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Modern View on the Taxonomy of the Genus Anemone L. Sensu Stricto (Ranunculaceae)

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Modern View on the Taxonomy of the Genus Anemone L. Sensu Stricto (Ranunculaceae) 植物研究雑誌 Originals J. Jpn. Bot. 83: 127–155 (2008) Modern View on the Taxonomy of the Genus Anemone L. Sensu Stricto (Ranunculaceae) 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 species. 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, flowers, leaves, 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 Hepatica 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 Anemonoides 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 South America 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
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