植物研究雑誌 J. J. Jpn. Bo t. 80:282-302(2005) A Taxonomic Revision of Anemone L. Section Omalocarpus DC. sensu sensu lato (Ranunculaceae): Part 1 Svetlana Svetlana N. ZIMAN a ，Friedrich EHRENDORFER b， Yuichi KADOTA¥ Carl S. KEENER d， Olga N. TSARENKO a， Elena BULAKH a，Bryan E. DUTTON e aN. aN. G. Kholodny Institute of Botany ，National Academy of Sciences ， Tereshchenkivska Tereshchenkivska 2，Kiev ， 01601 UKRAINE; E-mail: E-mail: [email protected] bInstitute bInstitute of Botany of the Vienna University ，Rennweg 14 ，Vienna ， 1030 AUSTRIA; CDepartment CDepartment of Botany ，National Science Museum ，Tokyo Amakubo Amakubo 4-1-1 ，Tsukuba ， 305-0005 JAPAN; dThe dThe Pennsylvania State University ，208 Mueller Laboratory ， University University Park ，Pennsylvania ， 16802 U. S. A.; eDepartment eDepartment of Biology ，Western Oregon State University ， Monmouth ，Oregon ，97361 U. S. A. (Received (Received on March 15 ，2005) The taxonomy of Anemone section Omalocarpus (Ranunculaceae) ， with members distributed distributed throughout the Northem Hemisphere and especially in East Asia ， was re- evaluated evaluated on the basis of a critical mo 中hological analysis of extensive herbarium mate- rials rials and living populations. A conspectus ，key ，mo 中hometric tables ， and figures illustrating illustrating taxonomic characters 訂 e presented. Three series and 11 species ，in c1 uding eight eight subspecies and 12 varieties of A. narcissiflora and four varieties of A. demissa 紅 e recognized recognized and discussed with respect to their relationships. This paper will be divided into into two parts. In Part I the results of the analysis the A. narcissiflora complex are re 田 ported. ported. Key words: Anemone narcissiflora ，Anemone sec t. Omalocarpus D C.， comparative mor- phology ， taxonomy. Because of its considerable diversity of the genus Anemone which ， for the A. species ，Anemone L. is one of the most diffi- narcissiflora complex ， included 10 species ， cult cult genera in the Ranunculaceae. The A. two subspecies and six varieties. However ， narcissiflora narcissiflora complex 陀 presented by A. this 佐eatment was too brief and did not in- narcissiflora narcissiflora and related taxa is taxonomi- clude data for the whole sec t. Omalocarpus. cally cally most complicated within Ane- Consequently in this paper we present a criti- mone. The complex is widely disjunctive and cal morphological analysis of extensive her- occur occur within the Northen Hemisphere while barium material of all the foregoing taxa other other species occur mainly in East Asia. with respect to their taxonomy ，distribution In In the previous papers (Ziman et al. 1994 ， and relationships. 1995 ，1997 ，etc.) we examined a part of the A. A. narcissiflora complex ， mainly in limits of Literature review the the former Soviet Union. Recently ， in the Anemone narcissiflora L. (error narcissi- “ Flora of China ，"羽 Tang ，Ziman and Dutton folia) was described by Linnaeus in 1753 (2001) (2001) published a taxonomic treatment of together with two allied species ， A. sibirica October October 2005 Journal of Japanese Botany Vo l. 80 No. 5 283 L. L. and A. fasciculata L. ， from Eurasia. scence distinction. He included A. narcissi- Willdnow (1797) later described A. flora and A. demissa in the first series ， and A. umbellata ， and Adams (1 805) described A. polyanthes ， A. elongata and A. tetrasepal αin speciosa speciosa (both from the Caucasus). the second one. De Candolle (1 817 ， 1824) recognized all In a comprehensive herbarium-based three three species within the A. narcissiflora study ， Juzepchuk (1 937) re-examined A. complex complex which he classified in sec t. narcissiflora and allied taxa collected within Omalocarpus Omalocarpus DC. on the basis of their com- the former Soviet Union. The A. narcissi- pressed ， suborbicular ， glabrous achenes ， flora complex was classified in se r. scapose scapose stems ， and umbelliferous inflore- Narcissiflorae Juz. (nomen nudum). scences. scences. Juzepchuk recognized A. sibirica ， A. During During the nineteenth century several spe- fasciculata and A. specios 仏 and he described cies cies allied to A. narcissiflora were described ten new species (A. shrenkiana Juz. ， A. [A. [A. ch ηisantha C. A. Mey. from the biarmiensis Juz. and others) in sec t. Caucasus ， A. αconitifolia Turcz. from Omalocarpus (= subgen. Homaloc 紅 pus Siberia ， and A. elongata D. Don ， A. DC.). Consequently Juzepchuk believed A. polyanthes polyanthes D. Don ， A. tetrasepala Royle and narcissiflora s. st r. did not occur within the A. A. demissa Hoo k. f. & Thoms. from the territory of the former Soviet Union ， but was Himalayas Himalayas (East Asia)]. replaced there with a series of vicariant The first sketch of sec t. Omalocarpus was races. prepared prepared by Hooker (1 872) ， and he included Within the flora of Europe only A. in in it five species (A. narcissiflora ， A. narcissiflora was recognized (Zapalowicz elongata ， A. demissa ， A. polyanthes and A. 1908 ，Hayek 1927 ， Pawlowski 1956 ， Chater tetrasepala) ，despite the fact that the author 1973 ，Pignatti 1988 ，etc.). of of the first monograph on Anemone ，Pritzel According to Sharma et al. (1 993) ， the (1841) ， regarded sec t. Omalocarpus (= flora of India included four species of sec t. Homalocarpus) Homalocarpus) as consisting of only A. Omalocarpus (A. demissa ， A. elongata ， A. narcissiflora narcissiflora and A. polyanthes. Pritzel had polyanthes and A. tetrasepala). However ， A. included included A. elongata and A. tetrasepala in narcissiflora was regarded as absent there. sec t. Anemonospermos ， and A. sibirica in Hara and Williams (1979) recognized A. sec t. Anemonanthea. demissa ， A. elongata and A. polyanthes in Early Early in the twentieth century ， Finet and the flora of Nepal ， but they also included in Gagnepain Gagnepain (1904) viewed A. narcissiflora in this flora two new species ，A. β/'scopurpurea a broad sense and regarded A. demissa and H. Hara (1973) and A. smithiana Lauener & A. A. polyanthes as its varieties. They did not Panigrahi (1975) ，a conclusion also reached accept accept de Candolle' s circumscription of by Chaudkhary (1988). Anemone and included A. elongata and A. In the flora of Pakistan ， three endemic tetrasepala tetrasepala in the so-called seventh section ， species were described recently by Qureshi their their “eighth section" consisting of A. and Chaudhri [(1 988): A. mirajabii R. A. dichotoma ， A. gelida ， A. narcissiflora ， A. Qureshi & M. N. Chaudhri ， A. lowariensis R. rivularis rivularis and A. begoniifolia. A. Qureshi & M. N. Chaudhri and A. In In the next monograph on Anemone ， abdurrehmanii R. A. Qureshi & M. N. Ulbrich Ulbrich (1 905 ， 1906) recognized two series Chaudhri]. All three species are very close to within within sec t. Omalocarpus (= Homalocarpus) ， A. polyanthes. However ， Riedl and Nasir viz. ， ser. Involucratae Ulbr. and ser. (1 991) ，who treated Anemone for the “ Flora Involucellatae Involucellatae Ulb r.， on the basis of inflore- of Pakistan ，" did not accept these species ， 284 植物研究雑誌第80 巻第5号 平成17 年 10 月 and and in their opinion ， only A. polyanthes and in the “ Flora of China." A. A. tetrasepala from sec t. Omalocarpus occur In treating Anemone for the flora of Japan ， in in Pakistan. Ohwi (1 953) in cI uded both A. nαrcissiflora In In the flora of Iran and adjacent territories ， and A. sikokiana Makino in sec t. Rechinger Rechinger and Riedl (1 992) recognized A. Omalocarpus. protracta protracta (instead of A. narcissiflora) ， A. In the flora of North America ， Britton poly α nthes and A. tetrasepal α， but these (1891) recognized A. narcissiflora to be dis- authors authors believed that A. low αriensis and A. tributed disjunctively in Alaska and the abdurrehmanii abdurrehmanii also occur there. They in- Rocky Mountains. Shortly thereafter ，plants cluded cluded all these species in sect. Anemonan- from the Rocky Mountains were described as thea thea (not Omalocarpus). A. zephyra A. Nelson (Nelson 1906). Only In In the flora of China ，Wang (1 974 ， 1980) Rydberg (1 917) accepted this species. Later included included eight species in sec t. Omalocarpus authors (e. g. ， Calder and Taylor 1968 ， (= (= Homalocarpus) [A. crinita ， A. schrenkian α Weber 1976 ，Dorn 1977 ， Scott 1994 ，etc.) (instead (instead of A. narciss 伊ora) ， A. demissa ， A. did not recognize A. zephyr αas a distinct imbricata ，and four endemics (A. cathayensis specles. Kitag. ， A. schantungensis Hand.-Mazz. ， A. The A. narcissiflora complex was already robusta robusta W. T. Wang and A. taipaiensis W. described earlier as the genus Homalocarpus T. T. Wang)]. Quite recently Wang described Schur (Schur 1866) ， but this segrega- generic three three na 町 ow endemics from China (A. tion was not supported by later authors. tengchongensis tengchongensis W. T. Wang ， A. multi- Recently Holub (1 973) again proposed to lobulata lobulata W. T. Wang & L. Q. Li and A. sep 訂 ate sec t. Omalocarpus from Anemone as nutantiflora nutantiflora W. T. Wang & L. Q. Li (Wang a separate genus (Omalocarpus). and and Li 1986 ，Wang 1996). According to the In his summaηof additions to the flora of descriptions ，all these plants are veηclose to the former Soviet Union ， Cherepanov (1 981 ， narcissiflora. A. narcissiflora. 1996) followed Holub (1 973) ， and he ac- Recently ， in the modern version of the cepted almost all the taxa of A. narcissiflora “ Flora of China" (2001) ，Wang ，Ziman and sensu Juzepchu k. In his treatment of Dutton Dutton (2001) published a taxonomic treat- Anemone ， Starodubtsev (1 989 ， 1991) also