the Eriocephalus Hook. f. & Thomson, and genera, and Pritzel (1841) did not recognize Rivularidium Jancz.), besides, he elevated Pulsatilla. Prantl (1887) regarded it as a most of Ulbrich’s subsections and series to genus, but included Hepatica in Anemone section level. Juzepchuk also described sev- and did not recognize Knowltonia and eral additional series (viz., Nemorosae Juz., Barneoudia. Finally, Janczewski (1892) Flaccidae Juz., Baicalenses Juz., Sylvestres accepted all the forementioned taxa (except Juz. and Rupicolae Juz.) but did not support Knowltonia)asgroups within the delimita- them with Latin diagnoses, like other taxo- tion of Anemone. nomic novelties. After Candolle, several new sections Nakai (1949) separated some species of within Anemone were described: Diplo- the section Anemonospermos as the genus calymnata Spreng. (Sprengel 1825), Sylvia Eriocapitella Nakai, and Wang (1974) ad- Gaudin (Gaudin 1828), Oriba (Adans.) mitted the generic state of the monotypic se- Spach, Anemonidium Spach and Phaeandra ries Anemoclema Franch. (Franchet 1886, Spach (Spach 1839), Eriocephalus Hook. f. 1888, 1890) as the genus Anemoclema & Thomson (Hooker and Thomson 1855) (Franch.) W. T. Wang. and some others. Sixty years after Ulbrich, Tamura (1967) The most detailed system of the genus re-shaped the Anemone system while the Anemone s. l. was worked out by Ulbrich revising the system of Ranunculaceae.Asa (1905, 1906). The author recognized whole, this author accepted all six sections of genera Pulsatilla, Knowltonia, Barneoudia Anemone sensu Ulbrich, however, he added and Capethia, but he regarded Hepatica as three Anemone sections, and only one of the subgenus of Anemone. Within subgenus them (the monotypic section Keiskea Euanemone, Ulbrich recognized six sections Tamura) was really new because sections already proposed, viz., Anemonanthea DC., Begoniifolia Tamura and Anemoclema Omalocarpus DC., Pulsatilloides DC., Tamura were the former series Begoniifolia Anemonidium Spach, Eriocephalus Hook. f. Ulbr. and Anemoclema Franch. Besides, & Thomson, and Rivularidium Jancz. This Tamura re-examined section Pulsatilloides, author described four subsections (Tuberosa and he ignored its subsections Brevistylae Ulbr., Stolonifera Ulbr., Brevistylae Ulbr. and Longistylae sensu Ulbrich but moved and Longistylae Ulbr.), and he also described the series Himalayicae sensu Ulbrich from June 2008 Journal of Japanese Botany Vol. 83 No. 3 129

Pulsatilloides to the section Omalocarpus as (Ulbr.) Ziman, Parviflora (Ulbr.) Ziman, the subsection Himalayicae. This author de- Flaccida (Ulbr.) Ziman, and Himalayicae scribed three series (Nemorosae Tamura, (Ulbr.) Ziman. She also separated some Flaccidae Tamura and Rupicola Tamura), species of the section Rivularidium into the although all of them were described in 1937 section Rivularis (Ulbr.) Ziman, and some by Juzepchuk (1937). Despite the fact that species of the section Anemospermos into the the Juzepchuk series were invalid, Tamura’s section Vitifolia Ziman. The first five sec- taxa were not supported by Latin diagnoses tions were supported by Latin diagnoses, and were invalid too. therefore, they may be considered as the Beside the recognition of the genera valid ones. Pulsatilla, Hepatica, Knowltonia and The novations of Starodubtsev (1989, Barneoudia, Tamura accepted also three 1991) represented a new stage of the further “narrow” or segregate genera Capethia, complication of the genus Anemone and the Miyakea and Eriocapitella. Like most of his system of the subtribe as a whole, and in fact predecessors, Tamura recognized the tribe he revised the huge forementioned taxo- Anemoneae and subtribe Anemoninae. nomic group. Beside admitting the genera After Tamura, further splitting of the Pulsatilla, Hepatica and Barneoudia, genus Anemone continued. So Löve and Starodubtsev followed Holub (1973) in Löve (1975) separated from it the monotypic corfirming the generic independence of series Richardsonia Ulbr. as the genus Anemonoides, Anemonastrum and Anemoni- Jurtsevia, Holub (1973) recalled the genus dium.Hemoved several species from section Anemonoides Mill. and described the genera Rivularidium to Anemonidium, and he ele- Anemonastrum Holub (former sect. vated the rank of three sections to the generic Omalocarpus DC. or genus Homalocarpus level [viz., Pulsatilloides (DC.) Starod., Schur) and Anemonidium (Spach) Holub Arsenjevia Starod. (former section Flaccida (former sect. Anemonidium), and Wang Juz.) and Tamuria Starod. (former sect. (1980) separated the narrow genus Keiskea Tamura)]. Within the genus Metanemone W. T. Wang. While working Anemone sensu strictissimo, Starodubtsev out his taxonomic scheme of the genus recognized three sections (Anemone, Anemone for the flora of China, this author Eriocephalus and Diplocalymnata) and ten mainly followed Ulbrich (1906) and Tamura subsections. (1967), but he included both Eriocapitella The results of the Starodubtsev study are and Anemoclema in Anemone. very valuable but they were based on the While discussing the phylogeny of the peculiarities of the fruits only, and his new family, Ziman (1985) proposed several al- concept of the tribe Anemoninae was based terations for the system of Anemone and mostly on material from the Russian Far Anemoninae, viz., she recognized the genera East. Moreover, our re-examination of his Pulsatilla, Hepatica, Knowltonia and herbarium materials in LE and VLA has Barneoudia but did not except the genera shown that his data were mostly inade- Eriocapitella, Anemoclema, Anemonidium, quately documented. In our opinion, some Anemonastrum, and Anemonoides (recogniz- taxonomic proposals of Starodubtsev within ing them as parts of the genus Anemone), the genus Anemone s. l. and the subtribe and Capethia, Oreithales and Miyakea (parts Anemoninae are debatable, especially his of the genus Pulsatilla). Beside, Ziman narrow generic concept, but all of them proposed to elevate the rank of four of provide a serious base for the further exami- Ulbrich’s series to sections Richardsonia nation of this taxon in detail. 130 植物研究雑誌 第83巻第3号平成20年6月

About at the same time as Starodubtsev, Anemone have been based on distinctions of Tamura (1991, 1995) proposed some im- fruits and perianth, and most authors paid the provements and alterations to the system great attention to the shape, size and pubes- of the family Ranunculaceae, subtribe cence of these organs. Some authors (e.g., Anemoninae and genus Anemone. This Tamura 1995) have also taken into consid- author confirmed the recognition of genera eration additional characters of leaves and Pulsatilla, Hepatica, Knowltonia, shoots, especially while distinguishing sec- Barneoudia, Metanemone and Oreithales tions and series. (instead of Capethia in 1967), and, like The anatomical studies of fruits of earlier, he recognized all six subgenera of Anemone taxa began with Janczewski Juzepchuk (1937). Beside that, Tamura (1890, 1892) and Smith (1926), and accord- elevated the rank of section Anemoclema ing to Starodubtsev (1991) and Tamura (Franch.) Tamura to subgenus Anemoclema (1963, 1995), the structure peculiarities of (Franch.) Tamura, series Hepaticifolia Ulbr. Anemone achenes are very important for to subgenus Hepaticifolia (Ulbr.) Tamura, their taxonomy. However, the literature data and series Rigida Ulbr. to subgenus Rigida on Anemone achenes (Ponomarenko and (Ulbr.) Tamura. He confirmed his own Berestetskaya 1981, Chaudhari and section Keiskea (within the subgenus Trifonova 1988, etc.) encompasses approxi- Anemonanthea), and he elevated subsections mately 60 species. Therefore, we tried to Tuberosae Ulbr., and Stoloniferae Ulbr., and include in our study the fruit morphology series Richardsoniae Ulbr., Alchimillefolia (and partly anatomy) of as many Anemone Ulbr., and Kilimandscharica Ulbr. to the species as possible. We examined ca. 110 level of sections. Anemone species in detail (including data on The data on serology (Jansen 1968, cross section and microsculpture of a sur- Chupov 1975, etc.), biochemistry (Hantula et face). Initially we published our results on al. 1989, Hoot 1995, Hoot et al. 1994, etc.), study of achenes (and other organs) of geographical patterns (Ziman 1985, Ziman A. narcissiflora and A. biflora aggregates and Keener 1989, etc.) were also important (Ziman et al. 1997, 1998) but afterwards we for clarification of some problems of published results for subgenera Anemonan- Anemone s. l. and they have added a wealth thea, Stoloniferae and Omalocarpus (Ziman of new information concerning the overall et al. 2004a, 2004b, 2004c, 2005, 2006a, relationships within all taxa of the subtribe 2006b), and section Himalayicae (Ziman Anemoninae. et al. 2007). We realized the value of com- However, many questions of the taxon- parative study of Anemone carpels and omy of the genus Anemone s. l. are still achenes because most of their morphological open. Thus, one very important task is the re- characters are in common, and their vision of the generally accepted characters, distictions help us to understand possible and more accurate definition of their diag- evolutionary trends. nostic and evolutionary significance, but also Most authors used five to eight achene the selection and examination of approaches characters (mainly of external morphology) and diagnostic characters of Anemone taxa. to distinguish Anemone species and their groups. According to Hoot et al. (1994), it is Morphologic characters essential expedient to use around 10 achene charac- for taxonomy of Anemone ters, and they proposed to add to the known Beginning from Candolle (1817), the characters shape of fruitlets and number of majority of the systems of the genus achenes in them. Considering our recent June 2008 Journal of Japanese Botany Vol. 83 No. 3 131 studies, we believe it possible to extend the Pulsatilloides)orshort (ca. 1 mm long, viz., number of the fruit characters of Anemone in Anemonanthea, etc.) or sometimes hardly species to 25. visible (viz., in Eriocapitella and It is well known that Anemone fruits are Stolonifera). Carpels and achene styles are apocarpic one-seeded achenes forming heads mainly straight or hooked (basally or (fruitlets). We paid the attention to the com- apically) but sometimes uncinate, and they mon carpel and achene shape (viz., ovoid, are narrow-conic (viz., in Rivularidium), globose or ellipsoid) but also the peculiari- sharply narrowed (viz., in Omalocarpus) ties of their basal (sessile, stalked or at least or sometimes funnel-like (viz., in narrowed) and apical (rounded or narrowed) Eriocapitella). parts. We concluded that Starodubtsev has Carpel and achene stigmas are mainly exaggerated the taxonomic significance of linear but sometimes capitate or subcapitate the stalked Anemone achenes when he sepa- (viz., in Eriocapitella, Kilimandscharica and rated several species (viz., A. keiskeana and Stolonifera)ordilated, and the latter corre- A. deltoide) into genus Tamuria because late with very short styles (sessile or Eriocapitella and Kilimandscharica species subsessile stigmas). also have the distinctly stalked carpels and Carpels and achenes are covered with achenes, moreover, several other Anemone hairs but sometimes they are glabrous or species (viz., A. antucensis, etc.) are charac- nearly so. The hairs are long (2–2.5 times terized by carpels and achenes basally nar- longer than an achene diameter) or short rowed into stalks. (shorter than an achene diameter). Besides, Sometimes Anemone carpels and achenes carpel and achene villosity may be dense are laterally compressed and ribbed or even (lanate) or sparse. The carpel and achene winged, and Anemone species having the hairs are typically monomorphic but some- winged reproductive organs are separated times they are dimorphic, especially in their into the Omalocarpus group. However, spe- basal and apical parts. In most taxa hairs cies characterized by compressed carpels and cover achene bodies only but sometimes achenes (with more or less distinct lateral styles too (viz., in A. baissunensis, A. ribs) are also found within Anemone s. str., capensis, etc.). Rivularidium, Begoniifolia, Anemonidium In cross section Anemone achenes are and some other groups. suborbicular or elliptic to elongate, with nar- In Anemone species the ovoid shape of row excrescences in which the vascular bun- carpels and achenes predominates but there dles are settled. Pericarp is well-developed are also globose, cylindroid and spindle-like and differentiated into exocarp, mesocarp carpels and achenes. The length of carpels and endocarp. A cross section of seeds re- and achenes of Anemone species is fre- sembles that of fruits; the seed coat consists quently 2–10 mm, but there are taxa with of several layers, the latters usually being large (4–8 mm long, viz., in Omalocarpus very compressed. Besides, Anemone taxa and Rivularidium) and small (1–2 mm long, differ by the peculiarities of the micro- viz., in Anemone, Eriocephalus and others) sculpture of achene surface (smooth, carpels and achenes. plicated, tuberculated, waved, folded, etc.). The length and shape of carpel and achene At present about ten morphologi- styles are various, and they are typically per- cal characters are used for distinguishing sistent at maturity. Styles differ in Anemone Anemone species, and they pertain mainly to species by their length being sometimes long the tepal number, color and size, and some- (5–10 mm long and tail-like, viz., in times to the and receptacle shape 132 植物研究雑誌 第83巻第3号平成20年6月 but the recent detailed study of the micro- some of them have more than 10 morphology of perianth (Tamura 1962, anastomoses (viz., A. coronaria, A. Slavikova 1968, 1976, Gulanjan 1974, etc.) virginiana, A. hepaticifolia, etc.) or their has added important data to the understand- tepals lack anastomosing veins (viz., in sec- ing of the taxonomy and phylogeny of tions Himalayicae, Parviflora, Caroliniana, Anemone. etc.). As a result of the critical examination of However, the tepal size and colour and flowers within the genus Anemone,wedis- sometimes shape of many Anemone taxa tinguished the most essential ones, and we frequently vary, therefore, they could be published these results (Ziman and Bulakh regarded as characters of a specific level not 2002a, 2002b, 2003, 2004, Ziman et al. of a higher taxonomic value. 2004a, 2004b, 2004c, 2005, 2006a, 2006b, Stamens are inconstant in number (40 to 2007, Ehrendorfer et al. 2007). 100), and the shape of their filaments (linear For distinguishing Anemone taxa (sec- or dilated) is already used to distinguish tions, subsections and series), beside the some Anemone species. After the examina- forementioned characters of carpels (ovary, tion of the shape and size of all parts of sta- style and stigma shape and villosity), we mens including their filaments, connectives used also size, number and venation, but also and anthers, we came to the conclusion to peculiarities of filaments, connectives and use the former two characters in the anthers. The diagnostic characters of species Anemone taxonomy. are mainly tepal colour, size and hairiness, Within the subtribe Anemoninae, and tepal, but also ovary and style shape pe- staminodes were regarded as a diagnostic culiarities, and they were used in our key for character for several Pulsatilla groups, and distinguishing species (Ziman et al. 2002). had not been noted in flowers of Anemone The number of tepals is 5 to 8, rarely 4 or taxa. However, as a result of the precise 9, however, there are 10 or even more tepals examination of Anemone flowers, we found in several groups (viz., Carolinianae, staminodes of various shapes in several Pulsatilloides, Alchimillifolia, Altaicae, Anemone species (viz., in the sections Tuberosa, Hepaticifolia, Tuberosae). In Himalayicae and Rosulantes). these groups the tepals are usually sublinear, The data on the pollen grains of Anemone they occur in two whorls and differ in shape, species are sufficient. According to existing size, colour and venation (dimorphic literature on pollen structure (Kumazawa perianth). However, most Anemone species 1936, Mittra and Sharma 1963, Si and Chjan have ovate to obovate tepals with obtuse or [Xi and Zhan] 1964, Huynh 1970a, 1970b, acute bases and tips. Savitski 1982, etc.), their pollen grains are We obtained data on the pecularities of radial-symmetric, isopolar, globose, some- tepal venation (including anastomosing times ellipsoid or ovoid, mainly of a middle veins), initially in the Anemone narcissiflora size, with equatorial diameter 19–45 mm. (Ziman et al. 1997) and A. biflora aggregates There are colpi or pores on the surface of (Ziman et al. 1998) tepals but afterwards we pollen grains. Colpi are meridional or not studied the flower peculiarities in all meridional, narrow or wide, long or short, Anemone species and our results confirmed with pointed, rounded or blunt tips. Their the diagnostic significance of the foremen- number is 3 to many (up to 10), and germi- tioned characters. nated pores are mostly 18 to 26, and they are As happened, most species are character- irregularly arranged. Exine is 1.0–3.0 (–4.0) ized by 3–7 anastomosing veins on tepals but µm thick, sexine is thicker than nexine, or June 2008 Journal of Japanese Botany Vol. 83 No. 3 133 equal to it; and trifine is thin. Surface of going characters based on the literature data pollen grains differs in peculiarities of their only, and in other manuscripts on Anemone sculpture, tuberculate, granulate, waved, taxa the data on leaves, shoots and roots are spined, plicated, etc. The foregoing authors fragmented. recognized five types of pollen grains We began to study the vegetative organs withing the genus Anemone: tricolpate (life forms) of Anemone species about 30 (majority of species), polycolpate (three years ago (Ziman 1978, Ziman and Savitski groups), pantocolpate or rugose (nine 1980, Ziman 1985, 1986, etc.). Initially we groups), pantoporate (five groups), and studied about 25 Anemone species in detail spiroaperturate (two groups). in the natural populations, and also examined Many authors regard the foregoing pollen the life forms of nearly all species of this grain characters as very important for genus (ca. 150 species) from herbarium ma- taxonomy and phylogeny of many taxa terial. As a result, we came to the conclusion within the vascular including the that the diagnostic value and evolutionary genus Anemone. But as long ago as in 1936, significance of many characters of leaves, Kumazawa noted that in Anemone the vari- shoots and roots of Anemone taxa is essential ous types of pollen grains did not seem to be indeed. Therefore, we enlarged the number coincident with the differentiation of tradi- of the discussed characters of the vegetative tional diagnostic characters generally sup- organs of Anemone s. str. to about 40. ported by taxonomists. According to the All Anemone plants are perennial herbs, modern data, there are several groups of erect and mainly scapose. Anemone capensis Anemone species (viz., Stoloniferae, from the Pulsatilloides group is the only Multifidae, Rivulares, etc.) within which exception because it is a subshrub with three main types of pollen grains (tricolpate, lignified stem bases. rugose and pantoporate) are present includ- The plants with semirosetteous shoots ing intermediate types. Besides, a large vari- predominate but within some groups (viz., ability of pollen grains is observed within Anemonanthea, Richardsonia, Anemoni- some groups of species or even within the dium, Stolonifera and Keiskea) the adult same species or population. plants have solitary basal leaves with distinct The majority of taxonomists used several blades, and other leaves are scale-like. morphological characters of leaves and Most Anemone species are characterized partly of shoots to distinguish Anemone by sympodial renewal of shoots, however, species and groups of species. They used the main shoot is monopodial in plants of mainly size and shape of basal and some Eriocephalus and all Omalocarpus, involucral leaves, number and size of stems, Himalayicae and Parviflorae groups, and some peculiarities of underground shoots, their flowering scapes develop from axils of and pubescence of the vegetative organs. basal leaves every year. Moreover, Starodubtsev (1991) proposed to The adult plants of all Eriocapitella, take into consideration several additional Hymalayicae, Anemoclema, Pulsatilloides, characters of seedlings (number of cotyle- and also of several species of Rupicolae and dons, their size and position, and size of Rivularis groups have tap-roots; all other hypocotyl). Besides, this author paid atten- groups are characterized by adventitious tion to the number of involucral leaves and roots only. The underground shoots of the to structure of shoots (rosetteous or tap-rooted plants are represented by branches unrosetteous, monopodial or sympodial). of the caudex, and in plants having adventi- Unfortunately, Starodubtsev revised all fore- tious roots the underground shoots are 134 植物研究雑誌 第83巻第3号平成20年6月 , stolons or tubers but sometimes The involucral leaves are petiolate or ses- root-runners. sile and they are mainly similar to basal ones Anemone species differ in number of and smaller but sometimes larger (viz., in stems and their size, as well as peculiarities most Anemonanthea species). However, in of inflorescences. Racemose inflorescences some taxa the involucral leaves differ from predominate, but Anemonidium species are the basal ones by their shape and size. The characterized by a dichotomous inflores- number of involucral leaves is often regarded cence in which the main axis is extremely as diagnostic but our detailed study (Ziman reduced. In the Rivularis group the et al. 1997) has demonstrated the variability dichasium is observed in which the axillary of this character within section Omalo- branches overtop the main axis. The carpus. umbelliferous cymes are characteristic for Some peculiarities of ontogeny in most Omalocarpus species. Finally, some Anemone could be used as additional charac- Anemone groups (viz., Himalayicae, ters for distinguishing species and higher Rupicolae, Richardsonia, Parviflora, Crassi- taxonomic groups (Ziman 1985, Ziman et al. folia, etc.) have solitary flowers, due to re- 1997, 1998, etc.). Epigeal germination duction of various types of inflorescences. within Anemone species predominates but All the forementioned characters of shoots sometimes seedlings with underground small and roots of Anemone taxa are constant and cotyledons occur (viz., in the Anemonanthea may be regarded as having high taxonomic and Stolonifera groups). The majority of value. Anemone seedlings is characterized by the The basal leaves of Anemone species are short tube (result of the partial fusion of peti- usually long-petiolate with a broad blade. oles of the cotyledonary leaves), but within However, in a few species (viz., A. trulli- some groups (viz., Omalocarpus, Anemone folia, etc.) blades are narrow, and peti- s. str., etc.) these petioles are fused com- oles are indistinct. The leaves with cordate pletely. Long hypocotyl is observed in most base are predominate. The basal leaf blades Anemone groups, and in some of them (viz., are deeply palmately or ternately sected or Flaccidae, Rivularis, Omalocarpus, etc.) the parted, and sometimes ternately compound, short hypocotyl is combined with the long but several groups have pinnate leaf blades cotyledonary petioles. The position of the (viz., Anemoclema). Biternate or triternate, cotyledonary petioles during germination as well as bipinnate or tripinnate leaves, are (erect or bent) is also a diagnostic character. known in Anemone species. The palmately The size and shape of cotyledones, and juve- lobed basal leaf blades occur rather seldomly nile, immature and virginile leaves are also (viz., in A. hepaticifolia, A. palmata); the the rather important additional diagnostic leaves with entire blades are also rare (viz., characters. in several Himalayicae and Alchimillifolia The genus Anemone is the most diversed species). in petiolar anatomy within the family The basal leaf blade petioles are usually Ranunculaceae (Tamura 1962, 1964, narrow and sometimes basally vaginate. Trifonova and Zubkova 1990), and it is However, we found in several sections (viz., unique within the family in having all four Anemone s. str., Himalayicae, Hepaticifolia, types of the vascular bundle arrangement etc.) the sharply dilated (auriculate) basal (dorsiventral, dorsiventral scattered, radial parts of basal leaf petioles. These characters and radial scattered). Moreover, the are always constant, therefore, have a diag- dorsiventral scattered type was found in nostic value. Anemone only (A. rivularis). Our study of June 2008 Journal of Japanese Botany Vol. 83 No. 3 135 more than 30 populations of the Anemone et al. 1994, Ehrendorfer 1995, Ehrendorfer narcissiflora-aggregate (Ziman et al. 1997) and Samuel 2000, 2001, Schuettpelz and confirmed the taxonomic significance of a Hoot 2000, Schuettpelz et al. 2001, etc.). detailed anatomical investigation of this Ehrendorfer (1995) and Ehrendorfer and genus. Samuel (2000, 2001) have realized the very The analysis of the data on karyology of important aim to compare and synthsize the Anemone taxa (Zhukova 1961, 1965a, data on morphology and DNA analysis. 1965b, Madahar 1967, Baumberger 1970, Their strict consensus trees resulting from Starodubtsev 1987, 1991, etc.) has shown the survey of about 30 selected morphological presence of two basic types of chromosome characters of more than 40 Anemone species, sets,x=8andx=7.Baumberger (1970) and the cladograms from cpDNA restriction considered the former as the initial one site data differed substantially, especially in (three acrocentric, one submetacentric and the basal branching patterns. Therefore, pro- four metacentric chromosomes), with x = 7 posals on taxonomy of Anemone by these being derivative. There were several at- authors differ much from the systems of their tempts to group 8-chromosome (Anemone predecessors. s. str., Carolinianae, Eriocapitella, Virginia- The foregoing patterns in the study of nae, Rupicolae, Tuberosae, etc.) and 7- Anemone are very important but very com- chromosome (Crassifoliae, Richardsonia, plicated. Since sufficient data for discussion Anemonidium, Himalayicae, Omalocarpus, of the comparison and synthesis of molecular etc.) Anemone species into two large associa- data are absent, it seemed to us to realise the tions. But in fact, in this genus plants with detailed comparative morphological study of x=7were found within species normally Anemone for clarification of relationships of having x = 8 (viz., in Anemone s. str., their groups of species. Eriocapitella, Baldenses, Tuberosae, Anemonanthea, etc.) and vice versa (viz., in Diagnostic significance of the morphologic Anemonidium, Himalayicae, Omalocarpus, characters in genus Anemone etc.). Therefore, it seems to be more realistic Here we present the grouping of essential not to exaggerate the taxonomic significance morphological characters of Anemone taxa of the above chromosome sets, but to pay into sets for distinguishing the main groups peculiar attention to size and arrangement of of species (sections) on the basis of the chromosomes and to the occurrence of results of our comparative analysis of the polyploidy in various groups. Anemone species. The recent cytological investigations Taking into consideration the essential (Marks and Schweizer 1974, Cullis and carpel and fruit characters, we divided all Schweizer 1974, Hagemann et al. 1993, etc.) Anemone taxa into two large groups. The have shown the significance of the evolu- first group includes 12 sections (82 species) tionary dynamics of chromosome banding characterized by a densely pubescent carpel patterns and accumulation of highly repeti- and achenes, symmetrical, mainly ovoid and tive DNA in Anemone species. The rather sharply narrowed into substraight styles, important results of study of the taxonomic meanwhile, the second group includes 9 sec- relationships in some Anemone groups were tions (36 species) having glabrous (some- obtained recently on cpDNA analysis by re- times subglabrous) carpels and achenes, striction endonucleases and cpDNA analysis mainly asymmetrical, oblong-ovoid and and restriction endonucleases and cpDNA gradually narrowed into curved or hooked probes (Hantula et al. 1989, Hoot 1995, Hoot styles. These groups are artificial because 136 植物研究雑誌 第83巻第3号平成20年6月 they embrace diverse taxa, for instance, both 3. Sect. Eriocephalus of them include plants having sessile or Carpels and achenes sessile, ovoid or stalked, compressed or not compressed spindle-like, apically narrowed, not com- carpels and achenes, with linear or dilated pressed, densely pubescent (hairs 1–7 mm stigmas. Nevertheless, we believe this divi- long), styles narrow, 1–3 mm long; carpel sion is useful to produce a key for determina- stigmas linear or dilated. Tepals 5–9, ana- tion of Anemone plants at any taxonomic stomosing veins from solitary to more than level. 10. Plants with caudices or non-tuberous We accept the sections as the main taxo- rhizomes and sympodial or monopodial nomic groups within the genus Anemone and scapes. for their more precise distinction we used the Within this section we recognize four se- essential characters of flowers, achenes and ries (Rupicolae, Sylvestres, Virginianae and vegetative organs. Below we present the Multifidae) which differ by compressed or brief characteristics of the Anemone sections not compressed carpels and achenes, length which differ from ones worked out by of their hairs, linear or dilated carpel stig- Tamura (1995). mas, sympodial or monopodial scapes, many or solitary flowers, basal and involucral leaf 1. Sect. Anemone shape, and types of underground shoots Carpels and achenes sessile, ovoid or (caudices, rhizomes or runners). subglobose, mainly compressed, densely pu- bescent (hairs 2–5 mm long); styles narrow, 4. Sect. Parviflora 0.5–3 mm long; carpel stigmas linear. Tepals Carpels and achenes sessile, ovoid, not 5–18, typically having anastomosing veins. compressed (with hairs 0.1–0.3 mm long), Plants with tuberous rhizomes and styles narrow, ca. 1 mm long, carpel stigmas sympodial scapes. linear. Tepals 5, monomorphic, without Within this section we recognize four anastomosing veins. Plants with long creep- series (Anemone, Somaliense, Biflora and ing rhizomes and monopodial scapes. Carolinianae) which differ by ovoid or subglobose, compressed or not compressed 5. Sect. Anemoclema carpels and achenes, length of their hairs, de- Carpels and achenes sessile, spindle-like, ciduous or persistent tepals, with solitary or slightly compressed (hairs 3–5 mm long); numerous anastomosing veins (sometimes styles narrow, 5–10 mm long, stigmas linear. absent), length of styles and hairs, and Tepals 5, monomorphic, with more than 10 basally narrow or sharply dilated basal leaf anastomosing veins. Herbaceous semi- petioles. rosetteous plants with caudices and mono- podial scapes. 2. Sect. Eriocapitella Carpels and achenes basally distinctly 6. Sect. Pulsatilloides stalked, inversely-ovoid, apically dilated, Carpels and achenes sessile, spindle-like, lanate (hairs 3–7 mm long), styles funnel- not compressed (hairs 2–5 mm long); styles like, shorter than 1 mm, stigmas capitate. narrow, 4–5 mm long, stigmas linear. Tepals Tepals 5–20, anastomosing veins more than 15–25, dimorphic, with 5–9 anastomosing 10. Plants with caudices and monopodial veins. Non-rosetteous semi-shrubs with scapes. caudices and sympodial scapes. June 2008 Journal of Japanese Botany Vol. 83 No. 3 137

7. Sect. Alchimillifolia of involucral leaf petioles. Carpels and achenes sessile, oblong-ovoid, not compressed (hairs 2–4 mm long); styles 11. Sect. Tuberosa narrow, 5–10 mm long. Tepals 10–12, Carpels and achenes sessile, slightly com- monomorphic or dimorphic, with more than pressed, ellipsoid, sparsely puberulent (hairs 10 anastomosing veins. Herbaceous semi- ca. 1 mm long); styles 0.2–1.2 mm long, rosetteous plants with caudices and sympo- stigmas linear or subcapitate. Tepals 8–12, dial scapes. dimorphic, with 1–9 anastomoses. Plants with tuberous rhizomes and sympodial rhi- 8. Sect. Kilimandscharica zomes. Carpels and achenes shortly stalked, Within this section we recognize two se- cylindroid, not compressed (hairs 3–4 mm ries (Tuberosae and Caucasicae) which dif- long); styles narrow, ca. 1 mm long; stigmas fer by width of achene ribs, length of achene capitate. Tepals 10–15, dimorphic, with soli- styles, shape of carpel stigmas (linear or tary anastomosing veins. Herbaceous subcapitate), tepal length and number, length semirosetteous plants with short ascending of involucral leaf petioles and shape of rhi- rhizomes and sympodial scapes. zomes.

9. Sect. Anemonanthea 12. Sect. Stolonifera Carpels and achenes sessile, ovoid (hairs Carpels and achenes sessile, ovoid (hairs shorter than 1 mm long); styles narrow, 1– ca. 1 mm long), styles hardly recognizable; 1.5 mm long; stigmas mainly linear. Tepals carpel stigmas dilated or subcapitate. Tepals typicaly monomorphic (in some taxa dimor- 5–7, anastomoses 3–9 or absent. Plants with phic). Plants with long or short rhizomes and dimorphic rhizomes and sympodial scapes. sympodial scapes. Within this section we recognize two se- Within this section we recognize five se- ries (Baicalenses and Flaccidae) which dif- ries (Anemonanthea, Altaicae, Nikoenses, fer by presence or absence of scale-like basal Reflexae and Quinquefoliae) which differ by leaves, basal leaves with distinct blades de- number and venation of tepals, width of veloping after or before anthesis, and pres- involucral leaf petioles, and shape of rhi- ence or absence of stolons. zomes (monomorphic or dimorphic, long or short). 13. Sect. Keiskea Carpels and achenes stalked, ovoid (hairs 10. Sect. Rosulantes 0.1–0.3 mm long), styles distinct, stigmas Carpels and achenes sessile, ovoid, mainly linear. Tepals 5–22, anastomoses 5–13. subglabrous (hairs ca 0.1 mm long), styles Plants with long rhizomes and sympodial narrow, 1–2 mm long, stigmas linear or rhizomes. slightly dilated. Tepals 5, monomorphic, Within this section we recognize two se- anastomoses mainly absent. Plants with non- ries (Keiskea and Deltoidea) which differ by tuberous rhizomes and sympodial scapes. length of achene stalks, achene shape and Within this section we recognize two se- villosity, number of tepals, their shape and ries (Rosulantes and Exiguae) which differ venation, and shape of rhizomes. by achene shape (compressed or not com- pressed, with ribs or without them), villosity 14. Sect. Omalocarpus of tepals, presence or absence of staminodes, Carpels and achenes sessile, subglobose- the number of scapes and flowers, and length obovate, compressed, with lateral wings 1– 138 植物研究雑誌 第83巻第3号平成20年6月

2mmwide, subglabrous, styles mainly bent 2–10 mm long, stigmas linear. Tepals (4–) or hooked, 1–2 mm long, carpel stigmas lin- 6–10, with anastomosing veins or without ear. Tepals 5–10, having 1–7 anastomosing them. Plants with caudices or short rhizomes veins. Plants with short vertical rhizomes and sympodial scapes. and monopodial scapes. Within this section we recognize four se- Within this section we distinguish three ries (Rivularidium, Angustilobae, Mexicanae series (Involucratae, Involucellatae and and Jamesonii) which differ by the length of Fuscopurpurea) which differ by shape of achenes and their styles, tepal number and achene styles, tepal length, filament shape, venation, the length of involucral leaf peti- types of inflorescences and shape of basal oles and underground shoots (caudices or leaf blades. rhizomes).

15. Sect. Imbricata 19. Sect. Begoniifolia Carpels and achenes sessile, broadly ellip- Carpels and achenes sessile, rhombic- soid, compressed, with lateral wings ca. 2 ovoid, compressed, with solitary lateral ribs, mm wide, glabrous, styles bent, ca. 1 mm subglabrous, styles ca. 1 mm long, stigmas long, carpel stigmas linear. Tepals 5–9, with- linear. Tepals 5, having anastomosing veins. out anastomosing veins. Plants with short Plants with short rhizomes and sympodial rhizomes and monopodial scapes. scapes.

16. Sect. Himalayicae 20. Sect. Richardsonia Carpels and achenes sessile, ovoid, typi- Carpels and achenes sessile, spindle-like, cally not compressed, densely pubescent not compressed, glabrous, styles narrow, (hairs ca. 1 mm long) or subglabrous, styles 5–6 mm long, carpel stigmas linear. Tepals narrow, 1–2.5 mm long, carpel stigmas lin- 5–6(–8), with solitary anastomosing veins. ear. Tepals 5–8, without anastomosing veins. Plants with long horizontal rhizomes and Plants with short vertical rhizomes and sympodial scapes. monopodial scapes. Within this section we recognize three 21. Sect. Crassifolia series (Obtusilobae, Trullifoliae and Carpels and achenes sessile, ellipsoid, Rupestres) which differ by compressed or compressed, with paired lateral ribs, basally not compressed carpels and achenes, densely sparsely pubescent (hairs 1–2 mm long), pubescent or subglabrous, filaments shape, styles narrow, ca. 1 mm long, carpel stigmas and shape of basal leaf blades and petioles. linear. Tepals 6–7, having anastomosing veins. Plants with tuberous and stolon-like 17. Sect. Rigida rhizomes and sympodial scapes. Carpels and achenes sessile, ovoid, not compressed, glabrous but with minute 22. Sect. Hepaticifolia spinous projections, styles 1–2 mm long, Carpels and achenes sessile, cylindroid, stigmas linear. Tepals 5–7, vein anstomoses slightly compressed, smooth, subglabrous, more than 10. Plants with short rhizomes and styles narrow, 1–2 mm long, carpel stigmas sympodial scapes. linear. Tepals 5, without anastomosing veins. Anther connectives with large subglobose 18. Sect. Rivularidium projections. Plants with short rhizomes and Carpels and achenes sessile, oblong-ovoid, sympodial scapes. subglabrous, styles conic, apically hooked, June 2008 Journal of Japanese Botany Vol. 83 No. 3 139

23. Sect. Anemonidium dscharica from subgen. Pulsatilloides as the Carpels and achenes sessile, ellipsoid, monotypic subgen. Kilimandscharica, sect. compressed, with lateral wings 0.5–1.5 mm Stolonifera as subgen. Stolonifera and sect. wide, styles narrow, 2–6 mm long, carpel Keiskea as subgen. Keiskea from subgen. stigmas linear. Tepals 4–5, anastomosing Anemonanthea. Moreover, we moved sect. veins more than 10. Non-rosetteous plants Begoniifolia from subgen. Omalocarpus in with short rhizomes, stolon-like runners and subgen. Rivularidium.Werecognized the sympodial scapes. subgenera Pulsatilloides, Anemoclema and Rigida sensu Tamura (1991) but in fact Key to subgenera, sections, subsections, sensu Juzepchuk (1937). At last, we accepted series and species of Anemone monotypic sections Hepaticifolia, Crassi- Our Key for determination of Anemone folia and Richardsonia as the subgenera sections, subsections, series and species was Hepaticifolia, Crassifolia and Richardsonia. elaborated on the basis of the analysis of the about 70 characters of the achenes, flowers, 1a. Carpels and achenes more or less leaves, above-ground and underground densely pubescent, symmetrical, mainly shoots and roots. ovoid, sharply narrowed into sub- The most complicated step of our work straight styles ...... 2 with the key of Anemone taxa was the at- 1b. Carpels and achenes glabrous or tempt to follow Tamura (1995) in recogni- subglabrous, typically assymetrical, ob- tion of subgenera. Unfortunately, most of long-ovoid, mainly gradually narrowed them were described by a few achene char- into curved or hooked styles ...... 83 acters only, viz., for subgen. Rivularidium 2a. Carpels and achenes embedded into this author noted glabrous achenes with dense hairs, mainly longer than their hooked beaks, despite these characters oc- bodies; tepals densely pubescent ...... 3 curring in other Anemone subgenera. For 2b. Carpels and achenes covered with hairs several subgenera Tamura used few other shorter than their bodies; tepals scarcely characters, viz., in his opinion subgen. pubescent or glabrous ...... 34 Anemonanthea is characterized by horizontal 3a. Plants with tuberous rhizomes; scapes or tuberous rhizomes and subgen. Omalo- sympodial; carpels and achenes mainly carpus by erect, not tuberous rhizomes. compressed, 2–5 mm long; tepals typi- Meanwhile, these peculiarities are also found cally having anastomosing veins, in other subgenera. Therefore, while examin- mainly densely pubescent (subgen. 1. ing the forementioned characteristics of Anemone sect. 1. Anemone) ...... 4 subgenera sensu Tamura, we added to them 3b. Plants with caudices or non-tuberous also the most essential peculiarities of shoots rhizomes; scapes sympodial or mono- and roots (life forms) and took into consid- podial; carpels and achenes not com- eration the basic chromosome numbers (x = pressed; tepals having or nor having 8 and x = 7). anastomosing veins, mainly glabrous .... As a result, we accepted within genus ...... 19 Anemone 15 subgenera because we divided 4a. Carpels and achenes ovoid, not com- subgen. Anemone on three ones: subgen. pressed, with styles 1.5–3 mm long and Anemone sensu Adanson (1763), subgen. hairs 2–4.5 mm long; tepals mainly Eriocapitella (= genus Eriocapitella Nakai) elliptic-obovate, with 3–13 basal veins and subgen. Eriocephalus sensu Juzepchuk and 0–30 anastomosing veins; basal leaf (1937). Besides, we separated sect. Kiliman- blades 3-sected or 3-parted (rarely 3–5- 140 植物研究雑誌 第83巻第3号平成20年6月

lobed) ...... 5 involucral leaves with petiole-like 4b. Carpels and achenes subglobose, dis- bases; tuberous rhizomes irregular ... 10 tinctly compressed, with styles 0.5–1.2 9b. Basal leaves 1–2, little divided; mm long and hairs 4–5.7 mm long; involucral leaves subsessile; tuberous tepals linear-oblong or lanceolate, with rhizomes subglobose ...... 12 5 basal veins and without vein 10a. Tepals yellow, 10–15 mm long, with 1 anastomose; basal leaf blades 1–2- –3 anastomosing veins; filaments linear; ternate (subsect. 4. Carolinianae) ... 13 basal leaf petiolules equal ...... 5a. Tepals deciduous, 20–50 mm long, with ...... 5. A. biflora 3–9 basal veins and 1–17 anastomosing 10b. Tepals red or yellow, 15–30 mm long, veins; basal leaf petioles basally sharply with 7–30 anastomosing veins; fila- dilated (“ear-like”); basal and involucral ments dilated; basal leaf petiolules une- leaf blades pubescent (subsect. 1. qual (central petiolule distinctly longer Anemone) ...... 6 than lateral ones) ...... 11 5b. Tepals persistent, 10–30 mm long, with 11a. Tepals red or reddish; filaments slightly 5–13 basal veins and 0–30 anastomos- dilated; achenes 2.5–3 mm long, with ing veins; basal leaf petioles without hairs 2.5–3.5 mm long and glabrous “ear-like” basal parts; basal and styles ...... 6. A. bucharica involucral leaf blades glabrous ...... 8 11b. Tepals yellow; filaments sublanceolate; 6a. Tepals with 10 or more anastomosing achenes 3.5–5 mm long, with hairs 4– veins, mainly red; basal and involucral 5.5 mm long and apically pubescent leaves multi-divided; stolon-like rhi- styles ...... 7. A. baissunensis zomes present ...... 1. A. coronaria 12a. Achenes 3–3.5 mm long, with hairs 5– 6b. Tepals with 1–3 anastomosing veins, 6mmlong and styles 1.7–2.5 mm long; usually not red; basal and involucral tepals 15–30 mm long, white or bluish, leaves little divided; stolon-like rhi- with 7–15 anastomosing veins, pilose zomes absent ...... 7 outside; basal and involucral leaf blades 7a. Achene hairs 3.5–5 mm long and styles 3-parted ...... 8. A. tschernjaewi 3.5–4 mm long; flowers solitary; basal 12b. Achenes 1.7–2.3 mm long, with hairs leaves dimorphic, 3-sected ...... 1.7–2.3 mm long and styles 0.5–1.2 mm ...... 2. A. hortensis long; tepals 5–8 mm long, yellowish- 7b. Achene hairs 3–3.5 mm long and styles green, without anastomosing veins, 4.5–5 mm long; scapes few-flowered; glabrous; basal and involucral leaf basal leaves monomorphic, 3-lobed ...... blades 3-sected .... 9. A. serawschanica ...... 3. A. palmata 13a (4b). Achenes with ribs ca. 0.2 mm 8a (5b). Achene hairs 3–3.5 mm long; fruit- wide; scapes 1-flowered; leaf blades ing heads elongated; tepals blue or glabrous; tubers suborbicular; stolon- mauve, glabrous; basal leaf blades like rhizomes present ...... palmately 3-parted (subsect. 2...... 10. A. caroliniana Somaliense) ...... 4. A. somaliensis 13b. Achenes with ribs 0.2–0.5 mm wide; 8b. Achene hairs 2–5 (–6) mm long; fruit- scapes 1–few-flowered; leaf blades pu- ing heads globose; tepals yellow, red or bescent or glabrous; tubers elongate; whitish, pubescent; basal leaf blades 3- stolon-like rhizomes absent ...... 14 parted to 3-sected (subsect. 3. Biflorae) 14a. Achenes with ribs 0.3–0.5 mm wide; ...... 9 tepals sparsely pubescent; basal leaves 9a. Basal leaves more than 2, many-sected; 1–2-ternate, with petiolate scale-like June 2008 Journal of Japanese Botany Vol. 83 No. 3 141

“ears” ...... 15 Eriocapitella) ...... 20 14b. Achenes with ribs 0.2 mm wide or in- 19b. Plants with caudices or rhizomes; distinct; tepals densely pubescent; basal scapes monopodial or sympodial, leaves typically 2-ternate, without inflorescences typically few-flowered; “ears” ...... 16 carpels and achenes sessile or 15a. Achenes 2.7–3.5 mm long; tepals 8–12, subsessile, apically narrowed, densely 12–20 mm long, reddish-white; scapes pubescent , with narrow styles longer 1-flowered; basal leaves sparsely pubes- than 1 mm and linear stigmas (rarely in cent; involucral leaves basally connate.. carpels dilated (subgen. 3. Eriocepha- ...... 11. A. berlandieri lus) ...... 22 15b. Achenes 1.3–2.2 mm long; tepals 6–8, 20a. Carpels and achenes covered with hairs 10–12 mm long, white or blue; scapes 2–3 mm long; basal leaf blades 3–7- mostly 2–3-flowered; basal leaves lobed ...... 18. A. vitifolia glabrous; involucral leaves basally dis- 20b. Carpels and achenes covered with hairs tinct ...... 12. A. edwardsiana 3–7 mm long; leaf blades ternate .... 21 16a. Achene hairs 2–3.5 mm long; tepals 6 21a. Ovaries 1.5–2 mm long, on stalks ca. 1 –10 ...... 17 mm long, covered with hairs 2–4 mm 16b. Achene hairs 4–6 mm long; tepals more long; tepals 20–30 mm long, than 10 ...... 18 anastomosing veins 5–7; leaf blades 17a. Tepals 10–20 mm long, pink or white; abaxially strigose .... 17. A. hupehensis basal leaves glabrous; basal and 21b. Ovaries 0.5–0.7 mm long, on stalks ca. involucral leaves similar ...... 0.3 mm long, covered with hairs 4–6 ...... 13. A. tuberosa mm long; tepals 15–20 mm long, 17b. Tepals 6–12 mm long, greenish-white; anastomosing veins more than 30; leaf basal leaves pubescent; basal and blades abaxially tomentose ...... involucral leaves dissimilar ...... 19. A. tomentosa ...... 14. A. okennonii 22a. Plants with caudices or short rhizomes; 18a. Tepals 8–20, 2–5 mm, with 5–7 basal scapes sympodial or monopodial, veins and 1–2 anastomosing veins; fila- inflorescences 1- to several-flowered; ments linear; inflorescences few- carpels and achenes ovoid or spindle- flowered; basal and involucral leaves like, compressed or not compressed; dissimilar, basal leaves dimorphic ...... tepals 5–9, anastomosing veins from ...... 15. A. decapetala solitary to more than 10 (sect. 3. 18b. Tepals 10–15, 2–3 mm, with 3 basal Eriocephalus) ...... 23 veins and without anastomosing veins; 22b. Plants with long creeping rhizomes; filaments filiform; flowers solitary; scapes monopodial, flowers solitary; basal and involucral leaves similar, carpels and achenes ovoid, not com- basal leaves monomorphic ...... pressed; tepals 5, without anastomosing ...... 16. A. triternata veins (sect. 4. Parviflora) ...... 19a (3a). Plants with caudices; scapes ...... 20. A. parviflora monopodial, inflorescences compound, 23a. Carpels and achenes ovoid, basally di- 2–3-branched; carpels and achenes lated, with styles 2–3(–6) mm long, basally distinctly stalked, apically carpel stigmas linear; tepals with few dilated, lanate, with funnel-like styles anastomosing veins; scapes mono- shorter than 1 mm and capitate stigmas podial; involucral leaves subsessile; (subgen. 2. Eriocapitella sect. 2. plants with caudices (ser. 4. Multifidae) 142 植物研究雑誌 第83巻第3号平成20年6月

...... 30 3–5, with petioles 3–7(–12) cm long .... 23b. Carpels and achenes obovoid or ...... 24. A. virginiana spindle-like, with styles 0.5–1.5 mm 28b. Achene styles 0.5–1 mm long; fruiting long; carpel stigmas linear or globose; heads cylindroid; tepals 7–15 mm long, tepals with many anastomosing veins; with less than 10 anastomosing veins; scapes sympodial or monopodial; involucral leaves 5–9, with petioles 1– involucral leaves petiolate; plants with 3cmlong ...... 25. A. cylindrica rhizomes ...... 24 29a (24b). Achenes basally distinctly nar- 24a. Carpel stigmas mostly linear; tepals 5– rowed, with styles ca. 1 mm long; tepals 9; scapes few-flowered ...... 25 5–25 mm long, with wide bases and 24b. Carpel stigmas globose; tepals 5(–6); apices; involucral leaf petioles 1–2 cm scapes 1-flowered (ser. 3. Sylvestres) .... long ...... 26. A. sylvestris ...... 29 29b. Achenes basally not narrowed, with 25a. Achenes not compressed, 2–3 mm long, styles ca. 0.5 mm long; tepals 10–15 with hairs 4–7 mm long; basal and mm long, with narrow bases and wide involucral leaf blades 3-sected or 3- apices; involucral leaf petioles 4–10 cm parted; involucral leaf petioles 5–10 long ...... 27. A. ochotensis mm long (ser. 1. Rupicolae) ...... 26 30a (23a). Achene styles 1.5–2 mm long; 25b. Achenes compressed, 2–4 mm long, tepals 5–6, with narrow bases and api- with hairs 2–3 mm long; basal and ces, anastomosing veins present; scapes invlolucral leaf blades ternate; few-flowered, 10–40 cm long; basal involucral leaf petioles 1–5 cm long leaf blades 2–3-ternate; involucral (ser. 2. Virginianae) ...... 28 leaves similar to basal ones ...... 31 26a. Tepals 5–9, purplish or yellowish; 30b. Achene styles 3–6 mm long; tepals 5– carpel stigmas globose; basal and 15, with wide bases and apices, involucral leaf blades 3-sected; anastomosing veins absent; scapes 1- involucral leaf petioles ca. 1 cm long; flowered, 5–25 cm long; basal leaf stolon-like root-runners present ...... blades ternate-palmatid or pinnatifid; ...... 21. A. rupicola involucral leaves reduced ...... 33 26b. Tepals 5, white or greenish; carpel stig- 31a. Achenes subellipsoid, 3–4 mm long, mas linear; basal and involucral leaf with hairs 3–6 mm long; tepals 6–10, blades 3–5-parted; involucral leaf peti- 8–15 mm long, blue to reddish, with 0– oles less 1 cm long; root-runners absent 1 anastomosing veins; scapes typically ...... 27 2–7-flowered; basal leaf blades 2- 27a. Tepals white, 20–25 mm long; anther ternate; involucral leaves 3-parted to 3- connectives without projections; basal lobed ...... 28. A. multifida leaf blades densely puberulent ...... 31b. Achenes subglobose, 2–3 mm long, ...... 22. A. laceratoincisa with hairs 2–3 mm long; tepals 5–8, 10– 27b. Tepals greenish, 15–20 mm long; anther 20 mm long, white, with 1–3 anasto- connectives with slight projections; mosing veins; scapes 1–2-flowered; basal leaf blades subglabrous ...... basal leaf blades 3-ternate; involucral ...... 23. A. tibetica leaves ternate ...... 32 28a (25b). Achene styles 1–1.5 mm long; 32a. Fruiting heads oblong-ovoid; tepals 12– fruiting heads oblong-ellipsoid; tepals 20 mm long, elliptic-ovate; filaments 10–20 mm long, with more than 10 sublanceolate; scapes 10–15 cm long .... anastomosing veins; involucral leaves ...... 29. A. baldensis June 2008 Journal of Japanese Botany Vol. 83 No. 3 143

32b. Fruiting heads subglobose; tepals 10–12 anastomosing veins 5 to more than 10; mm long, elliptic-oblong; filaments lin- basal leaf blades 3-ternate or palmately ear; scapes 25–40 cm long ...... lobed and acutely dentate ...... 37 ...... 30. A. pavoniana 37a. Non-rosulate semishrubs; carpels and 33a (30b). Tepals 7–15, 8–20 mm long, achenes spindle-like; tepals 15–25, whitish-blue, sometimes dimorphic; linear-lanceolate, anastomosing veins filaments filiform; scapes 10–25 cm 3–9, densely pubescent; leaf blades 3- long; basal leaves palmatifid, villous .... ternate, subglabrous (subgen. 5...... 31. A. drummondii Pulsatilloides 6. sect. Pulsatilloides) ..... 33b. Tepals 5–6, 10–12 mm long, dark blue ...... 34. A. capensis to reddish, monomorphic; filaments di- 37b. Semi-rosulate herbaceous plants; lated; scapes 5–15 cm long; basal leaves carpels and achenes oblong-ovoid; pinnatifid, subglabrous ...... tepals 10–12, wide-lanceolate, anasto- ...... 32. A. multiceps mosing veins more than 10, sparsely 34a (2b). Plants with caudices or rhizomes; pubescent; leaf blades palmately-lobed, scapes sympodial, flowers solitary; villous (sect. 7. Alchimillifolia) ...... 38 carpels and achenes compressed or not 38a. Carpels and achenes 4–5 mm long, cov- compressed, covered by hairs 4–10 mm ered with dimorphic hairs 1–5 mm long; long, styles densely pubescent ...... 35 tepals monomorphic; filaments dis- 34b. Plants with rhizomes; scapes sympodial tinctly dilated; basal leaf blades 5–7 cm or monopodial, inflorescences 1– wide ...... 35. A. caffra several-flowered; carpels and achenes 38b. Carpels and achenes 5–10 mm long, typically not compressed, covered with covered with monomorphic hairs 3–6 hairs 1–3 mm long, styles glabrous ...... mm long; tepals dimorphic; filaments ...... 39 slightly dilated; basal leaf blades 12–20 35a. Plants with caudices; carpels and cm wide ...... 36. A. fanninii achenes sessile, oblong-ovoid or spin- 39a (34b). Plants with ascending or horizon- dle-like, 5–10 mm long; styles densely tal rhizomes; scapes sympodial or pubescent, stigmas linear; tepals with monopodial; involucral leaves petiolate, more than 5 anastomosing veins, similar to basal leaves and frequently of densely pubescent ...... 36 a larger size; carpels and achenes cov- 35b. Plants with tuberous rhizomes; carpels ered with hairs shorter than 1 mm (usu- and achenes shortly stalked, cylindroid, ally 0.1–0.2 mm) ...... 40 3–5 mm long; styles 1–2 mm long; 39b. Plants with short vertical rhizomes; stigmas capitate; tepals with solitary scapes monopodial; involucral leaves anastomosing veins, sparsely pubescent sessile, reduced (much smaller than (subgen. 6. Kilimandscharica sect. 10. basal leaves); carpels and achenes cov- Kilimandscharica) ..... 37. A. thomsonii ered with hairs longer than 1 mm 36a. Carpels and achenes slightly com- (subgen. 9. Omalocarpus sect. 15. pressed; styles straight; tepals 5, Himalayicae) ...... 73 anastomosing veins ca. 30; basal leaf 40a. Plants with short or long, sometimes blades pinnatipartite and obtusely den- tuberous rhizomes; involucral leaves tate (subgen. 4. Anemoclema sect. 5. petiolate, carpels and achenes sessile, Anemoclema) ...... 33. A. glaucifolia with distinct styles and mainly linear 36b. Carpels and achenes not compressed; stigmas; basic chromosome number x = styles apically curved; tepals 10–25, 8 (subgen. 6. Anemonanthea) ...... 41 144 植物研究雑誌 第83巻第3号平成20年6月

40b. Plants with short or long but non- wedge-like bases ...... 38. A. nemorosa tuberous rhizomes; involucral leaves 45b. Achene stigmas sublinear; tepals 5–8, sessile, carpels and achenes sessile or monomorphic, basally narrowed, lack- stalked, with distinct or hardly recog- ing anastomosing veins; central leaflets nizable styles and mainly capitate stig- of basal leaf blades 3-sected or 3- mas; basic chromosome number x = 7 .. parted, rhombic, with semicordate bases ...... 68 ...... 39. A. amurensis 41a. Achene styles mainly 1–1.5 mm long; 46a. Tepals basally narrowed, mainly blue or basal leaves several, scale-like, persis- whitish, densely puberulent; filaments tent; solitary leaves with distinct blades basally dilated ...... 40. A. caerulea and long petioles developing on rhi- 46b. Tepals basally rounded, mainly rose or zomes after anthesis (sect. 8. red, sparsely puberulent; filaments Anemonanthea) ...... 42 apically dilated ...... 41. A. uralensis 41b. Achene styles mainly 0.5–0.7 mm long; 47a (43b). Cymes 1–few-flowered; tepals basal leaves several, with distinct blades pubescent, usually with anastomosing and long petioles developing on repro- veins; achenes slightly compressed, ductive shoots before anthesis ...... 61 with lateral ribs ...... 48 42a. Tepals 5–8 (rarely 9–10); rhizomes 47b. Cymes 1-flowered; tepals glabrous or monomorphic, long and 1–3 mm in di- subglabrous, lacking anastomosing ameter, branced, mainly horizontal (ser. veins; achenes compressed or not com- 1. Anemonanthea) ...... 43 pressed ...... 50 42b. Tepals 5–12; rhizomes dimorphic, 48a. Basal and involucral leaflets 2–3-lobed short, thick (nodulose) or long, thin ...... (sometimes parted); involucral leaf peti- ...... 52 oles 3–5 cm long; tepals dimorphic, 43a. Carpel styles ca. 1 mm long; stigmas basally narrowed; achene styles 0.8–1.5 subcapitate or slightly dilated; mm long ...... 42. A. ranunculoides involucral leaf petioles 3–5 mm wide .... 48b. Basal and involucral leaflets undivided; ...... 44 involucral leaf petioles 1–3 cm long; 43b. Carpel styles longer than 1 mm; stigmas tepals monomorphic, basally rounded; linear; involucral leaf petioles 1–2 mm achene styles 1.5–2.5 mm long ...... 49 wide ...... 47 49a. Cyme 1-flowered; tepals 5, mainly 44a. Achenes (2–) 3–5 mm long; tepals 5–8, pilose, having 5–9 anastomosing veins glabrous; cymes 1-flowered; involucral ...... 43. A. udensis leaf petioles 1–3 cm long; involucral 49b. Cymes few-flowered; tepals 5–8, leaf blades similar to the basal leaf glabrous, having solitary anastomosing blades ...... 45 veins ...... 44. A. trifolia 44b. Achenes 1–2 mm long; tepals 5, pubes- 50a (47b). Central segments of basal leaf cent; cymes 1–2-flowered; involucral blades 3-lobed; tepals 5; achene hairs leaf petioles 3–5 mm long; involucral 0.2–0.3 mm long ...... 45. A. umbrosa leaf blades larger than the basal leaf 50b. Central segments of basal leaf blades blades ...... 46 serrate or coarsely toothed; tepals 5–7; 45a. Achene stigmas slightly dilated; tepals achene hairs ca. 0.01 mm long ...... 51 6–10, mainly dimorphic, basally 51a. Tepals 10–15 mm long, basally not nar- rounded, having 3–5 anastomosing rowed, white; achenes not compressed, veins; central leaflets of basal leaf without ribs, 4–5 mm long, sparsely blades 3-lobed, ellipsoid-oblong, with puberulent ...... 46. A. soyensis June 2008 Journal of Japanese Botany Vol. 83 No. 3 145

51b. Tepals 4–8 mm long, basally narrowed, 55b. Achenes sparsely puberulent; tepals 5– white-greenish; achenes slightly com- 7, spreading or bent, with rounded bases pressed, with narrow ribs, 2–4 mm and apices, 5–20 mm long, mainly with- long, densely puberulent ...... out anastomosing veins, subglabrous; ...... 47. A. debilis involucral leaf petioles 1–2 mm wide .... 52a (42b). Tepals 8–15, glabrous, spreading; ...... 56 cymes 1-flowered (ser. 2. Altaicae) ...... 56a. Achene hairs less than 0.1 mm long; ...... 53 tepals 5–7 1–2 mm, bent, sparsely 52b. Tepals 5–8, glabrous or pubescent, puberulent; cymes few-flowered; spreading or bent; cymes 1–few- bracteoles present (ser. 2. Reflexae) ...... flowered ...... 55 ...... 52. A. reflexa 53a. Ovaries and achenes basally narrowed, 56b. Achene hairs 0.1–2 mm long; tepals with narrow lateral ribs; achene hairs 10–25 4–10 mm, spreading, glabrous; 0.1–0.3 mm long; achene styles cymes 1-flowered; bracteoles absent puberulent; carpel stigmas subcapitate; (ser. 4. Quinquefoliae) ...... 57 achene stigmas slightly dilated; basal 57a. Achene styles sparsely puberulent; leaf petiolules 5–10 mm long; tepals tepals 5, having solitary anastomosing dimorphic ...... 48. A. altaica veins; involucral leaf petioles 0.5–2 cm 53b. Ovaries and achenes basally narrowed long ...... 58 or rounded, with ribs or lacking them; 57b. Achene styles glabrous; tepals 5–7, achene hairs 0.2–0.5 mm long; achene lacking anastomosing veins; involucral styles glabrous; carpel and achene stig- leaf petioles 1–3 cm long ...... 59 mas linear or sublinear; basal leaf 58a. Carpels and achenes covered with hairs petiolules 2–5 mm long; tepals 0.5–1 mm long; lateral leaflets often monomorphic ...... 54 deeply two-lobed; basal leaf blades 54a. Achenes basally rounded, not com- puberulent; involucral leaf blades and pressed, styles 0.5–1 mm long; tepals stems puberulent or subglabrous ...... 8–12, having 3–5 anastomosing veins; ...... 53. A. quinquefolia basal and involucral leaflets deeply 3- 58b. Carpels and achenes covered with hairs lobed; involucral leaf petioles 10–20  0.1–0.2 mm long; lateral leaflets all 3–5 mm, distinctly dilated ...... lanceolate, undivided; leaves and stems ...... 49. A. pseudoaltaica glabrous ...... 54. A. lancifolia 54b. Achenes basally narrowed, slightly 59a. Carpels and achenes densely covered compressed, styles 1–1.5 mm long; with hairs 1–2 mm long; scapes few, tepals 9–15, having 1–3 anastomosing puberulent; rhizomes mainly vertical .... veins; basal and involucral leaflets ...... 55. A. piperi medially 3-lobed or toothed; involucral 59b. Carpels and achenes covered with hairs leaf petioles 5–12 1–2 mm, basally 0.1–0.2 mm long; scapes solitary, slightly dilated ...... 50. A. raddeana glabrous; rhizomes horizontal or as- 55a (52b). Achenes densely puberulent; cending ...... 60 tepals 5 (–6), spreading, with narrow 60a. Achenes basally narrowed, slightly bases and apices, 15–30 mm long, hav- compressed, with narrow lateral ribs; ing 5–9 anastomosing veins, sparsely tepals 7–10 4–6 mm, mainly white or puberulent; involucral leaf petioles 3–5 blue ...... 56. A. grayi mm wide (ser. 3. Nikoenses) ...... 60b. Achenes basally rounded, non- ...... 51. A. nikoensis compressed, without ribs; tepals 10–20 146 植物研究雑誌 第83巻第3号平成20年6月

5–10 mm, red to blue ...... cral leaf blades 3-sected, smaller than ...... 57. A. oregana those in basal leaves ...... 61a (41b). Plants with non-tuberous rhi- ...... 62. A. scabriuscula zomes, 1–few-flowered cymes and 66a (61b). Carpels and achenes 3.2–3.6 mm gradually dilated basal leaf petioles; long, with ribs ca. 0.2 mm (or absent) achenes ovoid, subglabrous or glabrous; and curved styles 1–1.2 mm long; stig- tepals 5 (sect. 9. Rosulantes) ...... 62 mas linear; tepals 12–15, 15–20 mm 61b. Plants with tuberous rhizomes, solitary long; basal leaf petioles 10–20 cm long, flowers and basally sharply dilated involucral leaf petioles 1.5–3 cm long; basal leaf petioles; achenes ellipsoid, rhizomes cylindroid (ser. 1. Tuberosae) sparsely puberulent; tepals 8–12 (sect...... 67 10. Tuberosa) ...... 66 66b. Carpels and achenes 2–2.2 mm long, 62a. Achenes not compressed, without ribs; with ribs ca. 0.5 mm and slightly curved tepals puberulent, with or without styles 0.2–0.3 mm long; stigmas in anastomosing veins; staminodes pre- carpels subcapitate, in achenes dis- sent; scapes 2–3, few-flowered; tinctly dilated; tepals 8–10, 8–10 mm involucral leaf petioles 1–3 cm long; long; basal leaf petioles 5–8 cm long, bracteoles present, small (ser. 1. involucral leaf petioles 0.5–1.5 cm Rosulantes) ...... 63 long; rhizomes subspherical (ser. 2. 62b. Achenes slightly compressed, with nar- Caucasicae) ...... 65. A. caucasica row ribs; tepals subglabrous, lacking 67a. Carpels and achenes without lateral anastomosing veins; staminodes absent; ribs, scarcely pubescent; tepals of outer scapes solitary, 1-flowered; involucral whorl with 1–3 anastomosing veins; leaf petioles 0.5–1.5 cm long; basal leaf blades ternate, with petiolules bracteoles absent (ser. 2. Exiguae) ...... 3–5 mm long ...... 63. A. apennina ...... 64 67b. Carpels and achenes with lateral ribs 63a. Stigmas linear; tepals 5–10 mm long, and pubescent only at the base; tepals of lacking anastomosing veins; staminodes outer circle with 5–9 anastomosing between tepals and stamens ...... veins; basal leaf blades 3-sected, with ...... 58. A. stolonifera petiolules 1–2 mm long (frequently ab- 63b. Stigmas subcapitate; tepals 15–20 mm sent) ...... 64. A. blanda long, having 5–15 anastomosing veins; 68a (40b). Plants with dimorphic rhizomes staminodes between stamens and (short and stolon-like); involucral carpels ...... 59. A. davidii leaves 3–5, partly reduced; cymes 1–3- 64a. Carpels and achenes sparsely flowered; tepals 5–7; carpels and puberulent; involucral leaf petioles 3–5 achenes sessile (hairs ca. 1 mm long); mm wide, basally connate ...... styles hardly recognizable; stigmas ...... 60. A. exigua dilated or subcapitate (subgen. 7. 64b. Carpels and achenes glabrous; involuc- Stolonifera sect. 11. Stolonifera) .... 69 ral leaf petioles 1–2 mm wide, free ..... 68b. Plants with long rhizomes; involucral ...... 65 leaves 3, all reduced; flowers solitary; 65a. Carpel and achene styles nearly absent; tepals 5–13; carpels and achenes stalked stigmas subglobose; involucral leaf (hairs 0.1–0.3 mm long), styles distinct, blades ternate, larger than those in basal stigmas linear (subgen. 8. Keiskea sect. leaves ...... 61. A. griffithii 12. Keiskea) ...... 72 65b. Styles distinct; stigmas linear; involu- 69a. Scale-like basal leaves absent; basal June 2008 Journal of Japanese Botany Vol. 83 No. 3 147

leaves with distinct blades and basally long, achene bodies subglobose, hispid vaginate petioles developing before proximally, stigmas triangular-subulate anthesis; carpel stigmas subcapitate, (ser. 2. Deltoidea) ...... 71. A. deltoide achene stigmas slightly dilated (ser. 1. 73a (39b). Carpels and achenes ovoid, not Stolonifera) ...... 70 compressed, densely pubescent; leaf 69b. Scale-like basal leaves present; basal blades once 3-sected to 3-lobed or leaves with distinct blades and narrow sometimes undivided ...... 74 petioles developing after anthesis; 73b. Carpels and achenes ellipsoid, distinctly carpel stigmas capitate or subcapitate, compressed, sparsely pubescent or achene stigmas subcapitate or dilated subglabrous; leaf blades twice 3-sected (ser. 2. Flaccida) ...... 71 (ser. 3. Rupestres) ...... 82 70a. Rhizomes short, 8–10 mm in diameter, 74a. Filaments lanceolate (basally dilated, with long underground stolons; tepals apically narrowed); leaf blades typically 10–15 mm long, with 3–5 anastomosing wider than long, with bases cordate or veins; achene bodies basally narrowed, rounded (Ser. 1. Obtusilobae) ...... 75 slightly compressed, with narrow ribs ... 74b. Filaments ovate (basally and apically ...... 66. A. baicalensis narrowed); leaf blades typically longer 70b. Rhizomes short, 5–7 mm in diameter, than wide, with bases attenuate or trun- with long above-ground stolons; tepals cate (Ser. 2. Trullifoliae) ...... 79 8–10 mm long, without anastomosing 75a. Tepals pilose; staminodes absent; veins; achene bodies basally rounded, cymes mainly few-flowered; basal leaf not compressed, without ribs ...... petioles 1–2 mm wide; involucral leaf ...... 67. A. prattii blades 3-lobed to 3-parted ...... 71a. Basal leaves having 2–4 blades; scapes ...... 105. A. obtusiloba 1–3, cymes few-flowered; bracteoles 75b. Tepals mainly subglabrous; staminodes present; tepals 5–10 mm long, with 7– sometimes present; flowers solitary; 9 anastomosing veins; achenes sparsely basal leaf petioles more than 2 mm puberulent or glabrate, hairs 0.1 mm wide; involucral leaf blades undivided long ...... 68. A. flaccida or sometimes 3-lobed ...... 76 71b. Basal leaves having solitary blades; 76a. Tepals 5, 5–8 mm long; filaments scapes solitary, cymes 1-flowered; linear; basal leaf blade central segments bracteoles absent; tepals 7–15 mm long, 3-sected or 3-parted, leaf blades appear- lacking anastomosing veins; achenes ing pinnatifid, petioles 2–5 cm long ...... densely puberulent, hairs ca. 0.5 mm ...... 106. A. subpinnata long ...... 69. A. delavayi 76b. Tepals 5–8, 10–20 mm long; filaments 72a (68b). Rhizomes 1–2 mm in diameter; mostly lanceolate or ovate; basal leaf tepals 10–22, linear-lanceolate, having blade central segments 3-lobed or undi- few anastomosing veins, basally vided, petioles 5–20 cm long ...... 77 sparsely puberulent; achene stalks 1.5– 77a. Ovaries ellipsoid; staminodes absent; 2mmlong, achene bodies long- filaments ovate; basal leaf blades or- obovate, sparsely puberulent, stigmas bicular-ovate, all segments subsessile ... linear-lanceolate (ser. 1. Keiskea) ...... 107. A. patula ...... 70. A. keiskeana 77b. Ovaries narrow-ovoid; staminodes pre- 72b. Rhizomes 2–5 mm in diameter; tepals sent; filaments linear-lanceolate; basal 5, elliptic, without anastomosing veins, leaf blades ovate, central segments dis- glabrous; achene stalks 0.5–0.7 mm tinctly petiolulate ...... 78 148 植物研究雑誌 第83巻第3号平成20年6月

78a. Tepals 10–20 mm long, dimorphic; fila- (without wings or ribs), elongate; styles ments linear, 0.3–0.5 mm wide; basal up to 9 mm long; basic chromosome leaf blade central segment petiolules 1– number x = 8 ...... 84 2mmlong ...... 108. A. rockii 83b. Plants without caudices; scapes 78b. Tepals 5–10 mm long, monomorphic; sympodial or monopodial; carpels and filaments lanceolate, 0.5–1.0 mm wide; achenes distinctly compressed (having basal leaf blade central segment wide lateral wings), cylindroid or petiolules 5–10 mm long ...... suborbicular; styles up to 5 mm long; ...... 109. A. geum basic chromosome number x = 7 .... 102 79a (74b). Tepal anastomosing veins some- 84a. Plants with rhizomes; basal leaf blades times present; cymes 1–3-flowered; 3-parted; tepals up to 7, with more than basal leaf blades basally attenuate .... 80 10 anastomosing veins, glabrous; 79b. Tepal anastomosing veins absent; flow- achenes with minute spinous projec- ers solitary; basal leaf blades basally tions, basally sparsely pubescent; styles truncate ...... 81 1–2 mm long (subgen. 10. Rigida sect. 80a. Filaments elliptic; cymes 1–3-flowered; 16. Rigida) ...... 72. A. rigida basal leaf blades 3-parted, 3-cleft, or 3- 84b. Plants with caudices or rhizomes; basal lobed, broadly lanceolate or ovate; leaf blades from entire to ternate; tepals involucral leaf blades 3-lobed or 3- up to 15, mainly without anastomosing dentate ...... 110. A. trullifolia veins, sparsely pubescent (sometimes 80b. Filaments linear; flowers solitary; basal glabrous); achenes without projections, leaf blades usually undivided, oblong- glabrous or subglabrous; styles 1–9 mm linear to oblanceolate; involucral leaf long (subgen. 11. Rivularidium) ...... 85 blades undivided ...... 111. A. coelestina 85a. Plants with caudices or short rhizomes; 81a. Tepals 8–15 mm long; basal leaf blades basal and involucral leaf blades mainly 3-parted to 3-lobed; involucral leaf ternate; tepals 5 to 15, mainly without blades 3-lobed ...... anastomosing veins; carpels and ...... 112. A. yulongshanica achenes oblong-ovoid, compressed 81b. Tepals 5–10 mm long; basal leaf blades (with paired ribs) or not compressed, undivided or obscurely 3-lobed; typically glabrous; styles 2–9 mm long involucral leaf blades entire or 3- (sect. 17. Rivularidium) ...... 86 dentate ...... 113. A. subindivisa 85b. Plants with short rhizomes; basal and 82a (73b). Carpel and achene styles involucral leaf blades 3-lobed to entire; substraight and apically thin; tepals 5– tepals 5, having anastomosing veins; 10 mm long, glabrous or sparsely carpels and achenes rhombic-ovoid, puberulent; filaments lanceolate; compressed (lateral ribs solitary), staminodes absent; leaves subglabrous subglabrous; styles ca. 1 mm long (sect...... 114. A. rupestris 18. Begoniifolia) ...... 99 82b. Carpel and achene styles uncinate and 86a. Tepals 4–5 (–7), with 5–10 anastomos- apically thickened; tepals 7–15 mm ing veins; scapes compound, 2–3-branc long, puberulent; filaments linear; hed, many-flowered; involucral leaves staminodes present; leaves sparsely more than 3 ( and bracteoles) ..... puberulent ...... 115. A. polycarpa ...... 87 83a (1b). Plants with caudices or short rhi- 86b. Tepals 5–9(–15), mostly without anas- zomes; scapes sympodial; carpels and tomosing veins; scapes typically simple, achenes mainly slightly compressed few-flowered (sometimes flowers soli- June 2008 Journal of Japanese Botany Vol. 83 No. 3 149

tary); involucral leaves mostly 3 (ser. 4. vaginate ...... 76. A. angustiloba Jamesonii) ...... 93 91b. Achenes with hooked styles 0.3–0.5 87a. Achenes 3–8 mm long; tepals basally mm long; tepals 5, anastomosing veins densely barbate; basal and primary 7–9; basal leaf petioles basally sharply involucral leaves similar; plants with dilated ...... 77. A. sumatrana caudices (ser. 1. Rivularidium) ...... 88 92a. Tepals 12–15 mm long, red or whitish- 87b. Achenes 2.5–4 mm long; tepals pink, sparsely pubscent; connectives subglabrous or glabrous; basal and with projections; basal leaf petioles primary involucral leaves dissimilar shortly vaginate, blades ternate to (primary reduced and smaller); plants biternate, puberulent; involucral leaves with caudices or rhizomes ...... 90 sessile ...... 78. A. mexicana 88a. Achenes 5–8 mm long, with styles 2–3 92b. Tepals 6–12 mm long, white or yel- mm long; tepals 7–15 mm long, bluish lowish, glabrous; connectives without or reddish; basal leaf blades 5–10 5– projections; basal leaf petioles basally 15 cm, pubescent; involucral leaves 3- sharply dilated, blades palmately parted, parted ...... 73. A. rivularis glabrous; involucral leaves petiolate ..... 88b. Achenes 3–6 mm long, with styles 1–2 ...... 79. A. helleborifolia mm long; tepals 4–7 mm long, white; 93a (86b). Tepals 15–35 mm long; basal leaf basal leaf blades 4–7  10–12 mm, petioles basally sharply dilated subglabrous; involucral leaves pal- (auriculate) ...... 94 mately dissected or 3-lobed ...... 89 93b. Tepals smaller; basal leaf petioles 89a. Achenes narrowly ovoid, 5–6 mm long; basally vaginate or narrow ...... 97 tepals 5–7 mm long, puberulent 94a. Achenes 1–2 mm long, with styles 1–2 throughout; basal leaf blades 4–5 5– mm long; filaments dilated; involucral 6cmlong; involucral leaves on petioles leaves more than 3, on petioles 1–5 mm 6–7 mm long ...... 74. A. filisecta long, 3-parted or 3–5-lobed ...... 95 89b. Achenes ovoid, 3–4 mm long; tepals 4– 94b. Achenes 4–9 mm long, with styles 2–5 5mmlong, puberulent only along cen- mm long; filaments filiform; involucral tral vein; basal leaf blades 5–7 10–12 leaves 3, sessile, entire ...... 96 mm; involucral leaves sessile ...... 95a. Tepals with narrow bases and wide api- ...... 75. A. orthocarpa ces; scapes few-flowered; basal leaf 90a (87b). Achenes 3–4 mm long, with blades 2- or 3-ternate, sparsely pubes- styles 0.3–0.5 mm long; tepals 5–7, cent; involucral leaves 2-ternate ...... white, with 3–9 anastomosing veins; ...... 80. A. jamesonii involucral leaves 3, on petioles 10–15 95b. Tepals with wide bases and acute api- mm long; plants with caudices (ser. 2. ces; scapes many-flowered; basal leaf Angustilobae) ...... 91 blades palmately ternate, glabrous; 90b. Achenes 5–6 mm long, with styles 1–2 involucral leaves 3–5-lobed ...... mm long; tepals 5 (–6), with 5–9 ...... 81. A. peruviana anastomosing veins; involucral leaves 96a. Carpel and achenes with substraight mostle more than 3, on petioles 1–2 mm styles 4–6 mm long; tepals dimorphic, long; plants with rhizomes (ser. 3. glabrous; basal leaflets 3-parted or 3- Mexicanae) ...... 92 lobed ...... 82. A. sellowii 91a. Achenes with spirally uncinate styles 96b. Carpels and achenes with hooked styles ca. 1 mm long; tepals 5–7, anastomos- 2–3 mm long; tepals monomorphic, ing veins 3–5; basal leaf petioles basally sparsely pubescent; basal leaflets entire 150 植物研究雑誌 第83巻第3号平成20年6月

...... 83. A. assibrasiliana without anastomosing veins; basal leaf 97a (93b). Carpels and achenes not com- blades 4–8 4–7 mm .. 90. A. howellii pressed, basally sparsely pubescent, 102a (83b). Plants with dichotomic non- with substraight styles; tepals 20–35 rosulate stems and stolon-like runners; mm long, anastomosing veins 5–7, carpels and achenes ellipsoid, wings sparsely puberulent along central vein; 0.5–1.5 mm wide; styles substraight, 2– basal leaf blades 15–35 cm long, 3mmlong; tepals 4–5 (subgen. 12. glabrous ...... 84. A. moorei Anemonidium 19. sect. Anemonidium).. 97b. Carpels and achenes compressed, ...... 103 glabrous, with hooked styles; tepals 5– 102b. Semi-rosulate plants with rhizomes; 15 mm long, without anastomosing carpels and achenes oblong-ellipsoid or veins, subglabrous; basal leaf blades cylindroid; without wings; styles mainly 3–8 (–10) cm long, sparsely pubescent curved or bent; tepals 5 or more ... 104 ...... 98 103a. Achenes with styles 1–2 mm long; 98a. Achenes basally shortly stalked, with tepals 4, 7–15 mm long; stems many- styles 3–5 mm long; tepals 7–15 mm flowered; basal leaves scale-like only; long, elliptic; scapes 2–3-flowered; all leaves opposite, with distinct blades basal leaf blades 3-parted or 3-lobed; ...... 91. A. dichotoma plants with short rhizomes ...... 103b. Achenes with styles 3–6 mm long; ...... 85. A. antucensis tepals 5, 10–20 mm long; stems 1–2- 98b. Achenes sessile, with styles 1–2 mm flowered; basal leaves 2–3, scale-like long; tepals 4–6 mm long, linear- and 1–2 leaves with distinct blades; lanceolate; flowers solitary; basal leaf involucral leaves 3, with 3-cleft blades blades ternate or biternate; plants with ...... 92. A. canadensis short nodulose and long stolon-like rhi- 104a. Plants with short vertical rhizomes and zomes ...... 86. A. tenuicaulis monopodial scapes; carpels and achenes 99a (85b). Achenes 2–4 mm long, 1-ribbed; oblong-ellipsoid, with wings 1–3 mm tepals basally pubescent; basal leaf wide (subgen. 9. Omalocarpus) .... 105 blades 8–13 10–13 cm ...... 100 104b. Plants with short ascending or long 99b. Achenes 4–5 mm long, without distinct horizontal rhizomes and sympodial ribs; tepals subglabrous; basal leaf scapes; carpels and achenes cylindroid, blades 4–8 4–7 cm ...... 101 without wings ...... 116 100a. Achenes rhombic-ovoid, sparsely pu- 105a. Basal leaf blades 3-lobed to 3-sected, bescent; tepals white, anastomosing with subequal lobes or segments; basal veins 3–5; staminodes absent ...... and involucral leaves similar; ...... 87. A. begoniifolia inflorescences typically umbelliferous; 100b. Achenes globose-ovoid, glabrous; tepals mainly elliptic, having 1–7 tepals purple, anastomosing veins more anastomosing (sect. 3. Omalocarpus) .... than 10; staminodes between stamens ...... 106 and carpels present ...... 105b. Basal leaf blades pinnatifid, with large ...... 88. A. hokouensis central segment and smaller imbricate 101a. Tepals 10–15 mm long, white, lateral segments; involucral leaves re- anastomosing veins more than 10; basal duced; flowers solitary; teplas linear- leaf blades 3–5 3–5 cm ...... lanceolate, without anastomosing veins ...... 89. A. chapaensis (sect. 14. Imbricata) ...... 101b. Tepals 8–10 mm long, reddish-white, ...... 104. A. imbricata June 2008 Journal of Japanese Botany Vol. 83 No. 3 151

106a. Tepals linear-lanceolate, 6–10 mm ovate ...... 97. A. taipaensis long; basal leaf blades rhombic, 112b. Tepals red or pink, 15–20 mm long, glabrous, 2–3  2–4 cm (ser. 3. with 1–3 anastomosing veins; basal leaf Fuscopurpurea) ...... blades pentagonal or rhombic ...... 103. A. fuscopurpurea ...... 98. A. smithiana 106b. Tepals wider,10–25 mm long; basal 113a (107b). Achenes 8–12 mm long, with leaf blades of various shape, pubescent styles 1.5–2 mm long; tepals 4; basal (rarely subglabrous), much larger ... 107 leaf blades 3-lobed, coriaceous, 107a. Achenes with styles pressed to their glabrous ...... 99. A. tetrasepala bodies; tepals mainly 6–15 mm long, 113b. Achenes and styles smaller; tepals 5– filaments linear; inflorescences mainly 10; basal leaf blades 3-parted, herba- umbelliferous; basal leaf blades 3- ceous, pubescent ...... 114 sected (ser. 1. Involucratae) ...... 108 114a. Achenes orbicular-ovate, sparsely 107b. Achenes with straight or bent styles; puberulent, with curved styles; tepals 5– tepals 10–25 mm long, filaments di- 10, reflexed, with 7–9 anastomosing lated; inflorescences branched; basal veins, sparsely puberulent ...... leaf blades 3-parted or 3-lobed (ser. 2...... 100. A. polyanthes Involucellatae) ...... 113 114b. Achenes ovate-ellipsoid, glabrous, 108a. Tepals 15–20 mm long, glabrous; with substraight styles; tepals 5, scapes 70–120 cm long; basal leaf straight, without anastomosing veins, blades orbicular-ovate, subglabrous ...... subglabrous ...... 115 ...... 94. A. robusta 115a. Achenes 6–8 mm long, with wings 108b. Tepals smaller, pubescent; scapes 1.2–1.3 mm wide and styles 1–1.5 mm lower; basal leaf blades of various long; tepals 12–20 mm long, with shape, pubescent ...... 109 cuneate base and rounded apex ...... 109a. Basal leaf blades orbicular; involucral ...... 101. A. elongata leaves larger than basal leaves; tepals 115b. Achenes 5–6 mm long, with wings 6–10 ...... 95. A. cathayensis 0.5–0.6 mm wide and styles 0.6–1 mm 109b. Basal leaf blades non-orbicular; long; tepals 7–12 mm long, with involucral leaves smaller than basal rounded base and acute apex ...... leaves; tepals 5–7 ...... 110 ...... 102. A. shikokiana 110a. Carpels and achenes ovate to obovate; 116a (104b). Carpels and achenes cylindroid; tepals 5–7, subglabrous ...... 111 tepals 5, without anastomosing veins; 110b. Carpels and achenes broadly ovate to anther connectives with large subglo- suborbicular; tepals 4–5, pubescent ...... bose projections; inflorescences many- ...... 112 flowered; basal leaf petioles basally 111a. Tepals monomorphic, white or yel- sharply dilated; plants with short non- lowish; scapes mainly solitary, erect; tuberous rhizomes (subgen. 13. basal leaf blades wider than long ...... Hepaticifolia sect. 19. Hepaticifolia) ...... 93. A. narcissiflora ...... 116. A. hepaticifolia 111b. Tepals sometimes dimorphic, mainly 116b. Carpels and achenes spindle-like or blue or pink; scapes 3–5, ascending; ellipsoid; tepals 5–8, having ana- basal leaf blades longer than wide ...... stomosing veins; anther connectives ...... 96. A. demissa without projections; flowers solitary; 112a. Tepals white, 7–10 mm long, with 3– basal leaf petioles basally not sharply 5 anastomosing veins; basal leaf blades dilated; plants with long horizontal or 152 植物研究雑誌 第83巻第3号平成20年6月

short tuberous rhizomes ...... 117 References 117a. Carpels and achenes spindle-like, not Adanson M. 1763. Familles des Piantes. 640 pp. Paris. compressed, glabrous; styles 4–5 mm Baumberger H. 1970. Chromosomenzahlbestimmun- long; tepals 5-6, pubescent; plants with gen und Karyotypanalysen be den Gattungen long horizontal rhizomes (subgen. 14. Anemone, Hepatica und Pulsatilla. Ber. Schweiz. Bot. Ges. 80: 17–96. Richardsonia sect. 15. Richardsonia) .... Britton N. L. 1892. The American Species of Genus ...... 117. A. richardsonii Anemone and the Genera which have been referred 117b. Carpels and achenes ellipsoid, com- to it. Ann. N. Y. Acad. Sci. 6: 215–238. pressed, basally sparsely pubescent; Candolle A. P. de. 1817. Regni Vegetabilis Systema styles ca. 1 mm long; tepals 6–7, Naturale. 1. Paris. Candolle A. P. de. 1824. Anemone L. Prodromus glabrous; plants with short tuberous and Systematis Naturalis Regni Vegetabilis 1: 2–66. long stolon-like rhizomes (subgen. 15. Paris. Crassifolia 18. sect. Crassifolia) ...... Chaudhary R. P. 1988. A Synopsis of the genus ...... 118. A. crassifolia Anemone (Ranunculaceae)inNepal and the adja- cent Regions of the Himalaya. Bot. Zhurn. 73: As a result of the above study, we accept 1188–202 (in Russian). Chaudhary R. P. and Trifonova V. L.1988. Fruit mor- a structure of the genus Anemone s. str. dif- phology and comparative anatomical study of fering essentially from those of Ulbrich pericarp and testa within the Nepalese species of (1906), Juzepchuk (1937), Starodubtsev Anemone (Ranunculaceae). Bot. Zhurn. 73: 803– (1989, 1991) and Tamura (1991, 1995). 817 (in Russian). Recently we published the results of the Chupov V. S. 1975. Biosystematic study in the tribe Anemoneae 22 (in Russian). taxonomic revizion of Anemone sensu Cullis C. A. and Schweizer D. 1974. Repetitious DNA proprio (viz., Ziman 1985, Ziman et al. 1997, in some Anemone species. Chromosoma 44: 417– 1998, 2004, 2005, 2006, 2007, Wang et al. 421. 2001), but in this paper we concentrate all Ehrendorfer F. 1995. Evolutionary trends and patterns our proposals and conclusions on taxonomy in the Anemoninae. Pl. Syst. Evol. (Suppl.) 9: 283– of this genus. In our opinion, the genus 293. Ehrendorfer F. and Samuel R. 2000. Comments on Anemone includes 118 species, 20 subspe- S.B. Hoot’s interpretation of Southern Hemisphere cies and 27 varieties belonging to 23 series, relationships in Anemone (Ranunculaceae) based four subsections, 23 sections and 15 subgen- on molecular data [Amer. J. Bot. 87 (6, suppl.): era. In the genus Anemone we propose the 154–155]. Taxon 49: 781–784. new understanding of three subgenera Ehrendorfer F. and Samuel R. 2001. Contributions to a molecular phylogeny and systematics of Anemone (Stolonifera, Keiskea and Kilimandsharica), and related genera (Ranunculaceae-Anemoninae). series Fuscopurpurea, Altaicae and Mexi- Acta Phytotax. Sin. 39: 77–87. canae.Inaddition we published earlier sect. Ehrendorfer F., Ziman S. N., Keener C. S. et al. 2007. Parviflora (Ziman 1985) and Rosulantes Taxonomic revision and phylogeny of Anemone (Ziman et al. 2005), subsection Somalienses sect. Anemone (Ranunculaceae). Bot. J. Linn. Soc. (Ziman 2006) and 12 series (viz., Obtusi- (in press). Franchet M. A. 1886. Plantae Yunnannensis. Bul. Soc. lobae, Trullifolia, Nikoenses, Caucasicae Bot. France 33: 360–382. and others - Ziman et al. 2006–2007). Franchet M. A. 1888. Plantae Davidianae ex Sinarum Imperie. Ranunculaceae. Bul. Soc. Bot. France 35: 1–10. Franchet M. A. 1890. Dela flora du Nord de la Chine. J. Bot. 23: 1–11. Gaudin J. 1828. Anemone L. Flora Helvetica. 3: 490. Gay C. 1845. Historia fisica y politica de Chile. Botanica. Paris. June 2008 Journal of Japanese Botany Vol. 83 No. 3 153

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S. N. ジーマン, E. V. ブラーク, 門田裕一, C. S. キーナー：イチリンソウ属 [狭義] (キンポウゲ 科) の分類 的特徴を述べるとともに, 118種全種の検索表を 痩果, 花, 葉, 地上部, 地下部など約70の形態 提示した. 的形質の解析の結果, キンポウゲ科イチリンソウ (aウクライナ・M. G. コロドニイ植物学研究所, 属 [狭義] に15亜属23節 4 亜節23列118種が認識 b国立科学博物館植物研究部, された. このうち 6 亜属 2 節16列は新しく報告さ cペンシルバニア州立大学) れたものである. 本稿では, 各種内分類群の形態