REVISION OF ANEMONE SECT. HIMALAYICAE (RANUNCULACEAE) WITH THREE NEW SERIES

Sergei Mosyakin

The present paper documents the genus Anemone L. (Ranunculaceae) from India. A total of 26 taxa (which includes 24 species, one subspecies and one variety) has been recorded from Indo Himalayas. Western Himalaya is endowed with 9 taxa, Eastern Himalaya with 8 taxa, while 9 are common to both the flanks. Three species are present in Northeast India which are common to IHR and one species has also been reported from Western Ghats. This genus constitutes flowering plants of high altitude and maximum diversity has been observed between 2000–3000 m asl. The genus possesses high medicinal potential and needs urgent assessment of taxonomic and conservation status for its sustainable utilization.

This report comprises the first list of 1019 specimens of vascular plants (Pteridophytes: 25 families and 106 species; Gymnosperms: 9 families and 19 species; Angiosperms: 136 families and 894 species) reposited in the “Tribhuvan University Herbarium (TUH)”, Department of Botany, Post Graduate Campus, Tribhuvan University, Biratnagar, Nepal. These specimens have been collected by students and teachers of the Department of Botany, Post Graduate Campus from various locations of eastern Nepal (tropical to temperate climates; 60 to 3000 m, msl altitudes) since the year 1992 onwards. For the purpose of easy access to the specimens, families within a taxon, genera within a family, and species within a genus are arranged alphabetically. Scientific names provided by Hara et al. (1978, 1979, 1982), Iwatsuki (1988), Koba et al. (1994), Siwakoti (1995), Press et al. (2000), Jha and Jha (2000), and Thapa (2002) have been adopted for the nomenclature of the reposited specimens. DOI: http://dx.do...

During the monitoring of populations of Anemone sylvestris L. (Ranunculaceae), a protected species in Poland, we found that the seed set is impaired. The flower is considered an adaptation that has coevolved to achieve effective pollination and successful fertilization. Therefore we have focused on the morphological and anatomical characteristics of the flowers of A. sylvestris L. as a prelude to the study of the species’ pollination biology and plant breeding system. The large size of the flower (50.6 ± 16.4 mm in dimensions) and its bowl shape fulfil both the biotic pollination syndrome and the aerodynamic requirements for pollen dispersal and capture. The opening and closing of the perianth provide a shelter for beetles. The odourless perianth, absence of nectar, scarcity of pollen (approximately 200 000 pollen grains per flower) and its traits - small size (axis P = 18.52 ± 1.0 μm; E = 16.59 ± 0.9 μm), lack of balsam on the exine surface, starch accumulation in more than 95% of ...

E D I N B U R G H J O U R N A L O F B O T A N Y 64 (1): 51–99 (2007) E 51 Trustees of the Royal Botanic Garden Edinburgh (2007) doi:10.1017/S0960428607000765 REVISION OF ANEMONE SECT. HIMALAYICAE (RANUNCULACEAE) WITH THREE NEW SERIES S . N . Z I M A N 1, F . E H R E N D O R F E R 2, C . S . K E E N E R 3, W . T . W A N G 4, S . L . M O S Y A K I N 1, E . V . B U L A K H 1, O . N . T S A R E N K O 1, B . E . D U T T O N 5, R . P . C HAUDHARY6 & Y . K ADOTA7 The members of Anemone L. sect. Himalayicae (Ulbr.) Juz. (Ranunculaceae) are mainly distributed in the Himalaya of North India, Nepal and Bhutan and the neighbouring mountains of SW China at elevations between 1850 and 4800 m. Their taxonomy is re-evaluated on the basis of a critical morphological analysis of extensive herbarium material. The section is placed in Anemone subgen. Omalocarpus and differentiated into three new series: ser. Obtusilobae, ser. Trullifoliae and ser. Rupestres. A conspectus, keys to species, subspecies and varieties, descriptions of taxa, illustrations and distribution maps are presented. Eleven species with several infraspecific taxa are recognized and their synonymy, variability and relationships are discussed. In addition to the generally accepted species Anemone obtusiloba, A. trullifolia and A. rupestris, we recognize the following: A. polycarpa, A. rockii, A. geum and A. coelestina and four Chinese endemics, A. yulongshanica, A. patula, A. subpinnata and A. subindivisa. Anemone imbricata and A. fuscopurpurea are described but excluded from the section. The origins, morphological differentiations and eco-geographical radiations of Anemone sect. Himalayicae are discussed. Keywords. Anemone subgen. Omalocarpus sect. Himalayicae, A. obtusiloba complex, new series, phytogeography, Ranunculaceae, Sino-Himalayan region, taxonomy. I NTRODUCTION Anemone L. sect. Himalayicae (Ulbr.) Juz., also called the Anemone obtusiloba D.Don complex, is one of the most difficult taxonomic groups in the genus Anemone (Ranunculaceae Juss.), due to the uncertain number of taxa and their problematic relationships. There have been three serious attempts to revise this complex: by Brühl (1896), Lauener (1960) and Wang et al. (2001). However, Brühl used data from only about 10 collections and although Lauener examined more collections, he 1 N. G. Kholodny Institute of Botany, National Academy of Sciences, Tereshchenkivska str. 2, Kiev, 01601, Ukraine. 2 Institute of Botany, University of Vienna, Rennweg 14, Vienna A-1030, Austria. 3 Pennsylvania State University, 208 Mueller Laboratory, University Park, PA 16802, USA. 4 Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 5 Department of Biology, Western Oregon University, 345 North Monmouth Avenue, Monmouth, OR 97361, USA. 6 Central Department of Botany, Tribhuvan University, Kirtipur, Kathmandu 5927, Nepal. 7 Department of Botany, National Science Museum, 4-1-1 Amakubo, Tsukuba, Ibaraki 305, Japan. 52 S. N. ZIMAN ET AL. dealt only with a limited number of morphological characters. More recently, Wang et al. in the Flora of China provided a treatment of the genus Anemone which, for the A. obtusiloba complex, included 11 species, seven subspecies and 13 varieties. In this paper we amplify this treatment by Wang et al. on the basis of a critical morphological analysis of extensive herbarium material, a thorough literature survey and a discussion of the differentiation and relationships of the taxa involved. We recognize three new series, 11 species, five subspecies and 14 varieties. M ATERIALS AND M ETHODS Our study is based mainly on c.2000 herbarium specimens received on loan from the following herbaria: BBG, BC, BCC, BM, BRA, E, GH, K, KATH, KRA, KRAM, KW, LE, P, PRG, US, VAB, W and WU (Holmgren et al., 1990), including 240 specimens collected in the high-mountain areas of India, Nepal and China by the early collectors Delavay, Forrest, Handel-Mazzetti, J. D. Hooker, Kingdon-Ward, Ludlow & Sheriff, Polunin, Potanin, Rock, Stainton et al., Thomson and T. T. Yu, together with 60 specimens used by Lauener (1960). In addition, we have examined the types of most species, subspecies and varieties in the section. From these specimens about 300 flowering and fruiting samples were studied in detail, using light and scanning electron microscopy, in order to obtain comparative data on basal and involucral leaves, flowers and achenes. H ISTORICAL S URVEY Anemone obtusiloba D.Don was described (Don, 1825) from the Nepal Himalaya, and shortly thereafter other authors published A. trullifolia Hook.f. & Thoms., A. rupestris Wall. ex Hook.f. & Thoms. and A. falconeri Thoms. (Hooker & Thomson, 1855). Later Anemone coelestina Franch. (Franchet, 1885), A. imbricata Maxim. (Maximowicz, 1889) and A. gelida Maxim. (Maximowicz, 1890) were described from the Himalaya and adjacent territories of China as taxa allied or closely related to A. obtusiloba. Within the Himalayan members of the group Brühl recognized only two species, Anemone obtusiloba and A. rupestris. He reduced the other taxa to subspecies and regarded the recognition of Anemone falconeri as uncertain. Shortly thereafter Finet & Gagnepain (1904) recognized four species within this complex (Anemone obtusiloba, A. trullifolia, A. glauciifolia and A. reflexa). In his extensive revision of Anemone Ulbrich (1906) recognized five species within the A. obtusiloba group (A. obtusiloba, A. rupestris, A. trullifolia, A. coelestina and A. imbricata). After Ulbrich’s revision a number of new species allied to Anemone obtusiloba were described: A. bonatiana H.Lév. (1909), A. geum H.Lév. (1915), A. polycarpa W.E.Evans (1921), A. rockii Ulbr. (1929), and A. ovalifolia (Brühl) Hand.-Mazz. (1931). Overlapping and variable characters have complicated the taxonomy of the Anemone obtusiloba complex. Pritzel (1841), who produced the first monograph on REVISION OF ANEMONE SECT. HIMALAYICAE 53 Anemone, classified A. obtusiloba in section Anemonospermos DC. (1824). Maximowicz (1890) placed Anemone imbricata into section Anemonanthea DC. and A. gelida into section Omalocarpus DC. After an examination of the achene morphology in Anemone, Janczewski (1892) classified A. obtusiloba in section Pulsatilloides DC., together with A. glauciifolia Franch. from China, and A. capensis L. and A. alchemillifolia E.Mey. from South Africa. Brühl (1896) regarded such a taxonomic association as unjustified and suggested a relationship between the groups of Anemone obtusiloba and A. narcissiflora. Moreover, Finet & Gagnepain (1904) included Anemone gelida and A. imbricata in the same section as A. narcissiflora. Like Janczewski (1892), Ulbrich (1906) included the species of the Anemone obtusiloba complex in section Pulsatilloides, but separated them as series Himalayicae Ulbr. After examining c.160 specimens, including 30 types, from 120 localities, Lauener (1960) noted almost continuous variation in many characters, especially of ‘sepals’ (often termed tepals in the literature, but referred to as ‘petaloids’ in this paper), pistil and involucral leaf characters, as well as indumentum. Lauener believed that four species within the Anemone obtusiloba complex (A. obtusiloba, A. rupestris, A. trullifolia and A. imbricata) could be distinguished on the basis of the shape, dissection and teeth characters of the basal leaves. Tamura (1967) took Brühl’s opinion into consideration and consequently removed Anemone obtusiloba and allied species from section Pulsatilloides, and re-classified them in section Omalocarpus as subsection Himalayicae (Ulbr.) Tamura. Tamura (1991, 1995) also followed Juzepchuk (1937) in raising the rank of section Omalocarpus to subgenus Omalocarpus (DC.) Juz. and of subsection Himalayicae to section Himalayicae (Ulbr.) Juz. In these publications, Tamura erroneously called the section ‘Himalayica’, despite the original spelling ‘Himalayicae’ by Ulbrich (1906), Juzepchuk (1937), and even by Tamura himself in his earlier publication (Tamura, 1967). According to Wang (1974, 1980), a total of 10 species of section Himalayicae occur in China: the previously mentioned Anemone obtusiloba, A. rupestris, A. trullifolia and A. rockii, and six Chinese endemics: A. patula C.C.Chang & W.T.Wang, A. yulongshanica W.T.Wang, A. subpinnata W.T.Wang, A. liangshanica W.T.Wang, A. lutienensis W.T.Wang and A. subindivisa W.T.Wang. However, Wang placed Anemone imbricata in section Omalocarpus. In North India and Nepal four species of section Himalayicae were recognized: Anemone obtusiloba, A. rupestris, A. trullifolia and A. geum, with A. falconeri transferred to Hepatica (Hara & Williams, 1979; Chaudhary, 1988; Sharma et al., 1993). In addition, Hara (1973) described Anemone fuscopurpurea as a taxon allied to A. obtusiloba which was later placed into the monotypic section Fuscopurpurea Tarasevich & Chaudhary (1987). According to Qureshi & Chaudhri (1978), only Anemone obtusiloba and two closely related species, A. multisepala Qureshi & Chaudhri and A. neelamiana Qureshi & Chaudhri, occur in Pakistan. They place these species into section Anemonanthea. In 54 S. N. ZIMAN ET AL. contrast, Rechinger & Riedl (1992) maintain that Anemone obtusiloba belongs to section Omalocarpus and that it occurs in both Pakistan and Afghanistan. In revising the subtribe Anemoninae Spach, Starodubtsev (1989, 1991) suggested raising section Pulsatilloides sensu Ulbrich to the status of genus, Pulsatilloides (DC.) Starod., with several East Asian and African species. He included in this genus the section Himalayicae (Ulbr.) Starod. with the following three species: Pulsatilloides obtusiloba (D.Don) Starod., P. trullifolia (Hook.f. & Thoms.) Starod., and P. gelida (Maxim.) Starod. However, he transferred Anemone imbricata to section Imbricatae Starod. in the genus Anemonastrum Holub. On the basis of morphological and molecular analyses Hoot et al. (1994) verified the correct placement of the Anemone obtusiloba complex in section Omalocarpus (‘Homalocarpus’) with five species: A. obtusiloba, A. rupestris, A. trullifolia, A. coelestina and A. geum. The treatment of the Anemone obtusiloba complex in the Flora of China by Wang et al. (2001) differed from Wang (1974, 1980) in that A. liangshanica and A. lutienensis were reduced to varieties of A. trullifolia, whereas A. coelestina, A. geum and A. polycarpa were recognized as species. P LACEMENT AND D EFINITION OF A NEMONE SECT. H IMALAYICAE In our treatment we mainly follow Hoot et al. (1994) and Tamura (1995) in accepting the placement of the Anemone obtusiloba complex as section Himalayicae (not ‘Himalayica’) within the subgenus Omalocarpus. Anemone subgen. Omalocarpus has two sections: Omalocarpus and Himalayicae. Members of both sections share numerous and mostly synapomorphic morphological characters: short, erect or ascendent monopodial rhizomes with several axillary flowering stems (scapes), rosulate basal leaves with vaginate petioles, sessile involucral leaves, umbellate cymose inflorescences, a perianth of 5–8 petaloids, sessile achenes in globose or ovoid heads and the synplesiomorphic feature of tricolpate pollen grains. Furthermore, both sections evidently have the same and apomorphic chromosome base number of x 5 7 (see Discussion). In agreement with Tamura (1967, 1995) we regard the taxa in section Himalayicae as characterized by having reduced inflorescences with only 1(2–3) flowers. In our opinion, other diagnostic characters of this section include the three reduced sessile involucral leaves (bracts) which are always distinctly smaller than the basal leaves, and petaloids with 3–5(–7) basal veins, usually lacking anastomoses. D IFFERENTIAL C HARACTERS WITHIN THE S ECTION For the taxonomic differentiation of the section, we first examined c.80 morphological characters and then narrowed down the list to 16 quantitative and 19 qualitative features. Several quantitative characters, such as the range of basal leaf number REVISION OF ANEMONE SECT. HIMALAYICAE 55 (mostly 5–12) or the average number of basal leaves (7–8), were too similar across the taxa to be useful. They depend largely on the plant’s age and ecological conditions. The same applies to the number of scapes and petaloids, the length and width of petaloids, and the length of involucral bracts, scapes, pistils and stamens. We consider the most important quantitative characters to be the length and width of petioles and blades of basal leaves. These vary from blades orbicular-ovate (wider than long) with long narrow petioles to blades oblong (longer than wide) with short, broad, or almost absent petioles. There are great differences in the degree of leaf division from twice 3-sect or 3-lobed with leaflets on petiolules or fused to completely undivided leaves. Great variability is also observed with respect to the indumentum of the basal leaves, involucral bracts, scapes and pedicels, from villous and densely pubescent to practically glabrous. Flowers differ in many qualitative and quantitative characters, for example in the shape of petaloids which may be obovate, elliptic or oblong-elliptic. In addition, petaloid colour varies considerably within most taxa and several colour variants frequently occur in the same population. For example, flowers of whitish-blue, orange-brown, deep blue-violet, dark red, and white with violet tinges are found in single populations of Anemone obtusiloba in the Himalaya. Further differences between taxa exist with respect to petaloids in one or two whorls (and then often dimorphic), in their veins and vein anastomoses, in the presence of staminodes and in the stamens and their filaments. The quantitative characters of achene and style length are also important. Styles may be conical, apically straight or slightly curved and rarely even uncinate. To summarize, the most important characters for taxonomic differentiation within Anemone sect. Himalayicae are: N N N N N N N N N N N base of leaf blades (cordate or rounded to truncate, attenuate or cuneate) division of leaf blades (from twice 3-sect to undivided) leaflets (with petiolules or sessile) inflorescences (several- or solitary-flowered) shape of inflorescence bracts (lobed, with teeth or undivided) petaloids (monomorphic or dimorphic) anastomosing petaloid veins (present or absent) shape of filaments (linear or dilated, i.e. lanceolate) staminodes (present or absent) achenes (ovoid or sometimes¡compressed, without or with distinct lateral ribs) indumentum of leaves, stems, pedicels, petaloids, pistils and achenes. C ONSPECTUS, K EY AND R EVISION OF A NEMONE SECT. H IMALAYICAE The following conspectus differs from Starodubtsev (1991) and Tamura (1995) in the circumscription of many taxa and some nomenclatural details. We recognize 11 species (with five subspecies and 14 varieties) in section Himalayicae, which occur mainly in the Himalaya of Nepal and North India and in adjacent areas of China. 56 S. N. ZIMAN ET AL. They are classified into three new series: Obtusilobae, Trullifoliae and Rupestres. For the three narrow endemics A. patula, A. subpinnata and A. subindivisa from China (Sichuan) no ripe achenes are yet available. Their placement into the series is still provisional. Two other species (Anemone imbricata and A. fuscopurpurea) are briefly described and discussed, but are excluded from section Himalayicae and the Key. Anemone L., Sp. Pl. 538 (1753); Gen. Pl. 241 (1754). – Type: Anemone coronaria L. Subgenus Omalocarpus (DC.) Juz., Fl. URSS 7: 256 (1937). – Type: Anemone narcissiflora L. Section Himalayicae (Ulbr.) Juz., Fl. URSS 7: 256 (1937). – Anemone subsect. Brevistylae Ulbr. ser. Himalayicae Ulbr., Bot. Jahrb. Syst. 37: 201 (1906). – Anemone subsect. Himalayicae (Ulbr.) Tamura, Sci. Rep. (Osaka Univ.) 16: 27 (1967). – Pulsatilloides (DC.) Starod. sect. Himalayicae (Ulbr.) Starod., Vetrenitsy 124 (1991). – Type: Anemone obtusiloba D.Don. Series Obtusilobae Ziman, Ehrendorfer & Bulakh, ser. nov. Achenia ovoidea, non compressa, sine costis lateralibus, plus minusve dense pilosa, styli paulo flexi, 1–2.5 mm longi. Foliorum radicalium petiola 1–2(3) mm lata, laminae in parte basali cordatae sive rotundatae, plerumque distincte trisectae foliolis apicalibus petiolulatis, tripartitis, trilobatis sive integerrimis. – Type: Anemone obtusiloba D.Don. Achenes ovoid, not compressed, without lateral ribs, more or less densely pubescent, styles slightly bent, 1–2.5 mm long. Basal leaves with petioles 1–2(3) mm wide, blades cordate or rounded at base, mostly 3-sect with apical leaflets petiolulate, 3-parted to 3-lobed or undivided. 1. Anemone obtusiloba D.Don A. subsp. obtusiloba a. var. obtusiloba b. var. potentilloides Lauener c. var. leiophylla (W.T.Wang) Ziman, Ehrendorfer & Bulakh B. subsp. megaphylla W.T.Wang C. subsp. nepalensis Chaudhary 2. Anemone patula C.C.Chang & W.T.Wang 3. Anemone polycarpa W.E.Evans 4. Anemone subpinnata W.T.Wang 5. Anemone rockii Ulbr. a. var. rockii b. var. pilocarpa W.T.Wang c. var. multicaulis W.T.Wang 6. Anemone geum H.Lév. REVISION OF ANEMONE SECT. HIMALAYICAE 57 Series Trullifoliae Ziman, Ehrendorfer & Bulakh, ser. nov. Achenia ovoidea, non compressa, sine costis lateralibus, plus minusve dense pilosa, styli erecti, 2–2.5 mm longi. Foliorum radicalium petiola (2–3)4–10 mm lata, laminae in parte basali attenuatae, cuneatae sive truncatae, plus minusve tripartitae foliolis sessilibus nec petiolulatis vel indivisae. – Type: Anemone trullifolia Hook.f. & Thoms. Achenes ovoid, not compressed, without lateral ribs, more or less densely pubescent, with straight styles 2–2.5 mm long. Basal leaves with petioles (2–3)4–10 mm wide, blades attenuate, cuneate or truncate, more or less 3-divided or undivided, leaflets never petiolulate. 7. Anemone trullifolia Hook.f. & Thoms. a. var. trullifolia b. var. liangshanica (W.T.Wang) Ziman & B.E.Dutton c. var. lutienensis (W.T.Wang) Ziman & B.E.Dutton 8. Anemone coelestina Franch. a. var. coelestina b. var. linearis (Brühl ex Hand.-Mazz.) Ziman & B.E.Dutton c. var. holophylla (Diels) Ziman & B.E.Dutton 9. Anemone subindivisa W.T.Wang 10. Anemone yulongshanica W.T.Wang a. var. yulongshanica b. var. truncata (H.F.Comber) W.T.Wang Series Rupestres Ziman, Ehrendorfer & Bulakh, ser. nov. Achenia ellipsoidea, distincte compressa, costis lateralibus distinctis, plerumque glabra, styli curvati, 1.2–1.8 mm longi. Foliorum radicalium petiola 1–2 mm lata, laminae in parte basali subcordatae, bis trisectae, foliolis petiolulatis, foliolum apicale trisectum. – Type: Anemone rupestris Wall. ex Hook.f. & Thoms. Achenes ellipsoid, distinctly compressed, with lateral ribs, mostly glabrous, styles curved, 1.2–1.8 mm long. Basal leaves with petioles 1–2 mm wide, blades basally subcordate, twice 3-sect, leaflets petiolulate, apical leaflet 3-sect. 11. Anemone rupestris Wall. ex Hook.f. & Thoms. A. subsp. rupestris B. subsp. gelida (Maxim.) Lauener Species of Anemone subgen. Omalocarpus excluded from section Himalayicae: 12. Anemone imbricata Maxim. 13. Anemone fuscopurpurea H.Hara 58 S. N. ZIMAN ET AL. Key to species 1a. Achenes ellipsoid, distinctly compressed, with lateral ribs, mostly (sub)glabrous; basal leaves with blades twice and deeply 3-sect with 3-sect apical leaflets ___ ______________________________________________________ 11. A. rupestris 1b. Achenes ovoid, not (or sometimes slightly) compressed, without lateral ribs, more or less densely pubescent; basal leaves with blades less deeply and often only once 3-sect to 3-cleft with 3-parted to 3-lobed or undivided apical leaflets, but leaves sometimes also completely undivided ________________________ 2 2a. Basal leaves mostly deeply divided and at least the terminal leaflets with distinct petiolules, blades basally cordate or rounded, often wider than long or as wide as long; indumentum variable to reduced; leaf petioles only 1–2(–3) mm wide _____________________________________________________________ 3 2b. Basal leaves more shallowly divided and the lobes without distinct petiolules, or completely undivided, blades basally mostly attenuate, cuneate or truncate, often longer than wide; indumentum villous to densely pubescent; leaf petioles mostly broader and (2–)4–10 mm wide ________________________________ 8 3a. Involucral bracts mostly 3-parted to 3-dentate; scapes 1- or 2(–3)-flowered _ 4 3b. Involucral bracts undivided or partly 3-lobed; flowers solitary ____________ 6 4a. Bases of basal leaves rounded, apical leaflets with petiolules 10–15 mm long; staminodes present; achene styles uncinate ________________ 3. A. polycarpa 4b. Bases of basal leaves cordate (rarely subtruncate), apical leaflets with shorter petiolules; staminodes absent; achene styles straight or slightly curved _____ 5 5a. Apical leaflets on petiolules 1–2 mm long, lateral leaflets subsessile; scapes 1– 3-flowered; filaments linear _____________________________ 1. A. obtusiloba 5b. Apical leaflets with petiolule only 0.5–1 mm long; flowers solitary; filaments lanceolate ________________________________________________ 2. A. patula 6a. Basal leaves longer than wide, apparently pinnatifid, apical leaflets 3-sect, on 3– 5 mm long petiolules, lateral leaflets subsessile, unequally 3-parted; petioles of basal leaves 2–5 cm long _______________________________ 4. A. subpinnata 6b. Basal leaves about as long as wide, mostly 3-sect to 3-cleft, apical leaflets 3lobed or undivided; petioles of basal leaves 5–20 cm long _______________ 7 7a. Apical leaflets with petiolules 1–2 mm long; petaloids dimorphic, 10–20 6 6– 12 mm; filaments sublinear (0.3–0.5 mm wide) ________________ 5. A. rockii 7b. Apical leaflets with petiolules 5–10 mm long; petaloids all similar, 5–12 6 4– 9 mm; filaments lanceolate (0.5–1 mm wide) __________________ 6. A. geum 8a. Blades of basal leaves with attenuate base; scapes 1–3-flowered; petaloids sometimes with anastomosing veins __________________________________ 9 8b. Blades of basal leaves with truncate base; flowers solitary; petaloids without anastomosing veins _______________________________________________ 10 REVISION OF ANEMONE SECT. HIMALAYICAE 59 9a. Blades of basal leaves 3-parted to 3-lobed, rhombic-ovate to obovate; involucral bracts 3-lobed or 3-dentate; scapes 1–3-flowered; involucral bracts mostly 3-lobed or 3-dentate; filaments ovate-lanceolate _____ 7. A. trullifolia 9b. Blades of basal leaves weakly 3-lobed to undivided, oblong-linear to oblanceolate; involucral bracts undivided; flowers solitary; filaments linear ______________________________________________ 8. A. coelestina 10a. Blades of basal leaves undivided or obscurely 3-lobed; involucral bracts entire or 3-dentate; petaloids 5–10 6 5–8 mm, with 3–5 veins ___ 9. A. subindivisa 10b. Blades of basal leaves mostly 3-parted or 3-lobed; involucral bracts 3-lobed; petaloids 8–16 6 8–10 mm, with 3 veins ____________ 10. A. yulongshanica 1. Anemone obtusiloba D.Don, Prodr. Fl. Nepal 194 (1825). – Anemone obtusiloba D.Don subsp. obtusiloba (typica) Brühl, Ann. Roy. Bot. Gard. Calcutta 5: 78 (1896). – Anemone obtusiloba D.Don subsp. genuina Ulbr., Bot. Jahrb. Syst. 37: 242 (1906). – Type: Hab. In Gosaingsthan Nepaliae, Wallich s.n. (lecto BM, designated here by Ziman and Mosyakin). Figs 1, 18. Anemone govaniana Wall., Numer. List 166: 4688 (1831), nom. nud., non Lindl. (1844). – The specimen of A. obtusiloba annotated as A. govaniana India, 10–11000 ft, Herb. Falconer 29 is deposited at K in a type folder. Anemone mollis Wall., Numer. List 166: 4689 (1831), nom. nud. Anemone discolor Royle, Ill. Bot. Him. Mount. 11: 52 (1839). – Type: NW India, Himalayae Alpibus. Choor-Urukta et Kendarkanta supra 10000 pedes elevatis, Royle (lecto LIV). Anemone micrantha Klotzsch, Bot. Ergebn. Reise Waldemar 133 (1862). – Anemone obtusiloba D.Don subsp. micrantha (Klotzsch) Ulbr., Bot. Jahrb. Syst. 37: 242 F I G . 1. Anemone obtusiloba D.Don subsp. obtusiloba. A, basal leaf; B, flower parts (a, petaloid; b, stamen; c, carpels); C, achene (A–B, Potanin 1872, LE; C, Grey-Wilson & Philips 471, K). Scale bars (applies to Figs 1–17): A 5 1 cm; B–C 5 1 mm. 60 S. N. ZIMAN ET AL. (1906). – Type: Not designated. We have examined the likely syntypes Duthie s.n. from Kashmir (K), Potanin s.n. from Kansu (LE) and Delavay s.n. from Yunnan (LE). Anemone obtusiloba D.Don var. coerulea Ulbr., Bot. Jahrb. Syst. 37: 242 (1906). – Type: Kashmir, Falconer 29 (lecto K!, designated here). Anemone obtusiloba D.Don var. chrysantha Ulbr., Bot. Jahrb. Syst. 37: 242 (1906). – Type: Tibet, Kardang, Hans (lecto K!, designated here). Anemone rupestris Hook.f. var. villosa Marquand & Shaw, J. Linn. Soc. 48: 154 (1929). – Type: Tibet, Nyima La, 3960–4270 m, in open pastures in the Rhododendron shrub belt, 22 vi 1924, Kingdon-Ward 5821 (holo K!). Anemone obtusiloba D.Don var. polysepala W.T.Wang, Acta Phytotax. Sin. 12: 169 (1974). – Type: China, Yunnan, Weisi, 3780 m, 29 v 1940, Feng 4270 (holo PE!). Anemone multisepala Qureshi & Chaudhri, Pakistan Syst. 2: 15 (1978). – Type: NW Pakistan, Chitral Distr., Gujargol near Lowari Pass, Bioban, Lowari Top, 17 vii 1977, Shah & Khan 2439 (holo ISL). Anemone neelamiana Qureshi & Chaudhri, Pakistan Syst. 2: 15 (1978). – Type: Azad Kashmir, Muzaffarabad Distr., Upper Neelam Valley, Dudgai, 3800 m, 28 vi 1978, Iqbal & Abbasi 1827 (holo ISL). Anemone obtusiloba D.Don var. leiocarpa Tamura, Acta Phytotax. Geobot. 37: 153 (1986). – Type: Nepal, Bhojpur Distr., Salpa-Bhanjyang-Charapani, 3320 m, 26 vi 1976, Tabata 10924 (holo KYO!). Basal leaves 5–10 (rarely to 20); petioles 3–15(–20) cm 6 1–2 mm, villous or pubescent; blades 3-sect (sometimes 3-parted or 3-lobed), reniform-pentagonal to broadly ovate, 2–6 6 2–10 cm, densely spreading villous to sparsely pubescent or sometimes abaxially subglabrous; bases cordate (rarely subtruncate); leaflets (or lobes) overlapping or remote; apical leaflets of 3-sect leaves on petiolules 1–2 mm long, 3-parted, 3-cleft, or 2- to 3-lobed, rarely undivided, rhombic-ovate to broadly rhombic, subequal or larger than lateral leaflets; with margins obtusely or acutely dentate; lateral leaflets subsessile, unequally 2- or 3-parted or 2- or 3-lobed, obliqueflabellate to ovate (Fig. 1A). Scapes 2–5, 5–25(–35) cm long, spreading-villous or pubescent, 1- or 2(–3)-flowered. Involucral bracts 3-parted, 3-cleft, 3-lobed or 3dentate (rarely undivided), broadly rhombic, rhombic-obovate, oblong-ovate, lanceolate, or cuneate, 1–3 cm long, appressed pilose. Pedicels 1–3(–5) cm long, appressed pilose. Petaloids 5–7, obovate or elliptic, white, bluish or yellowish, 5–15 6 3–8 mm, abaxially pilose; basal veins 3–5, anastomosing veins absent (rarely solitary) (Fig. 1Ba). Staminodes absent. Stamens 2.2–3 mm long; filaments linear, slightly narrowed apically, 0.5–0.8 mm long; anthers ellipsoid, connectives narrow (Fig. 1Bb). Carpels ovoid, 1.2–2 mm long, usually villous or sometimes subglabrous, hairs 0.5–1 mm long; styles straight or curved, 0.5–1.5 mm long (Fig. 1Bc). Achenes broadly ovoid, rarely slightly compressed, but without ribs, 3–6 6 2–3.5 mm, densely strigose, softly pubescent or subglabrous (hairs 0.6–1.6 mm long); styles apically hooked or substraight, 1–2.5 6 1–1.5 mm (Fig. 1C). REVISION OF ANEMONE SECT. HIMALAYICAE 61 Distribution and habitat. Afghanistan, Pakistan, Kirgizstan, Mongolia, W and SW China (NW Yunnan, SW Sichuan, E and S Xizang, Gansu, Shanxi, Sikang), North India (Kumaon, Simla, Punjab, Kashmir), Nepal, Bhutan, Burma (Fig. 18). In forests, at forest margins, meadows and alpine grassland; 1850–4800 m. Nomenclatural notes. In the protologue of Anemone obtusiloba Don (1825) cites the locality and collection/collector information as ‘Hab. In Gosaingsthan Nepaliae. Wallich.’. We studied several specimens collected by Wallich, of which only the BM specimen has a label closely corresponding to the protologue data: ‘Gosaingsthan Nepalensium, Wallich s.n.’ (BM!). The plant itself is undoubtedly Anemone obtusiloba, but the specimen bears no notes made by Don, and thus its identity as the holotype is questionable. In the Kew herbarium (K) the folder with ‘type specimens’ contains several herbarium sheets. We select here the BM specimen as the lectotype of Anemone obtusiloba. Taxonomic remarks. Don (1825) characterized Anemone obtusiloba as having 3-sect villous basal leaves with cuneate bases, incised-crenate margins and obtuse terminal lobes, 3-lobed involucral leaves, solitary flowers, 5 obovate petaloids, and pubescent achenes. Hooker (1872) noted the variability of Anemone obtusiloba, especially with respect to the size, colour and villosity of flowers and shape of basal leaves. Consequently, he suggested that there might be several infraspecific taxa. Brühl (1896) confirmed the evident variability within Anemone obtusiloba and was the first author to devise an infraspecific classification that included six subspecies and five varieties. In addition to Anemone obtusiloba subsp. obtusiloba (5 ‘typica’), Brühl recognized six others: subsp. trullifolia (with three varieties: var. linearis, var. spatulata and var. rotundifolia), subsp. coelestina, subsp. ovalifolia (with two varieties: var. geochares and var. orthocaula), subsp. saxicola, subsp. omalocarpella and subsp. imbricata. The first three subspecies (subsp. trullifolia, subsp. coelestina and subsp. ovalifolia) later were accepted as separate species (see below), whereas subsp. saxicola was discussed by Lauener (1960: 198) under A. rupestris, and subsp. omalocarpella (Wang et al., 2001: 323) under A. polycarpa. According to Ulbrich (1906), Anemone obtusiloba consists of two subspecies and two varieties: subsp. obtusiloba (5 ‘genuina’) and subsp. micrantha (differing mainly in size), and vars. coerulea and chrysantha (differing mainly in petaloid colour). Lauener (1960), however, recognized three subspecies: subsp. obtusiloba (with two varieties: var. obtusiloba and var. potentilloides), subsp. ovalifolia, and subsp. rockii which differ mainly in their leaf shapes. Wang (1974, 1980) differentiated five infraspecific taxa of Anemone obtusiloba in China: subsp. obtusiloba and subsp. ovalifolia (as above), and in addition subsp. megaphylla, subsp. leiophylla and var. polysepala. According to Wang, subsp. megaphylla and subsp. leiophylla differ from subsp. obtusiloba in their basal leaves; and var. polysepala by its more numerous petaloids. Hara & Williams (1979) noted that in Nepal Anemone obtusiloba was represented by subsp. obtusiloba and subsp. omalocarpella. Tamura (1986) described Anemone obtusiloba var. leiocarpa from Nepal (endemic to the Bhojpur District), which 62 S. N. ZIMAN ET AL. differed from subsp. obtusiloba in its more or less compressed, slightly marginate, glabrous or sparsely hispid carpels. However, Chaudhary (1988) considered that in Nepal there were four subspecies of Anemone obtusiloba: the forementioned subsp. obtusiloba, subsp. omalocarpella and subsp. potentilloides, but also the new subsp. nepalensis, which he differentiated by leaf and flower characters. Anemone obtusiloba was considered by Qureshi & Chaudhri (1978, 1988) to include two varieties (var. obtusiloba and var. potentilloides). In addition they described two endemics from Pakistan, Anemone multisepala and A. neelamiana, closely related to A. obtusiloba. Anemone multisepala was stated to differ from A. obtusiloba in having narrowly obovate-cuneate leaflets and smaller but more numerous yellow petaloids, and from A. neelamiana in having white flowers with 8 petaloids. More recently Thothathri & Uniyal (1982) and Riedl & Nasir (1991) reduced these recently described Pakistani species to synonymy under Anemone obtusiloba due to the considerable overlapping variability. Our analysis leads us to agree with Lauener (1960) in recognizing Anemone trullifolia as a distinct species and not as a subspecies of A. obtusiloba. However, we treat Anemone rockii and A. geum (5 A. ovalifolia) not as subspecies but as species. We also agree with Lauener in regarding Anemone discolor and A. micrantha as synonyms of A. obtusiloba as there are no distinct characters to separate them. With respect to Anemone multisepala and A. neelamiana, we follow Thothathri & Uniyal (1982) and Riedl & Nasir (1991) in regarding these as synonyms of A. obtusiloba. In our opinion, Anemone obtusiloba subsp. saxicola and Anemone obtusiloba subsp. omalocarpella should be included in A. rupestris due to the similarity of their basal leaves and especially their achenes. Anemone obtusiloba subsp. leiophylla and Anemone obtusiloba subsp. potentilloides should rather be treated as varieties because of their weak distinguishing characters. Thus, we recognize three subspecies in Anemone obtusiloba: subsp. obtusiloba (with two varieties, var. leiophylla and var. potentilloides), subsp. megaphylla and subsp. nepalensis, whose differences are given in the following key. Key to subspecies and varieties 1a. Petioles of basal leaf blades 15–20 cm long; leaf blades 3–6 6 4–10 cm, sparsely pubescent; scapes 25–35 cm long; petaloids 5, 10–15 6 5–10 mm ___________ ________________________________________________ 1B. subsp. megaphylla 1b. Petioles of basal leaf blades 3–15 cm long; leaf blades 1–6 6 1–8 cm, pubescent or villous; scapes mostly less than 20 cm long; petaloids 5–7, 7–12 6 4–8 mm 2 2a. Scapes only up to 10 cm long; basal leaves with blades 1.5–2.5 6 2.5–3.5 cm, pubescent, with acute teeth _________________________ 1C. subsp. nepalensis 2b. Scapes usually longer than 10 cm; basal leaves with blades up to 4 6 6 cm, often villous and with obtuse teeth. 1A. subsp. obtusiloba _____________________ 3 3a. Basal leaves densely spreading-villous, mostly with obtuse teeth ____________ __________________________________________________ 1Aa. var. obtusiloba REVISION OF ANEMONE SECT. HIMALAYICAE 63 3b. Basal leaves pubescent, often with acute teeth __________________________ 4 4a. Petaloids blue; basal leaves with obtuse to acute teeth, apical leaflets 3lobed _________________________________________ 1Ab. var. potentilloides 4b. Petaloids white; basal leaves with acute teeth, apical leaflets 3-sect __________ ___________________________________________________ 1Ac. var. leiophylla 1A. subsp. obtusiloba Petioles of basal leaves 5–15 cm long; blades mainly 3-sect, sometimes 3-parted to 3lobed, broadly ovate or reniform-pentagonal, usually obtusely dentate, 1–4 6 1–6 cm, densely spreading-villous to sometimes sparsely pubescent; apical leaflets 3-lobed to 3-cleft, rhombic-ovate. Scapes 5–20 cm long. Involucral bracts 3-parted to 3-lobed or sometimes 3-dentate, rarely undivided, rhombic to cuneate, 1–2 cm long. Petaloids 5–7, white, blue or yellowish, 8–12 6 3–8 mm. Distribution and habitat. As the species. 1Aa. var. obtusiloba Petioles of basal leaves 5–12 cm long; blades 3-sect, broadly reniform-pentagonal, obtusely dentate, 2–4 6 2–6 cm, densely spreading-villous; apical leaflets 3-lobed, rhombic-ovate. Scapes 5–20 cm long. Involucral bracts 3-parted to 3-lobed, rhombic or cuneate. Petaloids 5–7, white, blue or yellowish, 8–12 6 5–8 mm. Distribution and habitat. As the species. Selection of herbarium specimens. AFGHANISTAN. Nuristan: Kamdesh, 2800 m, 22 vi 1959, Gilli 766 (W). PAKISTAN. NW Frontier: Hazara Distr., Murree Hills, 15 vi 1955, Webster & Sack 5719 (K); Mukshpuri, 2500 m, 3 v 1970, Ecker 19737 (W); Hazara, Pipe Line, 31 v 1975, Chaudhri & Qureshi 306 (W); Chitral Distr., Gujargol near Lowari Pass, Bioban, Lowari Top, 17 vii 1977, Shah & Khan 2439 (ISL). MONGOLIA. 1895, Przewalski s.n. (without geographic details and number) (LE); Archengsi, Cecerle Ich-Tamir, 1850 m, 26 vi 1978, Knapp 233179 (LE). CHINA. Yunnan: Ca-long-pin, 18 vii 1889, Delavay s.n. (LE); Lidjiang [Likiang], Mt. Yulung-Schan, vi 1914, Handel-Mazzetti 4078 (WU); Djinscha-djiang [Yangtze], Landsangdjiang [Mekong], Lenago, 4050 m, 7 vi 1916, Handel-Mazzetti 8856 (WU); Likiang, eastern slopes of Likiang Snow Range, 25 vi 1922, Rock 4767 (W); Chung-tien, Mts. West Hsiao Chung-tien, v 1932, Rock 24666 (K). Sichuan: Da-Dsian lu (Tatsien-lou, Tarsando), 18 v 1893, Potanin s.n. (LE); Mts. of Kulu, Muli Territory, 1932, Rock 23953 (K). Kansu: T’ao River basin, alpine meadows, vi 1925, Rock 12254 (K); Terra Tangutorum, 14 vii 1872, Potanin s.n. (LE); Reg. Tangutica, 1873, Plantae a Przewalski collectae (E); ibid., 1880 (E, LE); Valle Runwyz, 1885, Potanin s.n. (LE); Berg Shao Wu Taishan, Waldgreuse, vii 1879, Moellendorf 612 (WU). Xizang: Montes Tala-gui, 4 vi 1884, Przewalski 92 (LE); Rija simitra, 11 vi 1884, Przewalski 123 (LE); Kam, Djagyn-gol, 1900, Ladygyn 25 (LE); Dohpa, 13–14000 ft, 15 vii 1900, Ladygyn 264 (LE); 17 vii 1900, Ladygyn 645 (LE); Entok-gomba, 11500 ft, 26 iv 1901, Ladygyn 25 (LE). 64 S. N. ZIMAN ET AL. NEPAL. Yarpang, 16000 ft, 10 viii 1932, Sharma 3415 (BM); Jumla, 7500 ft, 4 v 1952, Polunin et al. 906 (K); Gandak Kosi Watershed, 12000 ft, 5 v 1953, Proud 150 (BM); Chhairo gaun, North of Tukucha, 10000 ft, 2 vi 1954, Stainton et al. 877 (BM); Ankhu Khola, Barand, 10200 ft, 5 vi 1962, Bowes-Lyon 103 (BM); Bagmati Zone, Gossain Kurda, 3000 m, 8 v 1967, Nicolson 3299 (BM); Dolpo Valley, 5 mi East of Ting Kyu, 4800 m, 2 viii 1973, Grey-Wilson & Philips 471 (K); Purangaun, above Garunga, North of Parbat, 2700 m, 4 v 1976, Wormald 116 (BM). BHUTAN. Phaksana, 10500 ft, 6 vi 1971, Ramesh Bedi 68 (E). INDIA. Kashmir: Keslitoar, 10000 ft, Thomson 4688 (K); Simla La, 13–14000 ft, 22 i 1924, Kingdon-Ward 5821 (BM); Apharwat, 12000 ft, 12 viii 1956, Polunin 56/220 (BM); Gulmarg, above Khelanmarg, 30 v 1971, Robson 1911 (BM); Upper Chenab: Pangi Valley, 23 v 1881, Ellis 1141 (WU); Dehra Dun, Kulu Valley, 8 vi 1949, Basa s.n. (W); Kulu Valley, Pulga, 17 vi 1950, Jain & Bharadwaja s.n. (W); Simla Hill State: Punjab, Kamru, Baspa Valley, 13000 ft, 26 vi 1939, Sheriff 7345 (BM); Simla, Huttoo, 10000 ft, 20 v 1956, Chiddall 38 (BM). KIRGIZSTAN. Tien-Shan: Upper part of the basin of Sary-Dzhas, close river Kaska-Ter, 3 vii 1902, Sapozhnikov s.n. (LE); Semirechenski Distr., close Przewalski, river Tjuz, 3 vii 1912, Sapozhnikov s.n. (LE); Basin of Sary-Dzhas, close river Gjuz, 15 vii 1959, Zviagina s.n. (LE). 1Ab. var. potentilloides Lauener, Notes Roy. Bot. Gard. Edinburgh 23: 187 (1960). – Type: supra Shupien, in herbosis excelsissimus, Jacquemont 665 (holo K!; iso BM!). – Anemone potentilloides Cambess. ex Besant, Gard. Chron. 177: 371 (1927), nom. subnud. – Anemone obtusiloba subsp. potentilloides (Lauener) Chaudhary, Bot. Zhurn. 73: 1188 (1988). Petioles of basal leaves 5–8 cm long; blades mainly 3-sect, reniform-pentagonal, obtusely to acutely dentate, 1–3 6 1–2 cm, pubescent; apical leaflets 3-lobed, broadly rhombic. Scapes 10–20 cm long. Involucral bracts mostly 3-lobed, oblongrhombic, 1–2 cm long. Petaloids 5–6, blue, 8–12 6 4–8 mm. Distribution and habitat. India (Himachal Pradesh, Lahul, Kashmir), Pakistan (Chitral, Hazara) (Fig. 18); 2700–4000 m. Selection of herbarium specimens. INDIA. Kashmir: Apharwat, 12000 ft, 12 viii 1956, Polunin 56/220 (BM); Gulmarg, above Khelanmarg, 30 v 1971, Robson 1911 (BM). PAKISTAN. NW Frontier: Hazara Distr., Murree Hills, 15 vi 1955, Webster & Sack 5719 (K); Mukshpuri, 2500 m, 3 v 1970, Ecker 19737 (W); Hazara: Pipe Line, 31 v 1975, Chaudhri & Qureshi 306 (W). 1Ac. var. leiophylla (W.T.Wang) Ziman, Ehrendorfer & Bulakh, stat. nov. – Anemone obtusiloba D.Don subsp. leiophylla W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 350 (1980). – Type: China, NW Yunnan, Gongshan Drung-Nu Zu Zi Zhixian, 1938, T.T. Yu 22076 (holo PE!). Petioles of basal leaves 5–12 cm long; blades 3-sect, reniform-pentagonal, acutely dentate, 1–4 6 2–6 cm, sparsely pubescent; leaflets remote, apical 3-cleft, broadly rhombic. Scapes 8–25(–35) cm long. Involucral bracts 3-dentate or undivided, oblong-rhombic, 1–2 cm long. Petaloids 5–7, white, 7–12 6 3–7 mm. REVISION OF ANEMONE SECT. HIMALAYICAE 65 Distribution and habitat. China (Yunnan) (Fig. 18). At forest margins and on grassy slopes; 2900–3000 m. Endemic of NW Yunnan (Gonshan Drung-Nu-Zi Zhixian). Known only from the type collection. 1B. subsp. megaphylla W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 350 (1980). – Type: China, SW Yunnan, Chengkang, Snow Range, grassy slopes, 3400 m, 24 vii 1938, T.T. Yu 16956 (holo PE!). Petioles of basal leaves 15–20 cm long; blades 3-parted, reniform-pentagonal, acutely dentate, 3–6 6 4–10 cm, sparsely pubescent; lobes overlapping, apical leaflet 3-cleft, broadly rhombic. Scapes 25–35 cm long. Involucral bracts 3-lobed, rhombic or rhombic-obovate, 2–3 cm long. Petaloids 5, blue, 10–15 6 5–10 mm. Distribution and habitat. China (SW Yunnan: Zhenkang Xian) (Fig. 18), on grassy slopes; c.3400 m. Known only from the type collection. 1C. subsp. nepalensis Chaudhary, Bot. Zhurn. 73: 1188 (1988). – Type: Nepal, Jaljale, 3500 m, 23 vi 1979, Sajiu & Tuladhar s.n. (holo KATH!). Petioles of basal leaves 3–5 cm long; blades 3-sect, rhombic-ovate, 1.5–2.5 6 2.5– 3.5 cm, pubescent, margins acutely dentate, apical leaflets 3-parted. Scapes 5–10 cm long. Involucral bracts 3-sect, ovoid, 1–1.2 cm long. Petaloids 5, white, basally bluish, c.10 6 8 mm. Distribution and habitat. Central Nepal (Okhaldhunga, Dhading, Baglung) (Fig. 18); 3500 m. Selection of herbarium specimens. NEPAL. Okhaldhungagaon, South of Dhorpatan, 10000 ft, 1 v 1954, Stainton et al. 350 (BM), 374 (BM), 376 (BM); Jaljale, 3500 m, 23 vi 1979, Sajiu & Tuladhar s.n. (KATH!). 2. Anemone patula C.C.Chang ex W.T.Wang, Acta Phytotax. Sin. 12: 169 (1974). – Type: China, Central Sichuan, Li Xian, 3500 m, 14 vi 1936, K.L. Chu 2796 (holo E!). Figs 2, 19. Basal leaves 5–9; petioles 3–10 cm 6 c.1 mm, subglabrous; blades 3-sect, orbicularovate, 0.5–3.5 6 1–4 cm, sparsely pubescent or glabrescent, bases deeply cordate; leaflets subsessile; apical leaflets 3-lobed, flabellate-rhombic, lobes flabellateobovate, margins lobulate, lobules narrowly ovate, sometimes denticulate; lateral leaflets 3-lobed, obliquely flabellate (Fig. 2A). Scapes 3–6, 5–25 cm long, subglabrous; flowers solitary. Involucral bracts 3-lobed to 3-dentate, rarely undivided, obovate, 1–2 cm long, puberulent or subglabrous. Pedicels 2–5 cm long, puberulent or subglabrous. Petaloids 5(–6), white or blue-purple, ovate or broadly 66 S. N. ZIMAN ET AL. F I G . 2. Anemone patula C.C.Chang ex W.T.Wang var. patula. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). obovate, 4–12 6 2–9 mm, puberulent, basal veins 3–7, anastomosing veins absent. Stamens 3–5 mm long; filaments lanceolate (Fig. 2Ba). Carpels 2–4 mm long, villous (Fig. 2Bb). Achenes unknown. Distribution and habitat. China (Sichuan) (Fig. 19). In Abies forests and in thickets; 3500–3530 m. Taxonomic remarks. Wang (1974) described Anemone patula as a species allied to A. rupestris but differing in the shape of the basal leaflets (subsessile and 3-lobed), larger petaloids and strongly villous carpels. Based on these characters, but also on its solitary flowers and lanceolate filaments, Anemone patula is closer to A. obtusiloba and keys out in series Obtusilobae. Additional study is needed in this species due to the limited herbarium material and lack of data on achenes. Wang (1980) recognized two varieties of Anemone patula, var. patula and var. minor, which differ in scape and leaf size, and the size, colour and villosity of the flowers. These varieties were later accepted by Wang et al. (2001). Key to varieties 1a. Petaloids white or blue-purple, 8–12 6 5–9 mm; scapes 10–25 cm long; leaf blades 1–3.5 6 1.5–4 cm _______________________________ 2a. var. patula 1b. Petaloids white, 4–6 6 2–3 mm; scapes 5–10 cm long; leaf blades 0.5–1 6 1– 2 cm __________________________________________________ 2b. var. minor 2a. var. patula Leaf blades 1–3.5 6 1.5–4 cm. Scapes 10–25 cm long. Petaloids white or blue-purple, 8–12 6 5–9 mm. REVISION OF ANEMONE SECT. HIMALAYICAE 67 Distribution and habitat. Central Sichuan (Li Xian). In Abies forests; 3500 m. Known only from the type collection. 2b. var. minor W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 350 (1980). – Type: China, Sichuan, Baoxing Xian, 3530 m, 1936, K.L. Chu 923 (holo PE!). Leaf blades 0.5–1 6 1–2 cm. Scapes 5–10 cm long. Petaloids white, 4–6 6 2–3 mm. Distribution and habitat. Central and western Sichuan (Baoxing Xian, Jinchuan Xian). In thickets; 3530 m. Known only from the type collection and a few specimens collected in the same area and cited in the protologue. 3. Anemone polycarpa W.E.Evans, Notes Roy. Bot. Gard. Edinburgh 13: 154 (1921). – Anemone rupestris subsp. polycarpa (W.E.Evans) W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 43 (1980). – Type: China, Yunnan, ‘Mekong-Salween divide. Alt. 12000 ft, ix 1914, Forrest 13320’ (holo E!). Figs 3, 19. Anemone rupestris Wall. ex Hook.f. & Thoms., Fl. Ind. (1855), pro parte (excl. type). Leaves 5–12, petioles 5–12 cm 6 c.1 mm, sparsely puberulent; blades twice 3-sect, ovate, 3–5 6 2–4 cm, sparsely puberulent, bases rounded; apical leaflets on petiolules 10–15 mm long, 3-parted, broadly ovate; lateral leaflets on petiolules 3–5 mm long, 3-lobed, ovate (Fig. 3A). Scapes 2–5, 5–20 cm long, 1–3-flowered, sparsely puberulent. Involucral bracts 3-parted, obovate-rhombic, 1–3 cm long, puberulent. Petaloids 5–6(–7), monomorphic, reddish-white, ovate-elliptic, 7–14 6 4–9 mm, F I G . 3. Anemone polycarpa W.E.Evans. A, basal leaf; B, flower parts (a, petaloid; b, staminode; c, stamen; d, carpel); C, achene (A, Flora of China, 2001; B, Forrest 20738, E; C, Stainton et al. 5403, WU). 68 S. N. ZIMAN ET AL. sparsely puberulent, basal veins 3–5, anastomosing veins absent (Fig. 3Ba). Staminodes present (Fig. 3Bb). Stamens 3–4 mm long; filaments linear (Fig. 3Bc). Carpels up to 80, long ellipsoid, 3–3.5 mm long, villous, styles hooked (Fig. 3Bd). Achenes fusiform, compressed, 3–4 6 2–2.5 mm, sparsely puberulent along medial line, hairs dimorphic (c.1 mm and 0.3 mm long); ribs absent; styles apically thickened and uncinate, 1–4 6 1.5 mm (Fig. 3C). Distribution and habitat. China (NW Yunnan, Sichuan, SE Xizang), Nepal, North India (Assam), Bhutan (Fig. 19). On grassy and rocky slopes; 3500–4800 m. Taxonomic remarks. Anemone polycarpa was described as a species close to A. obtusiloba (Evans, 1921), but differing mainly in the pinnatisect basal leaf lobes, numerous hairy carpels, and achenes with uncinate styles. Lauener (1960), however, proposed to reduce it to a synonym of Anemone rupestris, and both Wang (1980) and Chaudhary (1988) treated it as A. rupestris subsp. polycarpa (W.E.Evans) W.T.Wang. Sharma et al. (1993) regarded Anemone polycarpa as a distinct species and Wang et al. (2001) accepted this status but they included in it A. saxicola (Brühl) Tamura & Kitamura based on A. obtusiloba D.Don subsp. saxicola Brühl. Our examination of the holotype (Forrest 13320) from Yunnan, and also Stainton et al. 5403 from Nepal, suggests that Anemone polycarpa differs from A. rupestris in the 3parted involucral bracts, larger monomorphic puberulent petaloids, narrower filaments, presence of staminodes, less compressed but more elongate achenes without ribs and with few dimorphic hairs, and apically thickened uncinate styles. Consequently, we regard this taxon as a species which is closer to Anemone obtusiloba than to A. rupestris. Selection of herbarium specimens. CHINA. NW Yunnan: Mekong-Salween Divide, vi 1921, Forrest 19578 (K); Doker-la, Mekong, Salween Divide, ix 1921, Forrest 20738 (E); Mekong, Salween Divide, Sila, 4000 m, 16 viii 1938, T.T. Yu 22374 (E); Upper Kjukiang Valley (Clulung), South of Lungtsahmura, 3800 m, 10 viii 1938, T.T. Yu 1958 (E). Sichuan: Dongregro, 4500 m, 21 vii 1922, Smith 3262 (BM); Sung pan hsien, 8 viii 1928, Fang 4063 (E). Qinghai: Maqin Xian, Dawu Xiang, 3500 m, 31 vii 1993, T.N. Ho & Bartolomew 618 (BM). NEPAL. Central Nepal: Bhurungdi, Khola, 22 v 1954, Stainton et al. 5403 (WU); Lamjung Himal, 4500 m, 13 ix 1954, Stainton et al. 6326 (BM, E); South of Bhubnjeng Kharka, 3400 m, 9 viii 1974, de Haas 2137 (BM); Sagarmatha Zone, Solukhumbu Distr., Pike Peak, 3560 m, Miyamoto et al. 84048 (BM). INDIA. Assam: Tha Chu Valley, 11000 ft, 9 vii 1950, Kingdon-Ward 19594 (BM). BHUTAN. Shing Be Me La, 10500 ft, 16 v 1949, Ludlow & Sheriff 20639 (BM). 4. Anemone subpinnata W.T.Wang, Acta Phytotax. Sin. 12: 170 (1974). – Type: China, Szechuan: Mu-li, Wachin, Ching-chang, 3750 m, 21 vi 1937, T.T. Yu 6512 (holo PE!). Figs 4, 19. Leaves 4–7, petioles 2–5 cm 6 1–2 mm, villous; blades apparently pinnatifid, but in principle 3-sect, elliptic, c.2.5 6 1.8 cm, densely pubescent, bases subcordate or broadly cuneate; apical leaflets with petiolules 3–5 mm long, 3-sect, REVISION OF ANEMONE SECT. HIMALAYICAE 69 F I G . 4. Anemone subpinnata W.T.Wang. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). rhombic-obovate, margins incised-dentate with secondary leaflets 3-parted; lateral leaflets subsessile, unequally 3-lobed, obliquely cuneate (Fig. 4A). Scapes 2–5, 3– 10 cm long, villous, 1-flowered. Involucral bracts subequally 3-lobed, elliptic-ovate to oblong-ovate, c.1 cm long; lobes entire or apical lobes 3-dentate, puberulent, apically obtuse. Pedicels 1–4 cm long, puberulent. Petaloids 5, white, tinged blue, purplish or violet, elliptic-obovate, 5–8 6 4–6 mm, villous, basal veins 3–5, anastomosing veins absent. Stamens 2–4 mm long; filaments linear (Fig. 4Ba). Carpels cylindric, 2.7–3.2 mm long, villous; styles curved (Fig. 4Bb). Achenes unknown. Distribution and habitat. China, endemic of SW Sichuan, Muli, Wachin (Fig. 19). In alpine meadows; 3700 m. Taxonomic remarks. Wang (1974) described this species as a narrow endemic from Sichuan, taxonomically close to Anemone obtusiloba but differing in its subpinnate basal leaf blades, solitary flowers and strongly villous carpels. According to our data, Anemone subpinnata differs from A. obtusiloba also in its wider basal leaf petioles, entire (sometimes 3-lobed) involucral bracts, and linear filaments. Additional studies are needed due to the lack of data on achenes and the limited herbarium material available. Known only from the type collection and a few specimens collected in the same area and cited in the protologue. 5. Anemone rockii Ulbr., Notizbl. Bot. Gart. Berlin-Dahlem 10: 876 (1929). – Anemone obtusiloba D.Don subsp. rockii (Ulbr.) Lauener, Notes Roy. Bot. Gard. Edinburgh 23: 188 (1960). – Type: China, SW Kansu, southern slopes of Minshan, South of Shimen, in crevices of limestone cliffs, along streams, 10600 ft, vi 1925, Rock 12520 (lecto K!, designated here by Ziman and Mosyakin; iso E!). Figs 5, 20. Leaves 5–15; petioles 5–20 cm 6 3 mm, sparsely puberulent or glabrescent; blades 3sect, broadly ovate, 2.5–5 6 2.5–5 cm, glabrous or abaxially sparsely puberulent, bases cordate; apical leaflets on petiolules 1–2 mm long, 3-lobed, broadly ovate or rhombic, lobes incised lobulate; lateral leaflets subsessile, unequally 2- or 3-lobed, 70 S. N. ZIMAN ET AL. F I G . 5. Anemone rockii Ulbr. var. rockii. A, basal leaf; B, flower parts (a, petaloid; b, stamens; c, carpel); C, achene (A, Flora of China, 2001; B, Forrest 30471, BM; C, Polunin 906, E). obliquely flabellate; lobes and lobules contiguous, overlapping or remote (Fig. 5A). Scapes 2–10, 10–30 cm long, glabrous or sparsely puberulent; flowers solitary. Involucral bracts undivided or 3-lobed, obovate, 1–3 cm long, margins dentate or entire. Pedicels 2–6(–10) cm long, puberulent. Petaloids 5(–6–9), dimorphic, white, blue (rarely purplish), oblong-obovate, 10–20 6 6–12 mm, sparsely puberulent or glabrous, basal veins 3–5, anastomosing veins solitary or absent (Fig. 5Ba). Staminodes sometimes present. Stamens 4–5 mm long; filaments sublinear, 0.3–0.5 mm wide (Fig. 5Bb). Carpels 2.8–3.5 mm long, cylindric, densely or sparsely puberulent; styles straight (Fig. 5Bc). Achenes ovoid, slightly compressed, 3–4 6 1–1.5 mm, glabrous or sparsely to densely pubescent (hairs c.1 mm long); styles slightly bent, 1.3–1.4 mm long (Fig. 5C). Distribution and habitat. China (Yunnan, Sichuan, Gansu, Xizang), Nepal (Fig. 20). On grassy slopes; 2100–4000 m. Taxonomic remarks. Anemone rockii was separated from A. obtusiloba by Ulbrich (1929) on the basis of several collections made by Rock in SW Kansu, western China, which had more robust stems, more divided basal leaves with a deeper sinus and larger flowers. Having examined syntypes from K and BM, we note that Anemone rockii differs from A. obtusiloba also by its wider basal leaf petioles, subglabrous leaf blades, solitary flowers, undivided or 3-lobed involucral bracts, subglabrous usually dimorphic petaloids, linear staminodes, and densely pubescent achenes with slightly bent styles. We believe these distinctions merit recognizing Anemone rockii as a distinct species. According to Wang (1980) and Wang et al. (2001), Anemone rockii consists of three varieties. Although the majority of plants throughout the distribution of this species belong to Anemone rockii var. rockii, REVISION OF ANEMONE SECT. HIMALAYICAE 71 several plants from SW China with remote basal leaf blade lobes and distinctly pubescent achenes have been recognized as distinct varieties, var. pilocarpa W.T.Wang and var. multicaulis W.T.Wang, differing in the number and colour of their petaloids. Key to varieties 1a. Leaf blade lobes and lobules contiguous or overlapping; achenes glabrous or subglabrous ____________________________________________ 5a. var. rockii 1b. Leaf blade lobes and lobules remote; achenes densely pubescent __________ 2 2a. Petaloids 8–9, white __________________________________ 5b. var. pilocarpa 2b. Petaloids 5–6, white or blue __________________________ 5c. var. multicaulis 5a. var. rockii Basal leaf blade lobes and lobules contiguous or overlapping. Petaloids 7–8, white or purplish. Achenes glabrous or subglabrous. Distribution and habitat. China (Yunnan, Sichuan, Gansu, Xizang), Nepal (Fig. 20). On grassy slopes; 2100–4000 m. Selection of herbarium specimens. CHINA. Yunnan: Distr. of Chung-tien, 13000 ft, v 1932, Rock 24666 (BM). Sichuan: Distr. of Tchen-keou-tin, Farges s.n. (P); top of Mt. Wa, vii 1903, Wilson 3052 (K). Kansu: Upper Tebbu County, southern slopes of Minshan, South of Shimen in crevices of limestone clifts along streams, 10600 ft, vi 1925, Rock 12408 (BM), 12487 (BM), 12520 (K); gravelly slopes at foot of Shimen, 12000 ft, vii–viii 1925, Rock 13061 (BM, E, K); Minshan near Djirakana Shimen limestone crevices, x 1925, Rock 13626 (BM, E, K). Xizang: Mt. Kenichumpo, Salween and Irawady Divide, 14000 ft, vi 1932, Rock 21952 (BM), 21953 (BM). NEPAL. West Nepal: Chankheli Lagna, 30 iv 1952, Polunin et al. 4345 (E); Saldanggaon, v 1952, Polunin et al. 16 (E); Jumla, v 1952, Polunin et al. 906 (E); Pansala, vi 1952, Polunin et al. 880 (E); Myagdi, Dara, 20 vi 1952, Polunin et al. 4344 (E); Lete Kali Gandaki, 20 iv 1954, Stainton et al. 581 (E). East Nepal: Uttar Ganga, iv 1954, Stainton et al. 969 (E); Baglurg, South of Dhorpatan, v 1954, Stainton et al. 356 (E); Prangburg, vi 1954, Stainton et al. 877 (E). 5b. var. pilocarpa W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 350 (1980). – Type: China, Yunnan, Chengkou Xian, 2100 m, 27 viii 1935, C.C. Chang 2064 (holo PE!). Basal leaf blade lobes and lobules remote. Petaloids 8–9, white. Achenes densely pubescent. Distribution and habitat. China (Yunnan: Chengkou Xian). On grassy slopes; 2100 m. Nomenclatural notes. Plants of this taxon were collected by C. C. Chang in 1935. He initially annotated them as ‘A. pseudo-rockii sp. nova’, but this name was never published. In 1980 Wang published it as Anemone rockii var. pilocarpa. Known only from the type collection. 72 S. N. ZIMAN ET AL. 5c. var. multicaulis W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 350 (1980). – Type: China, Sichuan, Xichang Shi, 3000 m, 1977, W.T. Wang 40171 (holo PE!). Basal leaf blade lobes and lobules remote. Petaloids 6, white or blue. Achenes densely pubescent. Distribution and habitat. China (SW Sichuan: Xichang Shi). On grassy slopes; 3000 m. Known only from the type collection. 6. Anemone geum H.Lév., Bull. Acad. Int. Geogr. Bot. (Le Mans) 25: 25 (1915). – Anemone bonatiana var. geum (H.Lév.) H.Lév., Cat. Pl. Yun-Nan 219 (1917). – Type: China, Yunnan, lagune du haut plateau de Ta-Hai-Tse, 3200 m, v 1912, Maire s.n. (holo E!). Figs 6, 19. Anemone obtusiloba Franch., Plantae Delav. 8 (1889), non D.Don (1825). Anemone obtusiloba D.Don subsp. ovalifolia Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Anemone ovalifolia (Brühl) Hand.-Mazz., Symb. Sin. 7: 315 (1931). – Anemone geum H.Lév. subsp. ovalifolia (Brühl) Chaudhary, Bot. Zhurn. 73: 1188 (1988). – Type: NW China, Yunnan, Yungning, Handel-Mazzetti 3157 (holo WU!) Anemone obtusiloba D.Don var. geochares Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: China, Regio Tangutorum (N Tibet), 1880, Przewalski s.n. (lecto LE!, designated here by Ziman; iso E). Anemone obtusiloba D.Don var. orthocaula Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: India, Kumaon near the Lebung pass, 14–15000 ft, 1 viii 1886, Duthie 5272 (holo BM!; iso LE!). Anemone wardii Marquand & Shaw, J. Linn. Soc. 48: 154 (1929). – Type: Tibet, Doshong La, Pemako, 25 x 1924, Kingdon-Ward 6262 (holo K!). F I G . 6. Anemone geum H.Lév. A, basal leaf; B, flower parts (a, petaloid; b, staminodes; c, stamen; d, carpel); C, achene (A–B, Handel-Mazzetti 6671, WU; C, Smith 10642, BM). REVISION OF ANEMONE SECT. HIMALAYICAE 73 Anemone rupestris Hook.f. var. pilosa Marquand & Shaw, J. Linn. Soc. 48: 154 (1929). – Type: Temo La, 4270 m, in pastures among dwarf Rhododendron on the open moorland, 6 vi 1924, Kingdon-Ward 5747 (holo K!). Anemone obtusiloba D.Don subsp. ovalifolia Brühl var. polysepala W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 43 (1980). – Type: China, NW Yunnan, 3780 m, 29 vii 1970, Feng 4070 (holo PE!). Anemone obtusiloba D.Don subsp. ovalifolia Brühl var. angustilimba W.T.Wang, Fl. Reipubl. Popul. Sin. 28: 43 (1980). – Type: China, SW Sichuan, 25 viii 1936, W.T. Wang 660929 (holo PE!). Basal leaves 5–15, petioles villous, 3–15 cm 6 2–3 mm; blades ovate, 3-sect or sometimes 3-parted, 2–5 6 2–5 cm, densely long hairy or sparsely short hairy; bases subcordate or subtruncate; apical leaflets of 3-sect leaves on petiolules 5–10 mm long, 3-lobed or undivided, broadly rhombic, longer (sometimes much) longer than lateral leaflets which have petiolules 1–3 mm long and are 2- or 3-lobed (Fig. 6A). Scapes 2–5, 5–25 cm long, villous or pubescent; flowers solitary. Involucral bracts undivided or 3-lobed, 1–2 cm long, puberulent. Pedicels 1–2(–6) cm long, puberulent. Petaloids 5(–8), broadly ovate, white, yellowish, mauve or blue, 5–12 6 4–9 mm, abaxially pubescent; basal veins 3(–5), anastomosing veins absent (Fig. 6Ba). Staminodes sometimes present, c.2 mm long (Fig. 6Bb). Stamens 2–3 mm long; filaments lanceolate, 0.5–1 mm wide (Fig. 6Bc). Carpels narrowly ovoid, 2–3 mm long, pubescent or subglabrous, hairs 0.5–0.7 mm long; styles straight, 0.5–1.5 mm long (Fig. 6Bd). Achenes ovoid, sometimes slightly compressed, 3–4.8 6 1.6–2.3 mm, villous or subglabrous, hairs 0.8–1.3 mm long; styles straight, 1.8–2.6 mm long (Fig. 6C). Distribution and habitat. Pakistan, China (Quinghai, NW Yunnan, W Sichuan, Xizang, S Xinjiang), Nepal, North India (Assam, Himachal Pradesh, Uttaranchal, Sikkim) (Fig. 19). In scrub and alpine meadows; 1900–4800 m. Chaudhary (1988) reported the occurrence of this taxon in Bhutan (without localities) and in Burma (Shan, Mt. Victoria) but we were unable to find any herbarium collections from these localities. Taxonomic remarks. Brühl (1896) described typical Anemone obtusiloba subsp. ovalifolia as having ovate 3-sect basal leaf blades and slightly compressed hirsute or glabrous pistils and achenes. He recognized two varieties, var. geochares and var. orthocaula, which differ in having prostrate-procumbent or erect-ascendent scapes, respectively. Several years later Léveille (1915) described plants from China (Yunnan) having characters similar to ‘ovalifolia’ of Brühl as Anemone geum H.Lév. Shortly afterwards he (Léveille, 1917) reduced it to Anemone bonatiana var. geum. Handel-Mazzetti (1931) then described plants with ovate 3-sect basal leaf blades from Yunnan as Anemone ovalifolia (Brühl) Hand.-Mazz. by raising Bruhl’s variety to specific status. Under this name he included Anemone rupestris var. lobata and A. obtusiloba subsp. micrantha Ulbr. pro parte based on the similar shape of the 74 S. N. ZIMAN ET AL. basal leaves. Later Lauener (1960) reinstated this taxon as Anemone obtusiloba subsp. ovalifolia and treated A. geum as its synonym. Wang (1974, 1980) also considered Anemone geum as a synonym of A. obtusiloba subsp. ovalifolia, but recently Wang et al. (2001) accepted A. geum as a distinct species with two subspecies, subsp. geum and subsp. ovalifolia. After examination of specimens from Nepal and India, Hara & Williams (1979) and Sharma et al. (1993) recognized Anemone geum and included A. ovalifolia as a synonym. Chaudhary (1988) considered Anemone geum to include two subspecies, subsp. geum and subsp. ovalifolia, which, although occurring in the same area, differed mainly in rhizome shape and the pubescence of the basal leaves. According to the results of our examination of the type specimens of Anemone geum (Maire from Yunnan) and A. ovalifolia (Handel-Mazzetti from Yunnan), plus examination of other herbarium specimens (mainly from K, BM and E), we confirm the morphological similarity of these plants and believe all of them belong to A. geum. The species name by Léveille (1915) has priority over the combination A. ovalifolia (Brühl) Handel-Mazzetti (1931). Despite our recent recognition of Anemone geum subsp. geum and A. geum subsp. ovalifolia (Wang et al., 2001), here we reject these taxa. Wang (1980) recognized three varieties within Anemone geum (albeit using the name A. obtusiloba subsp. ovalifolia): var. ovalifolia, var. polysepala and var. angustilimba. Variety polysepala was described as differing from the other varieties by its greater number of petaloids and was distinguished further from var. angustilimba by the shape of its basal leaf blades. Following our examination of the type specimens of Wang’s var. polysepala (Feng 4070) and var. angustilimba (W.T. Wang 660929), we regard all these ‘differential’ characters as too variable to be of taxonomic significance. Therefore we conclude that it would be better to recognize A. geum as a species without distinct subspecies and varieties. Selection of herbarium specimens. PAKISTAN. Prov. Frontiere, Gittidas au Katawai, 4100 m, 15 vii 1953, Schmidt 445 (BM). CHINA. Yunnan: Close to Yungning, 3140 m, 23 vi 1914, Handel-Mazzetti 2730 (WU); Likiang, Mt. Julung Shan, 6 ix 1914, Handel-Mazzetti 661 (WU), 4080 (WU); Yulung Shan, Mt. Nguluho, 3400 m, 7 vi 1915, Handel-Mazzetti 6671 (WU); Haba and Dugwantsum, SE of Chung tien, 4350 m, 23 vi 1915, Handel-Mazzetti 6908 (WU); eastern slopes of Mt. Dyinaloko, North Peak of the Likiang Snow Range, 13000 ft, vi 1923, Rock 9016 (W); Likiang Snow Range, 3780 m, 29 vii 1970, Feng 4070 (PE). Sichuan: Mts. Daliang Shan (territory Lolo), East of Ningyuen, South Dsilimba, 3275 m, 26 iv 1914, Handel-Mazzetti 1757 (WU); Fumadia and Wolo-ho, close to Yenyuen and Yungning, 3300 m, 15 vi 1915, Handel-Mazzetti 3065 (WU); South of Tschescha and North of Yungning, close to monasterium Muli, 3800 m, 24 vi 1915, Handel-Mazzetti 7179 (WU); close to Doko, and South of Yungning, monasterium Muli, 4350 m, 4 viii 1915, Handel-Mazzetti 7405 (WU); Sung-pan, 3000 m, 6 vii 1922, Fang 2624 (E); 9 vii 1922, Smith 2448 (BM); Sungpan-hsien, 8 viii 1928, Fang 4068 (E); Hsioeh-shan, 4300 m, 19 vii 1932, Smith 3885 (BM); Hsioeh-Shan, 25 viii 1936, W.T. Wang 660929 (PE). Shansi: Yang ling kie (Ou t’ai chan), 30 vi 1922, Licent 6556 (BM). Chindu: Xian, Qingshuine Xiang, between Madou and Yushu, 4300 m, 11 viii 1996, T.N. Ho et al. 1613 (BM). Qinghai: Tongde Xian, Longmuer Xiume, between Jungong (Gyumgo) and Hebei, North side of Hung He, 3650 m, 22 vii 1993, T.N. Ho et al. 156 (BM); Maqin (Magen) Xian, Dawu Xiang, along Gegu He, 3500 m, 31 vii 1993, T.N. REVISION OF ANEMONE SECT. HIMALAYICAE 75 Ho et al. 618 (BM). Xizang: Doshong La, Pemako, 3050 m, 25 x 1924, Kingdon-Ward 6262 (K); Radja and Yellow River Gorges, 13000 ft, vi 1926, Rock 14105 (E), vi 1926, Rock 14196 (BM, E); Tsarung, Solo-la, alpine belt, 14500 ft, vi 1932, Rock 22245 (BM, E). Xinjiang: Kangting (Tachienlu), 3100 m, 15 vii 1934, Smith 10462 (WU); Yulingkong, Gomba La, 3700 m, 22 vii 1934, Smith 10701 (BM). NEPAL. East Nepal: Largeng Khola, 14500 ft, 21 vi 1950, Lowndes 1038 (BM); Sangda: North of Tukucha, 19 viii 1954, Stainton et al. 7316 (BM); Basia: Banjang, 4700 m, 6 viii 1973, Einarsson et al. 2728 (BM); Close to Dhorpatan, North side of Uttar Ganga, 3800 m, 3 iv 1974, Vickery 554 (BM); Cha Lundgpa, 15300 ft, 29 vii 1977, Miehe 391 (E), 393 (E). INDIA. Uttaranchal: Kutti Yangti Valley, 15000 ft, 1 viii 1886, Duthie 5272 (WU). Assam: Tulung La, 1935, Kingdon-Ward 11681 (BM); Luguthang, 12000 ft, Kingdon-Ward 11622 (BM). 7. Anemone trullifolia Hook.f. & Thoms., Fl. Ind. 1: 22 (1855). – Anemone obtusiloba D.Don subsp. trullifolia (Hook.f. & Thoms.) Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Anemone trullifolia Hook.f. & Thoms. var. typica Finet & Gagnep., Bull. Soc. Bot. France 11: 61 (1904). – Type: East Himalaya, Hab. Sikkim. Regio alpina, 11–15000 ft, 14 vii 1849, Hooker s.n. (lecto K!, second step designation here by Ziman and Mosyakin; iso E!). Figs 7, 20. Anemone obtusiloba D.Don subsp. trullifolia (Hook.f. & Thoms.) Brühl var. spatulata Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Types: Brühl reported several type localities and specimens: ‘Sikkim (H. f. and others), near Thaling, Tsumtong (King’s collectors!); Chumbi: on Pit-ze-la, Pan-ka-la, Oey-gung-la; Phari (King’s collectors!); Bhutan; N. W. Himalaya?’, without numbers and data on herbarium (probably CAL). As we have not seen these plants, we are unable to designate a lectotype. F I G . 7. Anemone trullifolia Hook.f. & Thoms. var. trullifolia. A, basal leaf; B, flower parts (a, petaloid; b, stamen; c, carpel); C, achene (A–B, Handel-Mazzetti 7179, WU; C, Smith & Cave 1711, E). 76 S. N. ZIMAN ET AL. Anemone obtusiloba D.Don subsp. trullifolia (Hook.f. & Thoms.) Brühl var. rotundifolia Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: Sikkim, near Thaling, 13000 ft, and in other localities (Cunningham, G. Gammie, King’s collectors!). Like var. spatulata, we have not seen any of these plants. Anemone chumulangmaensis W.T.Wang, Acta Phytotax. Sin. 12: 171 (1974). – Type: China, Tibet, Mons Chumulangma, Kama, 4380 m, 15 vi 1959, Exped. ad Montem Chumulongma 269 (holo PE!). Leaves 4–10; petioles flat, 1–3(–5) cm 6 3–5 mm, villous or densely pubescent, rarely puberulent; blades 3-parted to 3-lobed, spathulate, rhombic-obovate to obovate, 2–7 6 1–5 cm, villous or densely pubescent, bases attenuate (sometimes cuneate), margins distally dentate, apices rounded (Fig. 7A). Scapes 2–7, 3–15(–20) cm long, villous or densely pubescent, 1–3-flowered. Involucral bracts 3-lobed or 3-dentate, sometimes entire, narrowly obovate or lanceolate, 1–2.5 cm long, hirsute. Bracteoles sometimes present, small, paired. Pedicels 1–5(–8) cm long, pubescent or puberulent. Petaloids 5–6(–15), monomorphic or sometimes dimorphic (if more than 6), white, yellow, pinkish, purplish or blue, elliptic-obovate, 5–12(–15) 6 4–8(–10) mm, densely or sparsely pubescent, hirsute or villous, anastomosing veins 1–2(–3) or sometimes absent (Fig. 7Ba). Stamens 1.8–3.4 mm long; filaments ovate-lanceolate, 0.5–0.7 mm wide (Fig. 7Bb). Carpels cylindric-ovoid, 2–4 mm long, villous or pubescent (Fig. 7Bc). Achenes ellipsoid-ovoid or fusiform, slightly compressed, 3.2–4.4 6 1.6–2.2 mm, without ribs, villous, hairs c.1 mm long; styles straight, 2–2.6 mm long (Fig. 7C). Distribution and habitat. SW China (NW Yunnan, S Sichuan, SW Gansu, S Quinghai, S Xizang), East Nepal, India (Sikkim), Bhutan (Fig. 20). In forests and alpine meadows; 2500–4800 m. Nomenclatural notes. Hooker and Thomson cited two collections (syntypes) in the protologue: ‘Sikkim, alt. 11–15,000 ft, Hooker f.; and Bhotan, Griffith’. Specimens of the respective collections are deposited at K and E. Lauener (1960) cited the E specimens (‘Sikkim: 3350–4570 m, J.D.Hooker’ and ‘Bhutan: Griffith 1718’) as syntypes, while Chaudhary (1988) cited as a type the first specimen mentioned by Lauener (Hooker’s specimen ‘Sikkim, 3350–4570 m, J.D.Hooker’, but from K), which can be regarded as an effective lectotypification. However, in that citation Chaudhary changed feet to metres and failed to indicate the collection date explicitly identifying the lectotype. Since there are several of Hooker’s specimens representing at least two collections (‘Hab. Sikkim. Regio alpina. Alt. 11–15000 ped., 14 vii 1849 and 5 ix 1849. Coll. J.D.Hooker. Herbarium Hookerianum 1867’), we have made the second step lectotypification of one of these. Taxonomic remarks. Anemone trullifolia was described from the eastern Himalaya as a taxon very close to A. obtusiloba (Hooker & Thomson, 1855). Brühl (1896) recognized Anemone trullifolia as a subspecies of A. obtusiloba and also made A. coelestina, a species which had recently been described by Franchet (1885), another REVISION OF ANEMONE SECT. HIMALAYICAE 77 subspecies of A. obtusiloba. However, he did not note any differences between these two taxa except for the yellow or blue petaloid colour and the distribution (Anemone trullifolia in the Himalaya and A. coelestina in SW China, Yunnan). Brühl noted the differences in basal leaf shape within subspecies trullifolia and on this basis he described three varieties: var. linearis (with mainly linear entire leaves), var. spatulata (with short-spathulate 3-lobed leaves) and var. rotundifolia (with suborbicular 3lobed leaves). Anemone trullifolia and A. coelestina were maintained by Ulbrich (1906) under the former name as one ‘species collectiva’. In contrast, Finet & Gagnepain (1904) proposed two varieties for Anemone trullifolia, var. coelestina and var. souliei, and Diels (1912) recognized var. campestris and var. holophylla within this taxon. Lauener (1960), however, again separated Anemone trullifolia and A. coelestina as distinct species and distinguished them by the shape of the basal leaves and the hairiness of the pistils. Furthermore, within Anemone trullifolia, he recognized var. trullifolia, var. linearis and var. holophylla, differentiated on the basis of leaf shape. Wang (1974) described three narrow endemics close to Anemone trullifolia: A. chumulangmaensis W.T.Wang, A. liangshanica W.T.Wang and A. lutienensis W.T.Wang. Anemone chumulangmaensis was lumped with A. trullifolia var. trullifolia by Wang himself and we agree with this. Furthermore, after examining the type materials of Anemone liangshanica and A. lutienensis we have reduced both to varieties of A. trullifolia (Figs 7, 8 and 9). Key to varieties 1a. Leaf blades 3-lobed, ovate or broadly lanceolate; scapes 1–3-flowered; petaloid anastomosing veins 1–3 _______________________________ 7a. var. trullifolia 1b. Leaf blades 3-parted or 3-cleft, spathulate or rhombic; flowers solitary; petaloid anastomosing veins absent __________________________________________ 2 2a. Leaf blades 3-parted, spathulate; petaloids 7–9 6 5–6 mm, villous __________ __________________________________________________ 7b. var. liangshanica 2b. Leaf blades 3-cleft, rhombic; petaloids 9–13 6 6–8 mm, sparsely pubescent __ ___________________________________________________ 7c. var. lutienensis 7a. var. trullifolia Leaf blades 3-lobed, ovate or broadly lanceolate. Scapes 1–3-flowered. Petaloids 5–7, anastomosing veins 1–3. Distribution and habitat. China (NW Yunnan, SW Sichuan, S Xizang), Nepal, India (Sikkim, Himachal Pradesh), Bhutan (Fig. 20). Along streams in forests and in alpine meadows; 2500–4500 m. Selection of herbarium specimens. CHINA. Yunnan: Long-Ki, 1894, Delavay s.n. (WU); v 1906, Bailey (LE); Likiang Range, 1933, McLawrens 307 (BM); Xiao huadinba, above Farm, v 1981, SBEC 747 (E). Sichuan: Mts. [near] Muli, vii 1930, Forrest 28434 (BM); Mountains round 78 S. N. ZIMAN ET AL. Muli, vii 1930, Forrest 28484 (BM); ibid., 1931, Forrest 30041 (BM), 30471 (BM), 30817 (BM). Xizang: Mons Chumulangma, Kama, 4380 m, 15 vi 1959, Exped. Mt. Chumulangma 269 (PE). NEPAL. Jansala, 3000 m, 17 v 1973, Einarsson et al. 53 (BM). INDIA. Sikkim: Regio alpina, 11–15000 ft, 14 vii 1849, Hooker s.n. (E, K); Llonok, 15000 ft, 22 ix 1909, Smith & Cave 1711 (E); Zemu and Lhonakh Valleys, Yeumtang, 12000 ft, 13 ix 1947, Cave 167 (E); Himalaya: Jolinka, 14500 ft, vii 1931, Bentham s.n. (BM). Himachal Pradesh: Chamba, below Sathraundi, Bera Nullah, 10000 ft, 8 vi 1981, Sayers 3689 (BM). BHUTAN. 1867, Griffith 1718 (E, K); Mem La, Paro Valley, 13000 ft, 15 v 1949, Ludlow & Sheriff 16247 (BM, E); Mangde Chu, 14000 ft, 12 vi 1966, Bowes-Lyon 3440 (BM); Waitang, Isampa, 15000 ft, 2 vi 1949, Ludlow et al. 19202 (BM); Cho La, 12500 ft, 4 vii 1949, Ludlow et al. 20800 (BM). 7b. var. liangshanica (W.T.Wang) Ziman & B.E.Dutton, Fl. China 6: 324 (2001). – Anemone liangshanica W.T.Wang, Acta Phytotax. Sin. 12: 170 (1974). – Type: China, Sichuan, Lepo, Liangshan, Hwangmaokeng, 2800 m, 19 vi 1959, C.C. Hu et al. 761 (holo PE!). Fig. 8. Leaf blades 3-parted, spathulate. Flowers solitary. Petaloids white or purplish, 7–9 6 5–6 mm, villous; anastomosing veins absent. Distribution and habitat. China. Endemic to western Sichuan, Lepo (5 Leibo Xian) (Fig. 20). In alpine meadows; 3000–3600 m. Known only from the type collection and a few specimens collected in the same area and cited in the protologue. F I G . 8. Anemone trullifolia var. liangshanica (W.T.Wang) Ziman & B.E.Dutton. A, leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). REVISION OF ANEMONE SECT. HIMALAYICAE 79 F I G . 9. Anemone trullifolia var. lutienensis (W.T.Wang) Ziman & B.E.Dutton. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). 7c. var. lutienensis (W.T.Wang) Ziman & B.E.Dutton, Fl. China 6: 324 (2001). – Anemone lutienensis W.T.Wang, Acta Phytotax. Sin. 12: 170 (1974). – Type: China, Yunnan, Lichiang, Lutien, 28 v 1939, R.C. Ching 20548 (holo PE!). Fig. 9. Leaf blades 3-cleft, rhombic. Petaloids white or blue, 9–13 6 6–8 mm, sparsely pubescent; anastomosing veins absent. Distribution and habitat. China. Endemic to NW Yunnan (Lichiang, Lutien) (Fig. 20). In alpine meadows; 4000 m. Known only from the type collection and a few specimens collected in the same area and cited in the protologue. 8. Anemone coelestina Franch., Bull. Soc. Bot. France 32: 4 (1885). – Anemone obtusiloba D.Don subsp. coelestina (Franch.) Brühl, Ann. Roy. Bot. Gard. Calcutta 5: 78 (1896). – Anemone trullifolia Hook.f. & Thoms. var. coelestina (Franch.) Finet & Gagnep., Bull. Soc. Bot. France 51: 61 (1904). – Type: China, Prov. Yunnan, sommet du mont Hee-chan-men, supra Lan-Kong, 2 vi 1884, Delavay 3 (lecto P!, designated here by Ziman and Mosyakin). Figs 10, 20. Anemone bonatiana H.Lév., Feddes Repert. Spec. Nov. 7: 98 (1909). – Type: China, Yunnan: Lao Kouy-Chan, pres My Le, 1906, Herb. Bonati, Ngueou 607 (holo E!). Leaves 5–10; petioles 1–4 cm 6 8–10 mm, villous or densely pubescent; blades subtrilobed to undivided, oblong-linear to oblanceolate, 2–8 6 1–3 cm, villous, bases cuneate-attenuate; margins obtusely or acutely dentate (Fig. 10A). Scapes 2–8, 3–10 cm long, densely pubescent; flowers solitary. Involucral bracts undivided, linearlanceolate, 1–3 cm long, villous. Pedicels 1–3 cm long, villous. Petaloids 5–6, monomorphic, reddish blue or bluish white, yellow, reddish orange, reddish violet or 80 S. N. ZIMAN ET AL. F I G . 10. Anemone coelestina Franch. var. coelestina. A, basal leaf; B, flower parts (a, petaloid; b, stamen; c, carpel); C, achene (A, Delavay 1887, WU; B, Handel-Mazzetti 2437, WU; C, Forrest 1922, LE). bluish violet, broadly elliptic, 8–14 6 7–12 mm, densely pubescent, basal veins 3–5, anastomosing veins absent (sometimes solitary) (Fig. 10Ba). Stamens 2.5–3 mm long; filaments linear (Fig. 10Bb). Staminodes absent or occasionally present. Carpels ovoid, 3–4 mm long, villous; styles straight (Fig. 10Bc). Achenes ovoid, 3–4.5 6 2 mm, without ribs, villous, hairs c.1 mm long; styles substraight, c.2.5 mm long (Fig. 10C). Distribution and habitat. China (Yunnan, Sichuan, E and S Xizang), Nepal, Bhutan, North India (Sikkim, Assam) (Fig. 20). In Rhododendron forests and scrub, on grassy slopes, streamsides, alpine meadows; 2500–5000 m. Taxonomic remarks. Anemone coelestina was described by Franchet (1885) from Yunnan as a species close to A. trullifolia but differing from it by the long-ovate subtrilobed basal leaves, and 5 blue or white petaloids. Although Ulbrich (1906) and Handel-Mazzetti (1931, 1939) recognized Anemone coelestina as a species, other authors (e.g. Brühl, 1896; Finet & Gagnepain, 1904; Lauener, 1960; Wang, 1974, 1980; Chaudhary, 1988) regarded this taxon as a subspecies or variety of A. trullifolia, or included it in A. trullifolia as a synonym. After a comparative examination of many specimens we note that Anemone trullifolia and A. coelestina share important characters but that A. coelestina differs from A. trullifolia in having mostly undivided, oblong-linear to oblanceolate basal leaf blades with wider petioles (3–5 and 8–10 mm), undivided bracts, solitary flowers, fewer monomorphic petaloids (5–6 only as compared with 5–15 in A. trullifolia) without anastomosing veins, and linear filaments. Consequently, we recognize Anemone coelestina as a distinct species. Brühl (1896) described Anemone obtusiloba subsp. trullifolia var. linearis on the basis of its linear entire basal leaf blades. Handel-Mazzetti (1939) also recognized this variety and published it as Anemone trullifolia var. linearis (Brühl) Hand.-Mazz. This variety was also recognized by Lauener (1960) who assumed its proximity to Anemone coelestina. As a result of our examination of the type specimens of var. REVISION OF ANEMONE SECT. HIMALAYICAE 81 linearis (Pratt 493 in K and BM) and var. souliei (Soulie 7 in K) we note their similarity in the basal leaves with short wide petioles and undivided oblong-linear blades, undivided bracts, solitary flowers, and petaloids without anastomosing veins. Accordingly, we recognize only one variety (var. linearis). However, we believe this variety to be closer to Anemone coelestina than to A. trullifolia and include it under A. coelestina. Variety holophylla Diels was also described under Anemone trullifolia but was transferred to A. coelestina (Handel-Mazzetti, 1939). Nevertheless, Lauener (1960) and Wang (1974, 1980) included var. holophylla in Anemone trullifolia. However, as a result of our examination of the type material of var. holophylla (Forrest 2166 in BM, E, K) with its lanceolate, almost entire apically toothed basal leaf blades, and blue-violet petaloids without anastomosing veins, we regard these plants as closer to Anemone coelestina than to A. trullifolia, and thus transfer var. holophylla from A. trullifolia to A. coelestina. Key to varieties 1a. Leaf blades subtrilobed, ovate-oblong; petaloids bluish-white, anastomosing veins solitary; staminodes absent ______________________ 8a. var. coelestina 1b. Leaf blades undivided or rarely obscurely 3-lobed, linear to oblanceolate; petaloids bluish-violet, anastomosing veins absent; staminodes absent or sometimes present _________________________________________________ 2 2a. Leaf blades linear, margins acutely dentate; staminodes absent 8b. var. linearis 2b. Leaf blades oblanceolate, margins obtusely dentate; staminodes sometimes present ____________________________________________ 8c. var. holophylla 8a. var. coelestina Leaf blades subtrilobed, ovate-oblong. Petaloids bluish-white, anastomosing veins solitary; staminodes absent. Distribution and habitat. China (Yunnan, Sichuan) (Fig. 20). In alpine meadows and bushes; 3500–4800 m. Selection of herbarium specimens. CHINA. Yunnan: sommet du mont Hee-chan-men, environs de Ta-li, 11 vii 1883, Delavay 3 (P, WU); Mts. Koua-la-po, prei Ho-Kin, env. De Bali, 26 v 1884, Delavay 1853 (P); Mt. Hee Chang men, above Lang-Kong, 2 vi 1884, Delavay 48 (WU); 30 x 1884, Delavay 1034 (WU); Lang-Kong, 16 iv 1887, Delavay s.n. (LE, PE, WU); Lao Kouy-Chan, pres My Le, 1906, Ngueou 607 (E); Yangtze Watershed, slopes of Likiang Snow Range, v 1910, Forrest 5640 (BM, K); Lichiang Range, 10–11000 ft, vi 1913, Forrest 10135 (BM); Xiao Huadinba, above farm, 18 v 1981, SBEC 747 (E). SW Sichuan: Mts. Liukuliandse, between Yenyuen and Castle Kwapi, 18 v 1914, Handel-Mazzetti 2437 (WU); Yungning, 16 vi 1914, Handel-Mazzetti 3085 (WU); Mts. Kopati, Djago and Muli, vi 1928, Rock 16162 (LE); Mts. Mitsuga, West of Muli Gomba, 4875 m, vi 1928, Rock 16206 (E, K). 8b. var. linearis (Brühl) Ziman & B.E.Dutton, Fl. China 6: 325 (2001). – Anemone obtusiloba subsp. trullifolia (Hook.f. & Thoms.) Brühl var. linearis Brühl, Ann. 82 S. N. ZIMAN ET AL. F I G . 11. Anemone coelestina var. linearis (Brühl) Ziman & B.E.Dutton. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). Roy. Bot. Gard. Calcutta 5: 77 (1896). – Anemone trullifolia var. linearis (Brühl) Hand.-Mazz., Acta Horti Gothob. 13: 178 (1939). – Type: West Szechuan and Tibetan Frontier, chiefly near Ta chien Lu, 9–13000 ft, xii 1890, Pratt 493 (holo BM!). Fig. 11. Anemone souliei Franch., Bull. Soc. Bot. France 32: 4 (1885). – Anemone trullifolia Hook.f. & Thoms. var. souliei (Franch.) Finet & Gagnep., Bull. Soc. Bot. France 51: 62 (1904). – Type: Chine. Prov. Se-tchuen, Tongolo, pres Ta-tsien-lou, vii–viii 1891, Soulie 7 (holo P!). Leaf blades mostly undivided (rarely obscurely 3-lobed), linear, acutely dentate, 8–15 6 2–3 cm. Petaloid anastomosing veins absent. Staminodes absent. Distribution and habitat. China (NW Yunnan, W and S Sichuan, Xizang), Bhutan (Fig. 20). In Rhododendron forests and bushes, and in alpine meadows; 2800–4900 m. Selection of herbarium specimens. CHINA. Yunnan: 1930, Forrest 28855 (E); vii 1935, W.T. Wang (PE). Sichuan: Tibetan Frontier, chiefly near Ta chien Lu, 9–13000 ft, xii 1890, Pratt 493 (BM); Tongolo, pres Ta-tsien-lou, vii–viii 1891, Soulie 7 (P!); Hongyuan Co., Rangkouxing, 1 viii 1982, Zhao Qing Sheng 212 (E); Chazhenliangzi, 3700 m, 31 vii 1989, Zhao Qing Sheng 14 (E). Xizang: Pasho Distr., Kham, Valley of Du Chu, 14000 ft, 13 vii 1936, Hanbury-Tracy 30 (BM). BHUTAN. Chojo Dzong, 13500 ft, 30 vi 1949, Ludlow et al. 16687 (K). 8c. var. holophylla (Diels ex H.F.Comber) Ziman & B.E.Dutton, Fl. China 6: 324 (2001). – Anemone trullifolia Hook.f. & Thoms. var. holophylla Diels, Notes Roy. Bot. Gard. Edinburgh 5: 263 (1912). – Type: China: Yunnan, Eastern flank of the Lichiang Range, 3050–3350 m, v 1906, Forrest 2166 (lecto E!; iso BM, K!). Fig. 12. Anemone coelestina Franch. var. polygyna H.F.Comber and f. holophylla (Diels) H.F.Comber, Notes Roy. Bot. Gard. Edinburgh 27: 226 (1934). Leaf blades oblanceolate, margins obtusely dentate. Petaloid anastomosing veins absent. Staminodes sometimes present. REVISION OF ANEMONE SECT. HIMALAYICAE 83 F I G . 12. Anemone coelestina var. holophylla (Diels ex H.F.Comber) Ziman & B.E.Dutton. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). Distribution and habitat. China (NW Yunnan, Sichuan, Xizang), Nepal, North India (Assam) (Fig. 20). In Rhododendron forests and on grassy slopes; 2500–4900 m. Selection of herbarium specimens. CHINA. Yunnan: Likiang Range, v 1906, Forrest 2166 (BM, E); ibid., vi 1906, Forrest 2226 (BM); ibid., vii 1906, Forrest 2634 (E); Distr. Chung-tien, 13000 ft, Mts. West of Hsiao Chung-tien, v 1932, Rock 24664 (BM, K); Pei Ma Shan, Mekong-Jangtze Divide, SW of Atuntze, vi 1932, Rock 22764 (BM, K); Haba Shan, North of Yangtze loop, third Peak of Likiang Snow Range, 1932, Rock 24793 (BM, K). Sichuan: Hongyuan Co., Rangkouxiang, 3740 m, 31 vii 1989, Zhao Qing-Sheng 212 (BM); Hongyuan Co., Chazhen liangzi, 3700 m, 31 vii 1989, Zhao Qing-Sheng 14 (BM). Xizang: 1906, Forrest 362 (K); TafsiculiDaevo, 25 vi 1914, Shimprift 1797 (WU); Hills of Lhasa, 14000 ft, 26 vii 1943, Ludlow & Sheriff 9701 (BM). NEPAL. East Nepal: Tamur Valley: Mewa Khola, Topke Gola, 13500 ft, 14 v 1956, Stainton 262 (BM, E). INDIA. Assam: Mago, 11700 ft, 1935, Kingdon-Ward (BM); Chumbi and Phari, 1878, Dunboo s.n. (BM). 9. Anemone subindivisa W.T.Wang, Acta Phytotax. Sin. 12: 173 (1974). – Type: China, Szechuan, Muli, Mons Misuga, 3700–4000 m, 26 v 1937, T.T. Yu 5756 (holo PE!). Figs 13, 21. Leaves 3–7; petioles 2.5–4 cm 6 5–10 mm, villous; blades undivided or obscurely 3lobed, broadly ovate or orbicular-ovate, 3–4 6 2–3 cm, villous; bases roundedtruncate; margins obtusely dentate, apices rounded (Fig. 13A). Scapes 2–5, 3–10 cm long, villous; flowers solitary. Involucral bracts entire or subequally 3-dentate, lanceolate, 1–2 cm long, with obtuse apices, villous. Pedicels 0.5–1 cm long, villous. Petaloids 5–6, white, broadly elliptic, 8–12 6 5–8 mm, sparsely puberulent, basal veins 3–5, without anastomoses. Stamens 3–5 mm long; filaments lanceolate (Fig. 13Ba). Carpels 3–3.5 mm long, densely pubescent (Fig. 13Bb). Distribution and habitat. China. Endemic to SW Sichuan: Muli (Fig. 21). In Abies and Pinus forests and on grassy slopes; 2500–4000 m. 84 S. N. ZIMAN ET AL. F I G . 13. Anemone subindivisa W.T.Wang. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). Taxonomic remarks. Wang (1974) described this species from China (Sichuan) as very close to Anemone trullifolia but differing from it by the rounded-truncate basal leaf bases and mainly undivided blades. Our data confirm that Anemone subindivisa is a member of series Trullifoliae but we suggest it is closer to A. yulongshanica, differing from it by its undivided basal leaf blades, wider petioles, entire (sometimes 3-dentate) lanceolate involucral bracts, and smaller petaloids. We accept this taxon as a species but we note the necessity for further detailed study, especially of its mature achenes. Known only from the type collection and a few specimens collected in the same area and cited in the protologue. 10. Anemone yulongshanica W.T.Wang, Bull. Bot. Res. NE Forest Univ. 16: 159 (1996). – Type: China, Yunnan, Likiang Snow Range, 1937, T.T. Yu 15595 (holo PE!). Figs 14, 21. Leaves 5–9; petioles 4–13 cm 6 2–3 mm, pubescent or densely villous; blades 3parted or 3-lobed, pentagonal or broadly ovate, 1–5.5 6 1–6 cm, herbaceous or F I G . 14. Anemone yulongshanica W.T.Wang var. yulongshanica. A, basal leaf; B, achene (A, Flora of China, 2001; B, T.T. Yu 1559, PE). REVISION OF ANEMONE SECT. HIMALAYICAE 85 papery, pubescent or villous, bases truncate; apical lobes 3-lobed or undivided, rhombic-ovate or broadly rhombic, sparsely obtusely dentate; lateral lobes unequally 2-lobed, obliquely broadly cuneate or flabellate (Fig. 14A). Scapes 2–9, 3–15(–30) cm, villous or pubescent; flowers solitary. Involucral bracts unequally 3-parted or 3-lobed, narrowly rhombic or oblong-lanceolate, 0.7–3 cm long, pubescent. Pedicels 1–7 cm long, puberulent. Petaloids 5–6, white, yellowish or bluish, narrowly obovate or elliptic, 8–16 6 8–10 mm, sparsely appressed pubescent, basal veins 3, without anastomosing veins. Stamens 2–3 mm long; filaments lanceolate. Carpels 2.5–3.5 mm long, linear-lanceolate or narrowly ovoid, sericeous or pubescent. Achenes ovoid, 2.8– 3.7 6 1.8–2.2 mm, without ribs, villous, hairs c.1 mm long; styles 1.4–1.6 mm long, straight or slightly curved (Fig. 14B). Distribution and habitat. China (NW Yunnan, SW Sichuan) (Fig. 21). In Abies forests, on rocks and grassy slopes; 2600–3900 m. Taxonomic remarks. Diels (1912) noted that certain plants referable to Anemone trullifolia had basal leaves with truncate bases. However, Comber (1934) later regarded these variants as belonging to Anemone coelestina, not A. trullifolia. Moreover, Comber described them as A. coelestina var. truncata and var. polygyna (the latter with more carpels), and he considered that both varieties were close to A. coelestina var. holophylla. Eventually Wang (1996) described Anemone yulongshanica on the basis of a specimen of A. obtusiloba from Yunnan collected by T. T. Yu. It differs from Anemone obtusiloba s.str. mainly by the apical lobes of the basal leaves (3-lobed or undivided). According to our data Anemone yulongshanica has certain affinities to series Obtusilobae but should be regarded as a member of series Trullifoliae. It differs from Anemone trullifolia by the basally truncate or subcordate (not attenuate) basal leaf blades, narrower filaments, and smaller ovoid achenes with shorter styles. Wang (1974) divided this species into two varieties, with var. yulongshanica differing from var. truncata mostly by its larger basal leaves and bracts. Wang et al. (2001) maintained these varieties and both are recognized in this treatment. Key to varieties 1a. Basal leaf petioles 6–13 cm long, pubescent; leaf blades 3–5.5 6 4–6 cm, basally subcordate, apical lobes 3-cleft; involucral bracts 1.5–3 cm long; carpels linearlanceolate, sericeous _____________________________ 10a. var. yulongshanica 1b. Basal leaf petioles 4–5 cm long, densely villous; leaf blades 1–3.5 6 1–4 cm, basally truncate, apical lobes undivided; involucral bracts 0.7–2 cm long; carpels narrowly ovoid, pubescent ____________________________ 10b. var. truncata 10a. var. yulongshanica Basal leaf petioles 6–13 cm long, pubescent; leaf blades 3–5.5 6 4–6 cm, basally subcordate, apical lobes 3-cleft (Fig. 14A). Involucral bracts 1.5–3 cm long. Carpels linear-lanceolate, sericeous (Fig. 14B). 86 S. N. ZIMAN ET AL. F I G . 15. Anemone yulongshanica var. truncata (H.F.Comber) W.T.Wang. A, basal leaf; B, flower parts (a, stamen; b, carpel) (A–B, Flora of China, 2001). Distribution and habitat. China (Yunnan). Endemic to the Lichiang Snow Range. Known only from the type collection. 10b. var. truncata (H.F.Comber) W.T.Wang, Bull. Bot. Res. NE Forest Univ. 16: 159 (1996). – Anemone coelestina Franch. var. truncata H.F.Comber, Notes Roy. Bot. Gard. Edinburgh 18: 226 (1934). – Type: China, Yunnan, East flank of Lichiang Range, 11000 ft, v 1910, Forrest 5640 (holo E!; iso K!, P!). Fig. 15. Basal leaf petioles 4–5 cm long, densely villous; leaf blades 1–3.5 6 1–4 cm, basally truncate, apical lobes undivided (Fig. 15A). Involucral bracts 0.7–2 cm long. Carpels narrowly ovoid, pubescent (Fig. 15Bb). Distribution. China (Yunnan). Endemic to the Lichiang Snow Range. Known only from the type collection. 11. Anemone rupestris Wall. ex Hook.f. & Thoms., Fl. Ind. 1: 21 (1855). – Type: Sikkim, 4570 m, Hooker s.n. (lecto K!; iso E!). Figs 16, 21. Anemone obtusiloba D.Don var. lobata Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: Sikkim: on the Pa-Tang-La; Chumbi, at Ley-rong, King’s collector (iso CAL). Anemone rupestris Wall. ex Hook.f. & Thoms. var. wallichii Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: Nepal, Gossain Than, Wallich 4696 (holo CAL; iso K). Anemone obtusiloba D.Don subsp. saxicola Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Anemone saxicola (Brühl) Tamura & Kitamura in Kihara, Fauna Fl. Nepal. Himal. 125 (1955). – Type: Kashmir, Falconer 28 (holo CAL). Anemone obtusiloba D.Don subsp. omalocarpella Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: Nipal, Scully (holo CAL). Anemone bhutanica Tamura, Acta Phytotax. Geobot. 19: 75 (1962). – Type: Bhutan, Shukuna Mountain, 1 viii 1958, Nakao 101 (holo KYO). REVISION OF ANEMONE SECT. HIMALAYICAE 87 F I G . 16. Anemone rupestris Wall. ex Hook.f. & Thoms. subsp. rupestris. A, basal leaf; B, flower parts (a, petaloid; b, stamens; c, carpel); C, achene (A–B, Ludlow & Sheriff 20352, E; C, Handel-Mazzetti 1914, WU). Basal leaves 4–7; petioles 3–10 6 1–2 mm, sparsely puberulent or subglabrous; blades twice 3-sect, ovate, 1–5 6 1–6 cm, sparsely puberulent or glabrous, bases subcordate; apical leaflets with petiolules 5–10 mm long, 3-sect or 3-parted, broadly rhombic, secondary leaflets with petiolules 1–2 mm long, ultimate lobules narrowly ovate or linear-lanceolate; lateral leaflets on petiolules 2–5 mm or subsessile, 3parted or 3-lobed, subreniform (Fig. 16A). Scapes 2–6, 3–20 cm long, sparsely puberulent or subglabrous, 1–3-flowered. Involucral bracts undivided or 2- or 3lobed, ovate-oblong, cuneate-obovate or rhombic, 1–3 cm long. Pedicels 1–6 cm long, puberulent or glabrous. Petaloids 5–6(–8), white, blue or purplish or reddishwhite, oblong-elliptic or obovate, mainly dimorphic (outer and inner ones differ in size and colour), 5–10(–14) 6 3–6(–8) mm, subglabrous, basal veins, anastomosing veins absent (Fig. 16Ba). Stamens 3.5–4.3 mm long; filaments lanceolate or oblongelliptic (Fig. 16Bb). Carpels ovoid, compressed, 2.5–3.3 mm long, subglabrous; styles down-curved (Fig. 16Bc). Achenes broadly ellipsoid, compressed, 2.5–4 6 1.6–2 mm, subglabrous; ribs distinct; styles curved, sometimes basally bent, 1.2–1.8 mm long (Fig. 16C). Distribution and habitat. China (Yunnan, Sichuan, S Xizang, Sikang), Nepal, Bhutan, India (Sikkim, Assam, Kashmir) (Fig. 21). In Rhododendron bushes, alpine meadows and on rocky outcrops; 2700–4800 m. Taxonomic remarks. Anemone rupestris was described (Hooker & Thomson, 1855) as a taxon very close to A. obtusiloba but with smaller and more slender 1–3-flowered stems, with narrower segments to the more dissected leaves and is less hairy. Later, Hooker (1872) assumed this taxon to be a part of Anemone obtusiloba and not a 88 S. N. ZIMAN ET AL. distinct species. Maximowicz (1890) described Anemone gelida on the basis of its twice 3-sect basal leaves, 3-lobed or entire bracts, 6–8 glabrous petaloid sepals and compressed pilose achenes with short straight styles. Shortly afterwards Brühl (1896) proposed to reduce Anemone gelida to a subspecies of A. rupestris due to the overall similarity except for the number and colour of petaloids and the pubescent receptacle. He recognized within Anemone rupestris three varieties: var. lobata, var. wallichii and var. pusilla. More recently, other authors (Ulbrich, 1906; Lauener, 1960) included Anemone gelida as a subspecies of A. rupestris. Thus, according to Lauener, Anemone rupestris includes two subspecies, subsp. rupestris and subsp. gelida, the latter with two varieties, var. gelida and var. wallichii. On the other hand, Wang (1974, 1980) and Chaudhary (1988) both regarded Anemone rupestris as consisting of three subspecies: subsp. rupestris, subsp. gelida and subsp. polycarpa. After a comparative study of Anemone rupestris and A. gelida, we confirm their similarity, but differentiate A. gelida by its smaller basal leaves, scapes, petaloids, and strictly solitary flowers. Consequently, following Wang et al. (2001), we accept Anemone rupestris and A. gelida as subspecies. In contrast, the characters listed by Brühl (1896) to separate three varieties appear to be too variable and not sufficiently distinct to warrant the recognition of the varieties. The same applies to the varieties of Anemone rupestris subsp. gelida (Lauener, 1960). Recently we included Anemone obtusiloba D.Don subsp. omalocarpella Brühl as a synonym of A. polycarpa (Wang et al., 2001), but here we reject this opinion and accept it as a part of A. rupestris due to its ‘nearly flat, margined, quite glabrous carpels’ (Brühl, 1896: 78). Key to subspecies 1a. Leaf blades 2–5 6 1–6 cm; scapes 10–20 cm long, 1–3-flowered; petaloids 5–6, 8–10(–14) 6 4–6(–8) mm; flowers and achenes subglabrous ________________ __________________________________________________ 11a. subsp. rupestris 1b. Leaf blades 1–1.5 6 1–2 cm; scapes 3–12 cm long, 1-flowered; petaloids 5–8, 5–7 6 3–4 mm; flowers and achenes glabrous _______________ 11b. subsp. gelida 11a. subsp. rupestris Leaf blades 2–5 6 1–6 cm. Scapes 10–20 cm long, 1–3-flowered. Petaloids 5–6, 8– 10(–14) 6 4–6(–8) mm. Flowers and achenes subglabrous (Fig. 16). Distribution and habitat. China (Yunnan, Sichuan, Xizang), Nepal, Bhutan, North India (Fig. 21). On slopes and along streams; 2500–3000 m. Selection of herbarium specimens. CHINA. Yunnan: Pei-ma shan, ix 1932, Rock 23390 (GH); Yangtze Watershed, western slopes of Likiang Snow Range, 6 vi 1922, Rock 4219 (BM). Sichuan: Yunghing, 24 vi 1914, Handel-Mazzetti s.n. (WU). SE Xizang: Tsarung, northern slopes of Mt. Kenichunpo, North of Sikitung, Upper Salween River, v 1932, Rock 22179 (E, K); Shinden-Gompa, Nagong, 25 vii 1933, Kingdon-Ward 10641 (BM); Kongbo Prov., Lusha REVISION OF ANEMONE SECT. HIMALAYICAE 89 Chu, 12000 ft, 9 vi 1938, Ludlow & Sheriff 4723 (BM); 60 km North of Lhasa, Reting, 14000 ft, 27 vi 1944, Ludlow & Sheriff 11069 (BM). NEPAL. Gosainsthan, Gopain Shar, Wallich 4696 (K); Arun Valley, Chhoyang Khola, West of Num, 13000 ft, 22 vi 1956, Stainton 739 (E). BHUTAN. Me La, 9 vi 1949, Ludlow & Sheriff 20352 (E); Kangla Karcha La, Po Chu Drainage, 16000 ft, 20 vi 1949, Ludlow & Sheriff 16596 (E). INDIA. Sikkim: Hills to North and West of Dzongri camp, 4200 m, 23 vi 1983, Starling et al. 82 (E); West Distr., between Dzongri and Black Kabru (Kabur), 4050 m, 17 vii 1992, Long et al. 450 (E); Kashmir, 15000 ft, alpine belt, 1864, Falconer s.n. (K); Sikkim, 4570 m, Hooker s.n. (E, K); Sikkim, Himalaya, 15000 ft, 1832, Wallich 4676 (K), 5479 (K); Kashmir ad Pir Panjal, Jacquemont s.n. (K). 11b. subsp. gelida (Maxim.) Lauener, Notes Roy. Bot. Gard. Edinburgh 23: 199 (1960). – Anemone gelida Maxim., Acta Hort. Petropol. 11: 21 (1890). – Type: China, S Sichuan, Valle Kserntso, 11 viii 1885, Potanin s.n. (holo LE!; iso K!). Fig. 17. Anemone obtusiloba D.Don var. pusilla Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: Sikkim, Sei-Cho-La, W of Jongri; Kapur below Kinchinjanga; Pandimchu, 15000 ft, King’s collectors s.n. (iso CAL!). Leaf blades 1–1.5 6 1–2 cm. Scapes 3–12 cm long, 1-flowered. Petaloids 5–8, 5–7 6 3–4 mm. Flowers and achenes glabrous. Distribution and habitat. China (Yunnan, Sichuan, Xizang), Bhutan, Nepal, India (Fig. 21). In alpine meadows; 4800–5000 m. Selection of herbarium specimens. CHINA. NW Yunnan: Lu-djiang (Salween) and Djiou-djiang (Irrawadi), close to Tschiangschel, 4075 m, 4 vii 1916, Handel-Mazzetti 9262 (WU), 2730 (WU); Yangtze Watershed, western slopes of Likiang Snow Range, 6 vi 1922, Rock 4219 (E); F I G . 17. Anemone rupestris subsp. gelida (Maxim.) Lauener. A, basal leaf; B, flower parts (a, petaloid; b, stamen; c, carpel) (A, Handel-Mazzetti 1658, WU; B, Potanin 1885, LE). 90 S. N. ZIMAN ET AL. SW of Wei-Hsi, Mekong-Salween Divide, 4300 m, v 1928, Rock 16965 (K). SW Sichuan: Daliang-schan (territory Lolo), East of Nirgyuen, close Lauba, 2700 m, 25 iv 1914, HandelMazzetti 1658 (WU). BHUTAN. Shimitang, Bumthang Co., 10500 ft, 23 v 1949, Ludlow & Sheriff 18940 (BM). NEPAL. Dhimti Shir, 14000 ft, 1930, Lall Dhwoj 487 (BM); Lete: South of Tukucha, Kali Gandaki, 12500 ft, 7 vi 1954, Stainton et al. 1012 (BM); Bhurungdi Khola, 10000 ft, 15 vi 1954, Stainton et al. 5798 (BM); Rambrong, Lamjung Himal, 13500 ft, 5 vii 1954, Stainton et al. 6154 (BM); Inkhu Khola, 14000 ft, 17 vi 1964, Bowes-Lyon 2044 (BM); Dudh Kosi: Chaunrikharka, 13000 ft, 20 vi 1964, Bowes-Lyon 2087 (BM); Likhu Khola: East of Chaukharma, 12500 ft, McCosh 250 (BM). INDIA. Sikkim: Hills to North and West Dzongri camp site, 4200 m, 23 vi 1983, Starling et al. 82 (E). The remaining two species, Anemone imbricata Maxim. and A. fuscopurpurea H.Hara, described and discussed below, are rather isolated members of Anemone subgen. Omalocarpus but have sometimes been considered to belong to Anemone sect. Himalayicae. Until more information is available we prefer to exclude them from this section. 12. Anemone imbricata Maxim., Fl. Tangut. 8 (1889). – Anemone obtusiloba D.Don subsp. imbricata (Maxim.) Brühl, Ann. Bot. Gard. Calcutta 5: 78 (1896). – Type: China, S Tibet, Jangtze, 1884, Przewalski (E!, LE!). Leaves 4–7; petioles basally vaginate, 3–5 6 0.1–0.2 cm, villous; blades appear pinnatifid but are in principle 3-sect, elliptic-ovate, 2–3 6 1.2–2 cm, villous; bases cordate; central segments on petiolules 5–10 mm long, 3-sect or 3-parted; lateral segments shortly petiolulate or subsessile, unequally 3-parted; segments and lobes imbricate. Scapes 3–5(–10), 5–12 cm long, villous; cymes 1–2-flowered. Involucral bracts with 3-parted or 3-lobed blades, 1–2 cm long, villous. Pedicels 1–3.5 cm long, villous. Petaloids 5–7(–9), obovate-oblong or obovate, with acute apices, purple or blackish-purple (rarely whitish), sometimes dimorphic, 8–13 6 4–8 cm, glabrous or sparsely puberulent only along central veins; basal veins 3, without anastomoses. Stamens 3–5 mm long; filaments linear, slightly dilated; anthers ellipsoid. Carpels broadly ovate, 2.5–3 mm long, glabrous. Achenes broadly elliptic, 4.5–6 6 3–4 mm, glabrous, compressed, with ribs 0.5–0.7 mm wide; styles basally bent, c.1 mm long. Distribution and habitat. China (W Sichuan, SW Gansu, Qinghai, E Xinjang, Xizang). In bushes and on grassy slopes; 3200–5300 m. Taxonomic remarks. Maximowicz (1889), who described Anemone imbricata, noted its pinnatifid basal leaf blades and long-vaginate dilated petioles, 3-lobed involucral bracts, solitary flowers with 5–9 violet ‘sepals’, dilated stamens, and compressed glabrous achenes with conic-cylindric styles. Among students of the Anemone obtusiloba complex only Brühl (1896) lumped A. imbricata with A. obtusiloba, whereas Ulbrich (1906), Lauener (1960) and Wang (1974, 1980, etc.) recognized A. REVISION OF ANEMONE SECT. HIMALAYICAE 91 imbricata as a distinct species. Most authors regarded Anemone imbricata as a member of the A. obtusiloba complex but Starodubtsev (1991) made this species the type of a new section: Anemonastrum sect. Imbricata. As a result of our comparative study of Anemone imbricata and examination of the Przewalski type material (in LE and E) we suggest that this taxon does not belong to sections Himalayicae (Anemone obtusiloba complex) or Omalocarpus but should be placed in a monotypic section Imbricata of Anemone subgen. Omalocarpus. Selection of herbarium specimens. CHINA. Yunnan: Kunming, Dijing Pref., Degin Co., Bai Ma Shan, N of W Pass, 4530 m, vi 1993, Alden et al. 818 (E). SW Sichuan: Kulu Mts., Muli Territory, 15000 ft, Rock 23953 (BM), 23955 (BM); Djesi-La and Djesi-Longba, S of Tatsienlu, vi 1929, Forrest 12704 (K). Qinghai: Yushu Xian, between Yushu and Gyairong, 9 viii 1996, T.N. Ho et al. 2059 (GH). Xinjang: Kangting (Tanchienlu), Tapaoshan, 4700 m, 22 viii 1934, Smith 11512 (BM); Kuen-Lun, 24 vi 1894, Robowski 362 (LE). Xizang: Jangtze, 1884, Przewalski (E, LE); Russkoe Lake, 2 vi 1901, Ladygin 87 (LE); between Radja and Jupar, Wajo la, vi 1926, Rock 14146 (LE); Totuch Nira, 14300 ft, vii 1926, Rock 14371 (K); Alpine Region between Radja and Jupar Range, Wotila, 14500 ft, vi 1929, Rock 14231 (K); Nang La, 20 vi 1936, Ludlow & Sheriff 1840 (BM); N of Lhasa, 15000 ft, 24 vi 1942, Ludlow & Sheriff 8737 (BM); NW of Lhasa, Nyenchentang La, 10 vi 1943, Ludlow & Sheriff 9604 (BM). 13. Anemone fuscopurpurea H.Hara, J. Jap. Bot. 48: 353 (1973). – Type: E Nepal, Banduke pokhari Saju pokhari, 4200 m, 15 vi 1972, Kanai et al. 721252 (holo TI!; iso E!). Leaves 5–7; petioles 5–10 cm long, sparsely puberulent; blades 3-sect, rounded-ovate, 2–3 6 2–4 cm, glabrous, ciliate; segments subsessile; central segments 3-lobed, oblong-rhombic; bases cuneate; margins incised-dentate; lateral segments similar to central ones, but smaller. Scapes 2–4, mainly ascending, 5–15 cm long, basally villous; cymes 2–5-flowered. Involucral bracts 3-lobed, bases cuneate, margins dentate, 2–3 cm long, sparsely puberulent. Pedicels 4–5 cm long, villous. Petaloids 4–5, purple, oblong, basally narrowed, apically rounded, 6–10 6 2–5 mm, puberulent; basal veins 3–5, vein anastomoses 1–3. Stamens 2–2.5 mm long; filaments slightly dilated; anthers oblong. Carpels ovate, glabrous, styles curved, c.1 mm long. Data on achenes lacking. Distribution and habitat. East Nepal (Pokhari, Topke Gola); 3600–4400 m. Taxonomic remarks. Anemone fuscopurpurea was described (Hara, 1973) from the flora of the Himalaya (East Nepal) on the basis of several herbarium specimens without achenes. Hara regarded this taxon as close to both Anemone obtusiloba (sect. Himalayicae) and A. demissa (sect. Omalocarpus). Tarasevich & Chaudhary (1987) examined its pollen grains and placed Anemone fuscopurpurea in a monotypic section Fuscopurpurea. Based on a morphological study of the herbarium material available in E we suggest its affinities are to section Omalocarpus. Until more SEM data become available on the microsculpture of pollen grains from other members of subgenus Omalocarpus and until the fruits of Anemone fuscopurpurea can be studied, 92 S. N. ZIMAN ET AL. we prefer to exclude this species from section Himalayicae and maintain it in section Omalocarpus rather than describe a separate section. Specimens examined. NEPAL. Arun-Tamur, Topke Gola, 13500 ft, 11 v 1956, Stainton 253 (E); Topke Gola, 14000 ft, 4 vii 1971, Beer 8280 (E!). D ISCUSSION: O RIGIN AND E CO-GEOGRAPHICAL R ADIATION A NEMONE SECT. H IMALAYICAE OF The close affinities between Anemone sect. Himalayicae and sect. Omalocarpus, both accommodated within Anemone subgen. Omalocarpus, are supported not only by the numerous apomorphic morphological characters they share (see earlier), but also by their karyotype and chromosome base number x 5 7. This number can be derived by several structural changes from the basic karyotype of x 5 8, plesiomorphic for Anemoninae (Baumberger, 1970). That members of Anemone sect. Himalayicae are characterized by this apomorphic x 5 7 has been obscured by earlier, clearly erroneous counts of 2n 5 16 (e.g. Sobti & Singh, 1961). These have now been superseded by several more careful recent studies which have clearly shown n 5 7 or 2n 5 14 for Anemone obtusiloba (Bhattarai, 1989; further references in Starodubtsev, 1989), A. geum (5 A. obtusiloba subsp. ovalifolia; Yang & Wu, 1993; Huang et al., 1996) and A. trullifolia (Sarkar et al., 1978). Members of Anemone sect. Omalocarpus and sect. Himalayicae are apparently diploid throughout and no polyploidy has yet been recorded for this clade. Furthermore, Hoot et al. (1994) have presented very well-supported evidence from chloroplast DNA restriction analyses for the close relationships of Anemone narcissiflora and A. demissa (Anemone sect. Omalocarpus), and of A. obtusiloba and A. trullifolia (Anemone sect. Himalayicae). Available cladograms suggest a narrow sister relationship of the two species groups within a clearly separated major clade corresponding to the present Anemone subgen. Omalocarpus. This, in turn, is affiliated to a major clade of the genus Anemone s.l., characterized by the chromosome base number x 5 7. Thus, the former placement of Anemone sect. Himalayicae under the genus or section Pulsatilloides (with x 5 8; far from Omalocarpus), maintained until quite recently (e.g. Starodubtsev, 1991), is now definitively refuted. The affinities between members of Anemone sect. Omalocarpus and sect. Himalayicae are so close that their taxonomic separation still suffers from uncertainties. Fruit characters of Anemone rupestris (sole species of Anemone sect. Himalayicae ser. Rupestres) tend towards Anemone sect. Omalocarpus. Furthermore, the placement of two critical and also rather isolated species, Anemone fuscopurpurea H.Hara and A. imbricata Maxim., is still under dispute. They are put either into Anemone sect. Omalocarpus or into monotypic sections of their own (see earlier). This makes further studies on the alliance of Anemone subgen. Omalocarpus necessary. REVISION OF ANEMONE SECT. HIMALAYICAE 93 Members of Anemone sect. Himalayicae differ from those of Anemone sect. Omalocarpus predominantly with respect to derived, apomorphic characters which often can be interpreted as adaptations to high-montane or alpine habitats. This supports the hypothesis that Anemone sect. Himalayicae originated, possibly during the late Tertiary, Pleistocene and Postglacial eras, as a response to the lifting of the Himalayan, Tibetan and SW Chinese mountain systems (see Rowley et al., 2001, etc.), from Anemone sect. Omalocarpus-like ancestors growing at lower elevations. What can be said about the further morphological and eco-geographical differentiation within Anemone sect. Himalayicae? On comparison of taxa from Anemone ser. Rupestres with ser. Obtusilobae and finally with ser. Trullifoliae with respect to vegetative features, we can demonstrate a decrease in leaf size, a reduction in leaf division, an increasing fusion of leaflets combined with loss of petiolules, and an increase in the density of the indumentum. Noteworthy among reproductive features are reductions from several-flowered cymes to solitary flowers, a decrease in anastomosing veins in petaloids, development of staminodes, dilatation of initially linear filaments, and changes in the hairiness of petaloids, pistils and fruits. Trends F I G . 18. Distribution of Anemone obtusiloba. 94 S. N. ZIMAN ET AL. relevant for the achenes concern changes from ovoid to compressed, development of lateral ribs, and transformations of styles from straight to curved, bent or hooked. Within Anemone sect. Himalayicae, Anemone obtusiloba and A. polycarpa appear to have numerous plesiomorphic characters. Most other taxa, however, belong to an intermediate level, whereas Anemone coelestina and A. subindivisa apparently represent evolutionarily more advanced species. The taxa of Anemone sect. Himalayicae are concentrated in the mountains of SW China and the Himalayan ranges of Burma, Bhutan, Nepal and North India. All of them are high-mountain plants occurring in the cool temperate montane to the subalpine and alpine belts at 1850–4800 m elevation (Figs 18–21). From Anemone ser. Obtusilobae the polymorphic A. obtusiloba has reached the widest distribution area within the section and now extends from SW China (with two endemic infraspecific taxa) to the Himalaya (with subsp. nepalensis in the central and var. potentilloides in the western part), Burma, Pakistan, Afghanistan, the mountains of North China, Kirgizstan and Mongolia (var. obtusiloba which also overlaps with other infraspecific taxa). It occurs from open forests at lower elevations to scrub at F I G . 19. Distribution of Anemone patula, A. polycarpa, A. geum and A. subpinnata. REVISION OF ANEMONE SECT. HIMALAYICAE 95 F I G . 20. Distribution of Anemone rockii, A. trullifolia and A. coelestina. the timberline and to grassland, rocks and talus in alpine regions. Among the other members of Anemone ser. Obtusilobae, the species A. geum and A. rockii have also attained large areas of distribution and wide altitudinal ranges. The former reaches to the western Himalaya and from the mountains of SW China to the eastern Tienshan in Xinjang. Anemone rockii extends from Sichuan to the central Himalaya. In contrast, Anemone polycarpa is limited as an ultraoreophyte to altitudes above 3500 m and ranges through the Himalaya to SW China, whereas its close relatives A. patula and A. subpinnata occur as local endemics in the mountains of Sichuan. The taxa of Anemone sect. Himalayicae ser. Trullifoliae and ser. Rupestres exhibit a similar eco-geographical pattern but are more restricted and disjunct in the central Himalaya and the mountains of SW China. Populations in both areas and with a wide altitudinal range characterize the Anemone ser. Trullifoliae taxa A. trullifolia 96 S. N. ZIMAN ET AL. F I G . 21. Distribution of Anemone yulongshanica, A. subindivisa and A. rupestris. var. trullifolia and A. coelestina var. holophylla, whereas A. yulongshanica and A. subindivisa are endemics in SW China. The other taxa of Anemone ser. Trullifoliae are restricted to high altitudes: A. coelestina var. coelestina and var. linearis occur in both areas, whereas A. trullifolia var. liangshanica and var. lutienensis have been found locally in SW China only. The infraspecific differentiation of Anemone rupestris in the monotypic series Rupestres is comparable: subsp. rupestris occurs at a relatively wide range of altitudes in the central and eastern Himalaya and adjacent Tibet, whereas subsp. gelida is an ultraoreophyte with populations in the Himalaya and the mountains of SW China. In retrospect, one can conclude that the present eco-geographical distribution of taxa within Anemone sect. Himalayicae is due to an allopatric pattern of differentiation. This has apparently followed two parallel trends of expansion: (i) from the SW Chinese mountains and the eastern and central Himalaya to the western Himalaya, Afghanistan, the Tienshan and to mountain ranges in NW China and adjacent central Asia; (ii) from lower to higher and highest mountain regions. Comparable examples for an allopatric east/west differentiation along the Himalaya are afforded by the Galium serpylloides group (Rubiaceae–Rubieae: SchönbeckTemesy & Ehrendorfer, 1987) and many other taxa. Within Anemone sect. Himalayicae the highest concentration of local endemic taxa (seven) is found in SW China, whereas only one other endemic is documented for the central Himalaya. REVISION OF ANEMONE SECT. HIMALAYICAE 97 In most regions of the distribution area of the section more than one taxon has been found. Future field work should clarify if there is isolation between these taxa (due either to their occurrence in different habitats or to other mechanisms). 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Received 11 April 2003; accepted for publication 5 December 2006

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