Ranunculus L. is a herbaceous genus of the Ranunculaceae, and one of the genera which has become diversified in the alpine zone of the Himalaya (Ohba, 1988). Flowers of Ranunculus are bowl-shaped with petals having glossy proximal and mat distal portions. The bowl-shaped flower serves as a reflector which focuses solar beams and heat on both the gynoecium and androecium. The center of the flower is attractive to insects because of the partial difference in the texture of the petals. This morphology is consequently regarded as an adaptation to entomophily. It is an interesting subject how such an entomophilous plant group has differentiated in the alpine zone of an area where insects are less abundant. Although Ranunculus of the Himalaya has been extensively studied by several students since Hooker's 'Flora of British India', Vol. 1 (1872), the systematics of the genus are still unsatisfactory. I have examined many herbarium materials deposited in the Herbarium, the Uni versity of Tokyo (TI), from botanical explorations to the Himalaya and specimens received on exchange from other herbaria. Fortunately, in 1988, I also had an opportunity to make field observation of natural populations in the Makalu region of E Nepal. This paper, based on the results of those observations, discusses the morphological variation in some alpine species of the Himalaya and describes two new species and a new variety.

It has been known since the time of Linnaeus that the petal blades of Ranunculus have nectaries at the bases. The nectaries are naked, covered by scaly appendages or surrounded by cylindrical borders. Benson (1940, 1941-1942, 1948, 1954, 1955) was the first to thoroughly investigate the shape of the nectary in various taxa and evaluated its taxonomic significance in the genus. Benson's contribution is now widely accepted and nectary shape is often described in floristic manuals and revisions (e.g., Davis, 1960; Tutin, 1964; L. Liou [or L. Liu], 1980,1985). Kosuge and Tamura (1989) and Kosuge (pers. com.) reported that R. sceleratus L. from Japan shows a considerably wide range of variation in nectary shape. Based on my own observa tions, nectary shape varies only slightly within a species in the Himalaya as well as in Japan (Kadota, 1990), and hence the wide range of nectary shape variation in R. sceleratus of Japan is rather exceptional. It is therefore reasonable to use nectary shape as a key character. The morphology of petals, however, has been scarcely studied with regard to the Himalayan species. Consequently, in this paper, I attempt to describe the petals in detail paying particular attention to nectary shape. Two types of nectary are present in the alpine species of the region and are discussed here.

Nectaries of the first type are shallow pockets with scaly appendages, which are laterally free from the petal blades. According to Benson (1940), this type of nectary is considered to be more primitive. However, Tamura (1965) stated that nectaries of this type are found among more advanced groups as well as in primitive groups. The second type of nectary is characterized by cup-shaped nectary pits without scaly appendages. Nectaries are very variable in size and shape. They may be spot-like, pocket-like, pocket-like to forked, circular to forked, lunate or shallow or form a large pocket. Benson (1940) stated that cup-like structures were derived from the lateral margins of the scaly appendage becomming connate to the petal blade. The laterally free scales are devoid of veins. If cup-shaped structures are the result of connation, as suspected by Benson (1940), the "cup" ought to lack veins. Among the Himalayan species (Figs. 1-11), however, most of the "cups" are actually provided with veins among at least the Himalayan species. Kosuge and Tamura (1989) additionally showed that scales and nectary pits develop independently. Plants belonging to Trollius and Coptis, which are generally believed to be primitive in the Ranunculaceae, always have cup-shaped nectary pits. The evolutionary trend in petals proposed by Benson (1940) is thus problematic.

This paper discusses three types of attachment and dehiscence of the anthers: type I, basifixed and latrorse; type II, dorsifixed and latrorse; type III, dorsifixed and extrorse. In species of the woody Ranales, which are believed to be the most primitive among the insect-pollinated species, the anthers are dorsifixed and introrse, latrorse or extrorse. Type I anthers consequently seem to represent a more advanced condition, although little attention has been paid to the three types of stamens within Ranunculus. Hence the stamens of species from other regions need further study in order to understand the evolutionary process in the genus Ranun culus.

In the alpine Himalaya and the arctic circumpolar region, species with similar growth forms occur. For example, R, pegaeus Hand.-Mazz, (dwarf perennial with creeping stem in wet habitats) and R. pseudopygmaeus Hand.-Mazz. (dwarf peren nial with abbreviated scape in rather mesic habitats) correspond to R. hyperboreus Rottb. and R. pygmaeus Wahlb., respectively. Although R. pegaeus is superficially similar to R. hyperboreus, the former is significantly different from the latter in nectary shape. Ranunculus pseudopygmaeus is discriminated from R. pygmaeus by the difference in anther attachment. The superficial resemblance in gross morphology in these species may be the result of adaptation to similarly severe environmental conditions in both regions (i.e., an example of parallelism based on convergent evolution).

During field survey in the Eastem Himalaya, I observed several individuals putatively considered to be hybrids between R. pulchellus C.A. Mey. and R. pseudopygmaeus. The hybridity was assessed using percent pollen stainability (Kadota, 1978) and mor phological attributes. I will preliminarily report the expression of characters of the putative parental species in the hybrids.

Systematic Treatment

1) Ranunculus albertii Regel et Schmalh. in Acta Hort. Petrop. 5: 223 (1877) - Ovcz. in Fl. URSS 7: 402 (1937)-Icon. Conn. Sin. 1: 713, fig. 1425 (1972)-L. Liou in Fl. Reip. Pop. Sin. 28: 277, fig. 37 (1980)-Ranunculus sulphureus C. J. Phipps var. albertii (Regel et Schmalh.) Maxim, in Enum. Pl. Mongol. 19 (1889). [Fig. 1]

Perennial herb. Roots cord-like, not tuberous. Stem ascending, solitary, simple or branched in the upper part, 5-8 cm high, densely pilose with yellowish, smooth surfaced, subpatent and curly hairs; collar sparsely clothed with fibrous remains of petioles. Basal leaves obovate-cuneate, 10-22 mm long, 5-13 mm broad, simple, crenate, cuneate at the bases, sparsely pilose with whitish appressed hairs on the adaxial side, glabrous on the abaxial side, long petiolate; petioles 2.5-6 cm long, clearly longer than the blades, sparsely pilose or glabrous, slightly vaginate at the basal parts. Cauline leaves broadly obovate-obovate, dentate to medially 3-lobed, ca. 15 mm long, 8-12 mm broad, broadly cuneate, shortly petioled. Flowers yellow, ca. 15 mm in diameter, solitary or 2-5 in a loose raceme; pedicels sulcate, 1-2 cm long, densely pilose with yellowish, subpatent and curly hairs; bracts broadly ovate, 7-17 mm long, 5-21 mm broad, deeply 3-lobed, vaginate at the basal parts. Sepals 5, elliptic, ca. 5 mm long, ca. 3 mm broad, purplish, convex, membranous along the margin, patent, pilose with whitish appressed hairs on the abaxial side. Petals 5, broadly obovate, ca. 9 mm long and broad, overlapping each other; claws ca. 0.5 mm long; nectaries cup-shaped with thickened borders. Stamens ca, 20; anthers 1 mm long, basifixed and latrorse; filaments 1.5 mm long, flattened. Carpels glabrous. Receptacles sparsely pilose. Fruiting heads and achenes not seen.

Specimen examined. C NEPAL. Gandaki Zone. Managhot Dist., Phedi-Thanti, alt. 4400-4080 m (H. Ohba et al. 8330877, July 26, 1983, TI).

This species, new to the Flora of Nepal, was hitherto reported from Central Asia (Tien Shan, Altai and Pamir) and China (Xinjiang). Ovczinnikov (1937) and Liou (1980) described the basal leaf blades of R. albertii as orbicular-reniform or fan shaped. Our plants, however, have obovate or cuneate basal leaf blades. Since the locality in Nepal is far segregated from those in Central Asia and China, the Nepalese plants are probably infraspecifically distinct.

Ranunculus albertii is similar to R. altaicus Laxm. [= R. sulphureus Soland. var. altaicus (Laxm.) Trautv.; R. altaicus Laxm. subsp. altaicus (Kadota, 1990)] in basal leaf shape and scapose habit, but the former is distinguished from the latter by flower size (1.5 cm vs. 2.5-3.5 cm in diameter) and hair color of sepals and receptacles (whitish vs. brownish).

2) Ranunculus brotherusii Freyn in Bull. Herb. Boiss. 6: 885 (1898)-Ovcz. in Fl. URSS 7: 392, t. 24, f. 4 (1937)-Hand.-Mazz. in Acta Hort. Gothob. 13: 148 (1939) -Mukerjee in Bull. Bot. Surv. Ind. 2: 102 (1960)-Hara in Fl. E Himal. 90 (1966); in Fl. E Himal. 2: 31 (1971); in Hara & Williams, Enum. Fl. Pl. Nepal 2: 19 (1979)-Tamura in Acta Phytotax. Geobot. 23: 31, 105 (1968)-Inst. Bot. Bor.-Occ. Acad. Sin., Fl. Tsinling. 1(2): 271, fig. 232 (1974)-L. Liou in Fl. Reip. Pop. Sin. 28: 295 (1980)-Polunin & Stainton, Fl. Himal. 10, pl. 5, 38 (1984). [Fig. 2]

Ranunculus affinis R. Br. β. tanguticus Maxim., Fl. Tangut. 14 (1889)-Ranunculus tanguticus (Maxim.) Ovcz. in Fl. URSS 7: 392 (1937)-L. Liou in Fl. Reip. Pop. Sin. 28: 295, pl. 92, figs. 1-2 (1980); in Fl. Xizang. 2: 105, fig. 31, 1-2 (1985)-Ranunculus brotherusii Freyn var. tanguticus (Maxim.) Tamura in Acta Phytotax. Geobot 23: 31, 105 (1968).

Ranunculus affinis R. Br. β. tanguticus Maxim. lusus dasy carpus Maxim., Fl. Tangut. 14 (1889)-Ranunculus brotherusii Freyn var. dasycarpus (Maxim.) Hand.- Mazz. in Acta Hort. Gothob. 13: 149 (1939)-Mukerjee in Bull. Bot. Surv. Ind. 2:102 (1960)-Ranunculus tanguticus (Maxim.) Ovcz. var. dasycarpus (Maxim.) L. Liou in Fl. Reip. Pop. Sin. 28: 297 (1980)-Ranunculus brotherusii Freyn var. tanguticus (Maxim.) Ovcz. f. dasycarpus (Maxim.) L. Liou in Fl. Xizang. 2: 105 (1985).

Ranunculus affinis R. Br. var. capillaceus Franch. in Pl. Delav. 19 (1889)-Hand.- Mazz., Symb. Sin. 7: 306 (1931)-Ranunculus tanguticus (Maxim.) Ovcz. var. capillaceus (Franch.) L. Liou in Fl. Reip. Pop. Sin. 28: 297 (1980).

Perennial herb. Roots fibrous but somewhat thickened and fusiform. Stem solitary or caespitose, erect, ascending or sometimes decumbent, 1.5-20(-40) cm long, simple or branched, sparsely pilose with long, curly and whitish hairs or glabrous; collars clothed with fibrous remains of leaf bases. Basal leaves several, sparsely pilose with whitish and appressed hairs on both sides; blades reniform or roundish, 6-20(-50) mm long, 6-25(-55) mm broad, deeply 3-lobed or ternate; middle lobes ovate, medially to deeply 3-lobed; lateral lobes medially to deeply 2-lobed, cordate; petioles 1-8 cm long, vaginate, pilose with whitish curly hairs. Lower cauline leaves similar to the basal ones in shape, subsessile; petioles vaginate. Upper cauline leaves deeply 3-5-lobed; laciniae linear. Flowers yellow, solitary, 5-15 mm in diameter; pedicels sulcate, 1-6 cm long, elongated up to 9 cm long at fructescence, pilose with whitish curly hairs. Sepals ovate to broadly oblong, membranaceous, patent, shorter than the petals, pilose with appressed hairs. Petals 5, narrowly or broadly obovate, including claws 5-6 mm long; nectaries pocket-like or forked. Anthers ca. 1 mm long, dorsifixed and latrorse; filaments dilated. Fruiting heads ovoid to cylindrical, 4-6 mm long; receptacles glabrous. Achenes obovoid, ca. 1.5 mm long, glabrous or hirsute with short and spreading hairs; beaks incurved or sometimes straight, ca. 0.5 mm long.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Saba Distr., Thudam-Lamni Nama, alt. 3400-4200 m (H. Ohashi et al. 770706,773717, 773762, Aug. 14, 1977, TI); Lamni Nama, alt. 4200-4900 m (H. Ohashi et al. s.n., Aug. 15, 1977, TI); Lamni Nam-Tasagon, alt. 4200-4350 m (H. Ohashi et al. 770828, 773818, Aug. 16, 1977, TI); Tasagon-Topke Gola (Mechi Zone. Taplejung Distr.), alt. 4350-3600 m (H. Ohashi et al. 770911, 772514, Aug. 17, 1977, TI). Cha Ding Kharka-Numbuk, alt. 3650 m (M. Suzuki et al. 8850435, July 17, 1988,TI); Phematang (Tematang) Kharka-Yangri Kharka, alt. 3400 m (M. Suzuki et al. 8850486, July 18, 1988, TI); Yangri Kharka, alt. 3540 m (M. Suzuki et al. 8850533, 8850870-8850879, July 19, 1988, TI); Refuk Kharka-Ta Dasa, alt. 3710 m (M. Suzuki et al. 8850552, 8850889-8850892, July 20, 1988, TI); Lyang Mo Le Kharka-Merek, alt. 4300 m (M. Suzuki et al. 8850596. 8850912-8850914, July 21, 1988, TI); Merek (M. Suzuki et al. 8820801, July 22, 1988, TI); Makalu Base Camp, alt. 4600 m (M. Suzuki et al. 8850616, 8850924, July 23, 1988, TI). Sagarmatha Zone. Solukhumbu Distr., Thosa Kharka-Beni (Dambuk) Kharka, alt. 3670-3970 m (H. Ohba et al. 8571911-8571912, Aug. 22, 1985, TI); Beni Kharka -Yurigolcha, alt. 3970-4500 m (S. K. Wu 8581132, Aug. 23,1985, TI). Janakpur Zone. Ramechhap Distr., Baula Pokhari, alt. 3960 m (H. Ohba et al, 8570362, July 10, 1985, TI); Jama Pokhari, alt. 4220-4400 m (N. Kurosaki 8570572, July 19, 1985, TI); Botase Kharka-Koshing Kharka, alt. 4500-4000 m (H. Ohba et al. 8570590, 8570617, July 21, 1985, TI); Thare Og, alt. 4150 m (H. Ohba et al. 8570744, 8580434, July 23, 1985, TI); Thare Og - Thare Teng, alt. 4150-4260 m (H. Ohba et al. 8570774, July 24, 1985, TI). Dolakha Distr., Rolwaling Khola, Beding - Na, alt. 3600-4050 m (H. Ohba et al. 8331785, 8351288, Sept. 4, 1983, TI). C Nepal. Dhaulagiri Zone. Mustang Distr., Kagbeni - Muktinath, alt. 2800-3550 m (H. Ohba et al. 8330723, July 22, 1983, TI).

This species is very common in the alpine zone and is extraordinarily variable in gross morphology. A large population is located in an Abies-Rhododendron forest near Refuk Kharka (3710 m, M. Suzuki et al. 8850552), E Nepal. Plants from this population are characterized by their ascending to decumbent stems and membranous leaves. The stem posture and the leaf thickness are regarded as adaptations to the closed habitat. The Refuk Kharka population is also characterized by having abnormal petals. The petals are irregular in shape and size and some of their nectary pits are very small and vestigial or completely absent. The cause of this abnormality needs further study.

Liou (1980) stated that R. brotherusii is discriminated from R. tanguticus by its dwarf size and the deeply 3-lobed and not temate blades of the basal leaves. However, plant size and degree of lobe incision in the basal leaves also shows a considerable range of variation. Hence R. tanguticus is considered to be synonymous with R. brotherusii.

M. Suzuki et al. 8850433, 8850486, 8850533, 8850871, and 8850878 may be identified as var. dasycarpus (Maxim.) Hand.-Mazz., which is characterized by pubescent achenes. Based on the results of observation made in the Makalu region (1988), however, colonies of plants with hirsute achenes are sporadically found among the individuals with glabrous achenes and plants with this feature are therefore not given taxonomic recognition.

Although anthers of this species are fundamentally basifixed and latrorse, they show dorsi-ventrality to some degree; the filaments are not fully differentiated from the connectives (Fig. 2). This type of anther attachment may indicate an intermediate situation from 'dorsifixed and extrorse' to 'basifixed and latrorse'.

Tamura (1968) commented on the relationship between this species and R. yatsugatakensis Honda et Kumazawa and R. kitadakeanus Ohwi, both endemic to central Japan. The two Japanese species undoubtedly resemble R. brotherusii in gross morphology. Ranunculus yatsugatakensis occurs on wet rocks in the subalpine and alpine zones of Mts. Yatsugatake, and has round petals with lunate or somewhat forked nectaries and basifixed and latrorse anthers. Ranunculus kitadakeanus grows on or among rocks in the alpine zone of Mt. Kitadake adjacent areas and is characterized by obovate petals with small pocket-like nectaries and basifixed and latrorse anthers (Fig. 3). Ranunculus yatsugatakensis is accordingly more similar to this species than to R. kitadakeanus in respect to nectary shape. However, the Japanese species show a more advanced condition so far as the anther attachment is concerned. Both Japanese species are restricted to the narrow area and are very rarely encountered. The two species are thus considered to be relictual. Ranunculus brotherusii, however, is aggressive and very common, at least in the Eastern Himalaya. The relationship between R. brotherusii and the two Japanese species needs further study, including a comparison of chromosome numbers.

3) Ranunculus glabricaulis (Hand.-Mazz.) L. Liou in Fl. Reip. Pop. Sin. 28: 298, pl. 90, 6 (1980); in Fl. Xizang. 2: 105, fig, 28, 3 (1985)-Ranunculus hirtellus Royle ex D. Don var. glabricaulis Hand.-Mazz. in Acta Hort. Gothob. 13: 151 (1939). [Fig. 4]

Dwarf perennial herb. Roots proximally, slightly fusiform. Stem erect to ascending or decumbent, simple or branched, 1-5 cm long, glabrous; collar sparsely clothed with fibrous remains of leaf bases. Basal leaves 2-3; blades reniform to pentagonal in outline, 2-8 mm long, 3-12 mm broad, deeply 3-lobed or temate, glabrous on both sides; middle lobes oblong to obovate, tricuspidate to entire; lateral lobes obliquely obovate, entire or toothed; leaf bases shallowly cordate to truncate; petioles 0.5-3 cm long, glabrous, vaginate at the basal parts. Cauline leaves 1-3 or absent; blades deeply 3-lobed with entire lobes or lanceolate and entire, glabrous. Flowers solitary to several, 5-7 mm in diameter; pedicels sulcate, 0.5-1.5 cm long, sparsely hirsute with subpatent hairs or glabrous. Sepals oblong to broadly obovate, 3-4 mm long, navicular, patent, glabrous, acute or obtuse at the apices, purplish. Petals 5, obovate or sometimes oblong, including claws 2..5-4.5 mm long, nearly equal to the sepals in length, 1-2 mm broad, truncate or obtuse; claws ca. 0.5 mm long; nectaries cup-shaped. Anthers 0.7-1.0 mm long, basifixed and latrorse; filaments slightly flattened. Receptacles and carpels glabrous. Mature fruiting heads and achenes not observed.

Specimens examined. E NEPAL. Janakpur Zone. Ramechhap Distr., Dubi Kharka-Baula Pokhari, alt. 3700-3960 m (H. Ohba et al. 8580236, July 9, 1985, -n); Baula Pokhari-Chhu Ningma, alt. 3960-4040 m (H. Ohba et al. 8530228, July 11, 1985, TI); Jata Pokhari, alt. 4220 m (H. Ohba et al. 8570443, July 13, 1985, TI).

Ranunculus glabricaulis grows on open, grassy, rocky ground in the alpine zone. This species was hitherto known only from W Gansu, China. Hence this is the first report outside China.

Dwarf forms of R. hirtellus are similar to this species, but the former is discriminated from the latter by: 1) the glabrous stems, pedicels, leaves and sepals (sericeous vs. glabrous); 2) sepal color (green vs. purplish); and 3) the shape of the nectary pits (deeply cup-shaped or forked vs. shallowly cup-shaped).

Liou (1980, 1985) described fruits of this species as "fruiting heads ca. 5 mm in diameter; achenes many, ovato-orbicular, glabrous" (the original text is written in Chinese). In his description, he also stated "flowers %hellip;, 1-1.5 cm in diameter; sepals …, 5-7 mm long; petals 6-9 mm long." Our Nepalese plants, however, have smaller flowers, sepals and petals. The taxonomic status of the Nepalese plants of this species needs further study.

4) Ranunculus himalaicus Kadota, sp. nov. [Fig. 5]

Haec species Ranunculo glabricauli affinis, sed a R. glabricaule nectariis squa-matis, receptaculis hirsutis et caule sericeo facile distinguitur.

Herba nana perennis, rhizomate erecto 0.5-1 mm longo, radicibus cylindricis ca. 1 mm diametro. Caulis erectus, solitarius, simplex, 0.5-5 cm altus, sericeus, pilis albis cripsis, collo vestito. Folia basalia prelumque 3-5, ambitu late ovata, 1-5 mm longa, 1-4 mm lata, ternata vel profunde trilobata, glabrata vel sparse sericea, lobis mediis obovatis integris ad cuspidatis vel profunde trifidis obtusis, lobis lateralibus integris ad dentatis vel profunde divisis, laciniis anguste lanceolatis minus quam 0.5 mm latis, basi late cuneatis, petiolo 5-25 mm longo glabro basi vaginato. Folia caulina 1 vel absentia, ovata, profunde trilobata, basi breviter vaginata. Flores lutei, 5-7 mm diametro, solitari, pedicellis sucatis, 5-25 mm longis, sericeis, pilis albis cripsis, absque bracteolis. Sepala 5, obovato-elliptica, 3-3.5 mm longa, purpurea, patentia, glabra vel sparse villosa, pilis albis subpatentibus. Petala 5, obovata ad late elliptica vel suborbiculata, 2-4.5 mm longa, 1.6-2.6 mm lata, unguibus 0.1- 0.6 mm longis non incrassatis, nectariis vix depressis squamatis. Stamina ca, 20, antheris 0.6-0.8 mm longis dorsifixis et extrosis dehiscentibus vel basifixis et latrorsis, filamentis dilatatis. Carpella glabra et laevigata. Receptacula hirsuta, pilis albis rectis vel adscendentibus. Achenia matura mihi incognita.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Sabha Distr., above Makalu Base Camp, 85° 05'E, 27° 50'N, alt. 4900 m (M. Suzuki et al. 8850636, July 24, 1988, Holotype, TI); Merek, 85° 05'E, 27° 45'N, alt. 4620 m (M. Suzuki et al. 8850613, July 22, 1988, TI). Janakpur Zone, Ramechhap Distr., Jata Pokhari-NE of Panchi Pokhari, 86° 25'E, 27° 43'N, alt. 4600 m (M. Suzuki et al. 8570497, July 16, 1985, TI).

Ranunculus himalaicus grows in tundra of the subnival zone, flowering close to the ground surface. In the Makalu region, E Nepal, the species occurs sympatricly with R. brotherusii and R. pseudopygmaeus. The three species are in full bloom in late July. Putative hybrids among the three taxa, however, have not been found.

Ranunculus himalaicus is characterized by: 1) abbreviated scapes with sericeous whitish hairs; 2) ternate or deeply 3-lobed basal leaves; 3) small flowers 5-7 mm in diameter; 4) small, shallow nectary pits covered with diminutive scales; and 5) receptacles hirsute with whitish hairs. The new species belongs to Sect. Ranunculus on the basis of the nectary shape of the petals and the cord-like roots. This species bears a superficial resemblance to R. glabricaulis in its occurrence in the subnival zone and in its erect and very short scape and small flowers. Ranunculus himalaicus, however, is clearly distinguished from R. glabricaulis by having diminutive but definite nectary scales, hirsute receptacles and sericeous stems.

Most plants of this species have basifixed and latrorse anthers, but in some plants from even a single population may have dorsifixed and extrorse anthers (Fig. 5 c). There is no significant variation in attachment and dehiscence of anther sacs within an individual. The significance and cause of the intraspecific variation of attachment and dehiscence of the anthers remains unknown.

5) Ranunculus hirtellus Royle ex D. Don in Royle, Ill. Bot. Himal. 53 (1834)-Hook. f. & Thorns., Fl. Ind. 1: 34 (1855); Fl. Brit. Ind. 1: 18 (1872)-Pampan., Fl. Caracorum 113 (1930)-Hand.-Mazz. in Acta Hort. Gothob. 13: 151, fig. 4, nos. 16-17 (1939)-Baehni et al. in Candollea 13: 226 (1951)-Tamura & Kitamura, F. & Fl. Nep. Himal. 130 (1955) -Mukerjee in Bull. Bot. Surv. Ind. 2: 103 (1960)-Tamura in Kitamura, Fl. Pl. W Pakist. 61 (1964)-Banerji in Rec. Bot. Surv. Ind. 19: 19 (1966)-L. Liou in Fl. Reip. Pop. Sin. 28: 292, fig. 38 (1980); in Fl. Xizang. 2: 104 (1985). [Fig. 6]

Ranunculus affinis R. Br. a. typicus Fin. & Gagnep. in Bull. Soc, Bot. Fr. 51: 314 (1904)-Hand.-Mazz., Symb. Sin. 3: 306 (1931).

Ranunculus affinis R. Br. var. ternatus Diels in Not. Bot. Gard. Edinb. 7: 231 (1912)-Hand.-Mazz., Symb. Sin. 3: 306 (1931).

Ranunculus attenuatus Royle ex D. Don in Royle, Ill. Bot. Himal. 53 (1834).

Ranunculus choorensis Royle ex D. Don in Royle, Ill. Bot. Himal. 53 (1834).

Ranunculus glabellus Royle ex D. Don in Royle, Ill. Bot. Himal. 53 (1834).

Ranunculus longipetalus Hand.-Mazz. in Acta Hort. Gothob. 13: 160, tab. 1, figs. 1-7 (1939)-L. Liou in Fl. Reip. Pop. Sin. 28: 280, pl. 89, 1-2 (1980).

Ranunculus nervosus Royle ex D. Don in Royle, Ill. Bot. Himal. 53 (1834).

Ranunculus micronivalis Hand.-Mazz. in Sitzgsanz. Ak. Wiss. Wien. Math.-Nat. Kl. 57: 48 (1920); Symb. Sin. 7: 307, tab. 6, fig. 14 (1929); in Acta Hort. Gothob. 13: 160 (1939).

Ranunculus nivalis auct. non L.: Hook. f. & Thoms., Fl. Ind. 1: 35 (1835): Fl. Brit. Ind. 1: 19 (1872).

Ranunculus pedicellatus Hand.-Mazz. in Acta Hort. Gothob. 13: 161, tab. 1. figs. 10-11 (1939).

Ranunculus stenorhynchus Franch, in Nouv. Arch. Mus. Paris, Ser. 2: 189 (1885); Bull. Soc. Bot. Fr. 33: 373 (1886); Pl. David. 2: 7 (1888)-Fin. & Gagnep. in Bull. Soc. Bot. Fr. 51: 319 (1904).

Perennial herb. Roots proximally, slightly fusiform. Stem ascending, spreading to decumbent or erect, 5-16 cm long, branched or simple, pilose with ascending and/or patent whitish hairs; collar sparsely clothed with fibrous remains of leaf bases. Basal leaves usually 3 to several; blades reniform to orbicular or broadly ovate, 5-18 mm long, 6-22 mm broad, medially to deeply 3-lobed or sometimes ternate, densely sericeous with appressed hairs on the adaxial sides and along the margins, sparsely sericeous on the abaxial sides, or almost glabrous on both sides; middle lobes broadly oblong to obovate, tricuspidate or entire, obtuse; lateral lobes obliquely ovate, coarsely toothed or entire; leaf bases cordate to truncate; petioles 1.5-8 cm long, sparsely sericeous or glabrous, vaginate at the basal parts. Cauline leaves deeply 3 lobed or entire, sericeous or glabrous, vaginate and amplexicaul; lobes linearly lanceolate. Flowers solitary to several, 6-15 mm in diameter; pedicels sulcate, 0.5-6 cm long, pilose with appressed hairs and/or curly subpatent hairs. Sepals oblong to orbicular, 3-5.5 mm long, navicular, patent or .slightly recurved, sericeous with ascending hairs, membranous and sometimes purplish along the margins. Petals 5, oblong to broadly ovate, sometimes asymmetrical, obtuse to rounded or slightly emarginate, including claws 5-6 mm long, 1.5-5 mm broad; claws 0.5-1 mm long; nectaries cup-shaped or forked, sometimes borders of the nectary cups slightly projected. Anthers 0.8-1.2 mm long, basifixed and latrorse; filaments more or less flattened. Fruiting heads ovoid, ca. 4 mm long and broad; receptacles glabrous or pilose. Achenes suborbicular, 1.5-1.7 mm long, somewhat inflated, glabrous and smooth; beaks ca. 1 mm long, straight or recurved.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Sabha Distr., Khongma-Numbuk, alt. 3500 m (M. Suzuki et al. 8820544, July 17, 1988, TI); Yangri Kharka, alt. 3540 m (M. Suzuki et al. 8850534, July 19, 1988, TI); Ne Kharka, alt. 3650 m (M. Suzuki et al. 8820738, 8850563, 8850900-8850902, July 20, 1988, TI); Merek, alt. 4380 m (M. Suzuki etal. 8850925, 8860631, July 23, 1988, TI). Sagarmatha Zone. Solukhumbu Distr., Beni Kharka, alt. 3970 m (H. Ohba et al. 8572015, Aug. 25, 1985, TI). Janakpur Zone. Ramechhap Distr., Serdingma-Dubi Kharka, alt. 3400-3720 m (H. Ohba et al. 8570245, July 7, 1985, TI); Dubi Kharka-Baula Pokhari, alt. 3960 m (H. Ohba et al. 8530177, 8580223, July 9, 1985, TI); Baula Pokhari-Chhu Ningma, alt. 3960-4040 m (H. Ohba et al. 8530256, July 11, 1985, TI); Thare Og-Neju, alt, 4150-3651 m (H. Ohba et al. 8570957, July 30, 1985, TI). Rolwaling Khola, Na-Khabun, ait. 4050-4400 m (H. Ohba et al. 8351331, Sept. 6, 1983, TI). C NEPAL. Bagmati Zone. Rasuwa Distr., Shin Gompa-Gosainkund, alt. 3200-4300 m (M. Suzuki & S. Noshiro 8541045, June 1, 1985, TI).

In the Makalu region, this species is rather rare and is restricted to rocky slopes, resulting in a relatively narrow range of variation in scape height, size and shape of leaves and depth of the leaf division.

This species is discriminated from R. pulchellus by: 1) ovoid fruiting heads; 2) usually reniform basal leaves; 3) more slender roots; and 4) well developed nectaries.

6) Ranunculus makaluensis Kadota, sp. nov. [Fig. 7]

Herba perennis nana, radicibus dimorphis, tuberosis et fibrosis. Caulis adscendens, solitarius vel 1-3, simplex vel ramosus, 2-6 cm altus, sericeus, pilis albis adpressis crispis. Folia basalia ambitu renifbrmia vel orbiculata, 3-16 mm longa, 4-16 mm lata, non profunde trilobata, plus minusve sericea in pagina adaxiali vel utrinque glabra, pilis albis adpressis, ciliata, lobis mediis late ovatis integris vel dentatis obtusis, lobis lateralibus non profunde bifidis, basi non profunde vel profunde cordatis, petiolo 1-4 mm longo sparsim sericeo vel glabro basi vaginato. Folia caulina obovata vel suborbiculata grosse dentata, 2.5-5 mm longa et lata, basi late cuneata. Flores lutei, ca. 4 mm diametro, solitari vel 1-3 in laxo racemo, pedicellis sulcatis, (0.3 -) 1-3 cm longis dense sericeis, bracteolis 1-2 simplicibus lanceolatis 2-5 mm longis glabris vel sparsim sericeis. Sepala 5, ovata, ca. 2 mm longa, ca. 1 mm lata, glabra, concava, ad margine membranacea, adscendentia, post anthesin persistentia. Petala 5, elliptica, 2-3.4 mm longa, 1-1.8 mm lata, unguibus ca. 0.5 mm longis non incrassatis, nectariis leviter depressis squamatis. Stamina minus quam 10, antheris 0.4-0.7 mm longis, dorsifixis et extrorsis dehiscentibus, filamentis dilatatis. Capitula nuculae hemispherica vel late obovoidea, ca. 2 mm longa, ca. 3 mm diametro, receptaculiis glabris. Achenia plus minusve 10, suborbiculata, ca. 1 mm longa, glabra, tuberculata, lenticulata, rostris ca. 0.2 mm longis valde recurvatis, stigmatibus lanceolatis.

Specimens examined. E NEPAL. Koshi Zone, Sankhuwa Sabha Distr., Merek, 87°05'E, 27°45'N, alt, 4340 m (M. Suzuki et al. 8850675, July 27, 1988, Holotype, TI, Isotype, TNS); Merek, 87°05'E, 27° 45'N, alt. 4340 m (M. Suzuki et al. 8850677, July 27, 1988, TI); Ta Dasa-Merek, 87°05'E, 27°45'N, alt. 4340 m (M. Suzuki et al. 8820745, July 21, 1988, TI).

Ranunculus makaluensis grows on wet mossy, partially shaded rocks in the alpine and subnival zones in the Makalu region.

This species is characterized by: 1) its dwarf size; 2) dimorphic roots; 3) scaly petals; and 4) tuberculate and flat achenes. Hence this is a constituent species of ser. Xiphocoma (Stev.) L. M. Kemularia-Natadze [Fl. Gruzii 3: 240 (1975)] (= subgenus Ranunculastrum DC. sect. Xiphocoma (Stev.) Ovcz., Fl. URSS 7: 483 (1937), nom. invalid.)

The nectary pits are so small that they take the form of shallow depressions, and the scaly appendages of petals are very small (less than 0.1 mm long) and irregular in shape. Because of this nectaries of this species are suspected of representing a degenerate condition.

7) Ranunculus pegaeus Hand.-Mazz. in Acta Hort. Gothob. 13:141 (1939)-Hara, Fl. E Himal. 3: 37 (1975); in Hara & Williams, Enum. Fl. Pl. Nepal 2: 20 (1979)-L. Liou in Fl. Reip. Pop. Sin. 28: 280, pl. 88, figs. 7-8 (1980); in Fl. Xizang. 2: 101, fig. 28, 6-7 (1985).[Fig. 8, a-c]

Ranunculus hyperboreus auct. non Rottb.: Hook. f. et Thoms., Fl. Br. Ind. 1: 18 (1812), p.p.-Hand.-Mazz., Simb. Sin. 7: 308 (1931), p.p.

Dwarf perennial herb. Roots somewhat thickened proximally, fibrous roots few. Stem green or purplish brown, creeping and stoloniferous, rooting at the nodes, sometimes decumbent and not stoloniferous, glabrous; internodes 1-8 cm long. Blades of the basal leaves pentagonal to reniform, 2-9 mm long, 2-10 mm broad. medially to deeply 3-lobed or temate, glabrous, herbaceous or sometimes subcori-aceous, truncate or shallowly cordate at the bases; middle lobes lanceolate to broadly ovate, simple or rarely toothed, 1-5 mm long, 1-4 mm broad; lateral lobes simple or 2-parted; petioles 2-45 mm long, vaginate at the basal parts. Flowers yellow, solitary, 3-6(-8) mm in diameter; pedicels slightly sulcate, glabrous, 2-12 mm long. Sepals 5, broadly ovate, navicular, 1.5-2 mm long, ca. 1.5 mm broad, glabrous. Petals 5, elliptic, ascending, lustrous, 2-3 mm long, 1-1.5 mm broad, obtuse or truncate at the apices; claws ca. 0.5 mm long, not thickened; nectaries cup-shaped, ca. 0.3 mm in diameter, borders of the cups very slightly projecting, not concave on the abaxial sides and somewhat vestigial. Anthers 0.2-0.4(-0.6) mm long, basifixed and latrorse; filaments flattened. Fruiting heads broadly ovoid, 2-3 mm in diameter, 2 mm long; receptacles glabrous. Achenes broadly obovate, lenticulate, ca. 1.0 mm long, glabrous; beaks 0.3-0.5 long, straight.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Sabha Distr., Chhurchathanga - Thudam, alt. 3600-3400 m (H. Ohashi et al. 772260,774983, Aug. 11,1977, TI); Thudam-Lamni Nama, alt. 3400-4200 m (H. Ohashi et al. 775067, 770731, Aug. 14, 1977, TI); around Lamni Nama, alt. 4200 - 4900 m (H. Ohashi et al. 775097, Aug. 15, 1977, TI); Lamni Nama - Tasagon, alt. 4200-4350 m (H. Ohashi et al. 770838, Aug. 16, 1977, TI); Tasagon-Topke Gola, alt. 4350-3600 m (H. Ohashi et al. 772542, Aug. 17, 1977, TI); Topka Gola, alt. 3600 m (H. Kanai et al. 720614, June 19, 1972, TI). Makalu Base Camp, alt. 4800 m (M. Suzuki et al. 8850650, 8850934, July 26, 1988, TI). Janakpur Zone. Ramechhap Distr., Jata Pokhari-Panchi Pokhari, alt. 4220-4600 m (H. Ohba et al. 8570488, July 15, 1985, TI). C NEPAL. Dhaulagiri Zone. Throng Phedi -Throng La, alt. 4700 m (H. Ohba et al. 831075, July 25, 1983, TI).

In the protologue of this species, Handel-Mazzetti (1939) stated that it resembled R. gmelinii DC., which occurs widely in the arctic and subarctic regions. In appearance, R. pegaeus is rather similar to R. hyperboreus, which is also a circumpolar arctic element and extends to the northernmost region. However, R. pegaeus is clearly distinguished from R. hyperboreus by the nectary shape and the shape of the beak of the achenes.

In R. hyperboreus, the nectary scales are forked or circular and surround the nectary pits. This type of nectary is also found in R. gmelinii and is characteristic of Sect. Hecatonia (Lour.) DC. In R. pegaeus, the nectary scales are laterally connate to the ventral sides of the petal laminae and result in cup-shaped nectary pits or pockets, as in members of Sect. Auricomus Spach. Ranunculus hyperboreus has achenes with linear stigmatic surfaces and very short and strongly curved beaks. In R. pegaeus, on the other hand, the stigmatic surface is spot-like and the beaks are long and straight. The similarity of R. hyperboreus to R. pegaeus is therefore only superficial.

This superficial resemblance of the two species (e.g., dwarf, creeping and stoloniferous habit) is considered to be the result of parallel evolution; adaptation to severe environmental conditions in the arctic region and in the alpine zone of the Himalaya.

Var. curvistylis Kadota, var. nov. [Fig. 8 d]

Caules, folia et pedicelli sparsim sericeis. Rostra achenia valde recurvata. Anthera 0.7-1.9 mm longa, filamentis 1.6-1.8 mm longis.

Specimens examined. E NEPAL. Janakpur Zone. Ramechhap Distr., around Baula Pokhari, 86°23'E, 27°40'N, alt. 3960 m (H. Ohba et al. 8570354, July 10, 1985, Holotype, TI).

Stem, leaves and pedicels sparsely sericeous with yellowish, appressed, smooth-surfaced curly hairs. The specimens numbered 'H. Ohba et al. 8570354' are composed of 45 individuals. There is no significant variation in the diagnostic characters throughout the individuals.

8) Ranunculus pseudopygmaeus Hand.-Mazz. in Acta Hort. Gothob. 13: 161, tab. 1, fig. 14 (1939)-Hara, Fl. E Himalaya 3: 37 (1975); in Hara & Williams, Enum. Fl. Pl. Nepal 2: 20 (1979)-L. Liou in Fl. Reip. Pop. Sin. 28: 282, pl. 89, figs. 6-7 (1980); in Fl. Xizang. 2: 102 (1985). [Fig. 9]

Ranunculus hyperhoreus auct. non Rottb.: Franch., Pl. Delav. 21 (1889)-Hand.- Mazz., Symb. Sin. 7: 309 (1931), p.p.

Dwarf perennial herb. Roots fusiform. Stem erect or ascending, 1-6 cm long, simple, densely sericeous with appressed hairs; collar clothed with fibrous remains of leaf bases. Basal leaves usually 3-5; blades reniform to broadly ovate, 4-9 mm long, 3-9 mm broad, (shallowly-)medially-deeply 3-lobed, sericeous with appressed hairs on both sides; middle lobes ovate to oblanceolate, entire or tricuspidate; lateral lobes entire or shallowly incised; leaf bases round-truncate-shallowly cordate; petioles 1-2.5 cm long, sparsely sericeous or glabrous, vaginate at the basal parts. Cauline leaves 1-3. Lower cauline leaves similar to the basal in shape, shortly petioled. Upper cauline leaves deeply 3-lobed, sessile; lobes linear. Flowers yellow, solitary, 4-9 mm in diameter; pedicels sulcate, 5-25 mm, pubescent with appressed and ascending hairs and/or patent hairs. Sepals patent, oblong, 3-5 mm long, navicular, hirsute with ascending hairs on the abaxial side. Petals 5, obovate to broadly obovate, 4-5 mm long including claws, entire or very slightly emarginate; claws 0.5-1 mm long; nectaries small cup-shaped. Anthers 0.6-0.8 mm long, basifixed and latrorse; filaments not dilated or slightly dilated. Fruiting heads ovoid-globose, 4-5 mm long; receptacles glabrous. Achenes broadly ovoidal, slightly inflated, ca. 1 mm long, smooth and glabrous; beaks ca. 0.5 mm long, straight but incurved at the apical parts.

Specimens examined. E NEPAL. Mechi Zone. Taplejung Distr., Kokim Pokhari, alt. 3200 m (H, Kanai et al. 720313, June 11, 1972, TI). Koshi Zone. Sankhuwa Sabha Distr., Merek, alt. 4350 m (M. Suzuki et al. 8850595/3, July 21, 1988, TI); Merek, alt. 4400 m (M. Suzuki et al. 8850610, July 22, 1988, TI).

Ranunculus pygmaeus of the circumpolar arctic region superficially resembles this species in size (dwarf) and nectary shape (cup-shaped). However, the former differs from the latter in root shape and hairiness of the plant body: in R. pygmaeus, the roots are not fusiform but filiform and the plants are almost glabrous. Additionally, the anthers of R. pygmaeus are dorsifixed and have dorsiventrality.

9) Ranunculus pulchellus C. A. Mey. in Ledeb. Fl. Altai. 2: 333 (1830); in Ledeb., Icon. Pl. Fl. Ross. 2: tab. 111 (1830)-Pampan., Fl. Caracorum 113 (l930)-Ovcz., Fl. URSS 7: 374 (1937)-Hand.-Mazz. in Acta Hort. Gothob. 13: 146 (1939)-Kitagawa in J. Jap. Bot. 31: 306 (1956)-Popov, Fl. Sred. Sibir. 1: 256 (1957)-Tamura in Kitamura, Fl. Pl. W Pakist. 65 (1964)-N. T. Liou et al. in Fl. Pl. Herb. Chinae Bor.-Orient. 3: 193, pl. 83, figs. 3-4 (1975)-Hara in Hara & Williams, Enum. Fl. Pl. Nepal 2: 20 (1979)-L. Liou in Fl. Reip. Pop. Sin. 28: 270, pl. 86, figs. 1-2 (1980)-Polunin & Stainton, Fl. Himal. 10, 445-5 (1984). [Fig. 10]

Ranunculus affinis R. Br. var. stracheyanus Maxim., Fl. Tangut. 14 (1889) -Ranunculus pulchellus C. A. Mey. var. stracheyanus (Maxim.) Hand.-Mazz. in Acta Hort. Gothob. 13: 147, fig. 4, 1-8 (1939), p. p.-Mukerjee in Bull. Bot. Surv. Ind. 2: 104 (1960)-Tamura in Acta Phytotax. Geobot. 23: 33 (1968); in Acta Phytotax. Geobot. 23: 106 (1968)-Hara, Enum. Fl. Pl. Nepal 2: 20 (1979)-L. Liou, Fl. Reip. Pop. Sin. 28: 271 (1980); in Fl. Xizang. 2: 99 (1985).

Ranunculus petrogeiton auct. non Ulbr.: Tamura in Acta Phytotax. Geobot. 23: 32 (1968).

Ranunculus potanini Kom. in Rep. Sp. Nov. 9: 392 (1911)-Ranunculus pulchellus C. A. Mey. var. potanini (Kom.) Hand.-Mazz. in Acta Hort. Gothob. 13: 147 (1939)-Mukerjee in Bull. Bot. Surv. Ind. 2: 104 (1960)-Tamura in Acta Phytotax. Geobot. 23: 33, fig. 4, 9-11 (1968)-Hara. Enum. Fl. Pl. Nepal 2: 20 (1979).

Ranunculus pulchellus C. A. Mey. var. geniculatus Hand.-Mazz., Symb. Sin. 7: 305 (1931); in Acta Hort. Gothob. 13: 147 (1939)-Ranunculus longicaulis C. A. Mey. var. geniculatus (Hand.-Mazz.) L. Liou in Fl. Reip. Pop. Sin. 28: 269 (1980).

Perennial herb. Roots fusiform. Stem erect, ascending or sometimes decumbent, 3-35 cm long, simple or branched, sericeous with appressed curly hairs. Basal leaves 1-5 or more; blades broadly obovate to broadly oblong or reniform, tricuspidate or medially (to deeply) 3-lobed with toothed lobes, or sometimes subentire with obtuse teeth or entire, 4-42 mm long, 4-50 mm broad, sparsely sericeous or almost glabrous on both sides; leaf bases round to subtruncate; petioles 0.5-15 cm long, clearly longer than the blades, vaginate at the basal parts. Lower and middle cauline leaves deeply 3-lobed, shortly petioled. Upper cauline leaves narrowly lanceolate or linear, entire, sessile. Flowers yellow, solitary to several, 5-12 mm in diameter; pedicels sulcate, 0.5-8 cm long, sericeous. Sepals 5, patent, purplish along the margins, oblong, 3-6.5 mm long, sericeous on the abaxial sides. Petals 5(-8), obovate to broadly obovate, entire or slightly emarginate, including claws 4-9 mm long, 2-8 mm broad; claws 0.5-1 mm long; nectaries cup-shaped with slightly projected borders or rather forked. Anthers 0.8-1.6 mm long, basifixed and latrorse; filaments flattened. Fruiting heads cylindrical to ovoidal, 3-5 mm in diameter, 4-9 mm long; receptacles glabrous. Achenes ovoidal, lenticulate, 1.3-1.9 mm long, smooth and glabrous; beaks ca. 0.7 mm long, straight, recurved or incurved at the proximal parts.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Sabha Distr., Chhurchathanga -Thudam, alt. 3600-3400 m (H. Ohashi et al, 774980, Aug. 11, 1977, TI). Shipton Pass, alt. 3960 m (M, Suzuki et al. 8850436, July 17, 1988, TI); Phematang Kharka, alt. 3300 m (M. Suzuki et al. 8850506, July 18,1988, TI); Yangri Kharka, alt. 3540 m (M. Suzuki et al. 8850882-8850884, July 19, 1988, TI) ; Ne Kharka, alt. 3650 m (M. Suzuki et al. 8850560-8850562, 8850894-8850899, July 20, 1988, TI); Merek, alt. 4400 m (M. Suzuki et al. 8850610, July 22, 1988, TI); Makalu Base Camp 4800 m (M. Suzuki et al. 8850654, July 26, 1988, TI); Ne Kharka, alt. 3660 m (M. Suzuki et al. 8850687, July 28, 1988, TI); Yangri Kharka, alt. 3450 m (M. Suzuki et al. 8820967, 8850679, July 28, 1988, TI); Unshina Kharka-Bhainsi Kharka, alt. 3030 m (M. Suzuki et al. 8850742, July 28, 1988,TI). Sagarmatha Zone. Solukhumbu Distr., Beni Kharka-Yulingolcha, alt. 3970-4900 m (N. Kurosaki 8572119, Aug. 31, 1985, TI). Janakpur Zone. Ramechhap Distr., Yalung Kharka-Pam Lhang, alt. 4750-4300 m (H. Ohba et al. 8351363, Sept. 8, 1983, TI).

This species occurs very commonly in meadows and in Abies-Rhododendron forests, in the subalpine and alpine zones of the Himalaya. It shows a wide range of inter- and intra-populational variation in gross morphology, adapting to the differences in environmental conditions of habitats. Several "varieties" have hitherto been recognized within this species. However, as stated above, the 'varieties', except for var. sericeus, should be included within var. pulchellus.

Handel-Mazzetti (1939) stated that var. stracheyanus and var. potanini are characterized by pilose pedicels with ferruginous hairs. It is true that most plants have sericeous pedicels with whitish appressed hairs. However, in my experience, the hairs change color from whitish to ferruginous during drying over firewood. The ferruginous color of the hairs is consequently considered to be the result of discoloration by heating during the drying of specimens. The difference in hair color is hence insignificant in the delimitation of infraspecific taxa.

There occur large populations of Ranunculus from place to place in the area ranging from 4100 m to 4400 m along Barun Khola in the Makalu region, E Nepal. These populations are generally composed of four entities. The first, second and third forms are R. pulchellus, R. pseudopygmaeus, and R. brotherusii, respectively. M. Suzuki et al. 8850566 (Lyang Mo Le Kharka, 4130 m) and M. Suzuki et al. 8850595/2 (Merek, 4350 m) belong to the fourth form, which is similar to R. pulchellus in appearance. However, the plants differ from R. pulchellus by the following way: 1) petals are absent or indefinite in number and/or size; 2) the anther sacs contain no pollen grains or contain pollen grains which are abnormal in shape and show significantly low stainability (e.g., 0%, 7% and 12%); 3) the nectaries are diminutive, shallow depressions and are vestigial; 4) the sepals are yellowish and more membranous. Consequently, we can consider these plants to be of hybrid origin. One of the putative parental species is undoubtedly R. pulchellus. The other parental species is thought to be R. pseudopygmaeus from the sericeous and medially to deeply 3-lobed basal leaves and the short elliptic anther sacs.

Var. sericeus Hook. f. et Thoms., Fl. Brit. Ind. 1: 17 (1872)-Maxim., Fl. Tangut. 13 (1889); Enum. Pl. Mong., 18-Tamura in Acta Phytotax. Geobot. 23: 106 (1968)-Hara in Hara & Williams, Enum. Fl. Pl. Nepal 2: 20 (1979)-Ranunculus membranaceus Royle ex D. Don, in Royle III. Bot. Himal. 1: 53 (1839)-L. Liou in Fl. Reip. Pop. Sin. 28: 269, fig. 36 (1980); in Fl. Xizang. 2: 98, fig. 27, 2-3 (1985)-Ranunculus pulchellus C. A. Mey. var. membranaceus (Kom.) Mukerjee in Bull. Bot. Surv. Ind. 2: 104 (1960). [Fig. 11]

Ranunculus longicaulis C. A. Mey. in Ledeb., Fl. Altai 2: 308 (1830); in Ledeb., Icon. Pl. Fl. Ross. 2: tab. 117 (1830)-Ovcz. in Fl. URSS 7: 375 (1937)-Ranunculus pulchellus var. longicaulis (C. A, Mey.) Trautv. in Bull. Soc. Nat. Mosc. 33: 68 (1860)-L. Liou in Fl. Reip. Pop. Sin. 28: 267, pl. 86, fig. 6 (1980).

Ranunculus nephelogenes Edgw. in Trans. Linn. Soc. 20: 28 (1846)-Ranunculus longicaulls C. A. Mey. var. nephelogenes (Edgw.) L. Liou in Fl. Reil. Pop. Sin. 28: 269 (1980); in Fl. Xizang. 2: 98, fig. 27, 1 (1985).

Stem erect to ascending or divaricate, 5-12 cm long, densely or sparsely sericeous with appressed whitish hairs. Blades of the basal leaves lanceolate to narrowly oblong to narrowly obovate, 2-25 mm long, 2-10 mm broad, entire to tricuspidate to deeply 3-lobed, usually densely sericeous on the abaxial sides and along the margins and sparsely sericeous on the adaxial sides, or almost glabrous on both sides; leaf bases cuneate to narrowly round. Flowers 10-14 mm in diameter. Fruiting heads ovoidal to narrowly ovoidal, 3-5 mm long; receptacles pilose with whitish curly hairs or glabrous. Achenes ca. 1.0 mm long.

Specimens examined. E NEPAL. Koshi Zone. Sankhuwa Saba Distr., Merek, alt. 4380 m (M. Suzuki et al. 8850603, 8850916-8850920, July 22, 1988, TI); Merek, alt. 4350 m (M. Suzuki et al. 8850605, July 22, 1988, TI); Merek, alt. 4400 m (M. Suzuki et al. 8850676, 8850942, July 27, 1988, TI); Makalu Base Camp, alt. 4780 m (M. Suzuki et al. 8850655, July 26, 1988, TI). Janakpur Zone. Ramechhap Distr., Yalung Kharka-Pam Lhang, alt. 4750-4300 m (H. Ohba et al. 8351364, Sept. 8, 1983, TI).

Var. sericeus was originally characterized by the "densely silky leaves." However, the leaf indumentum shows a considerable range of variation. In fact, plants bearing almost glabrous leaves have been collected (M. Suzuki et al. 8850655). This variety, therefore, is discriminated from var. pulchellus not by leaf indumentum, but by the shape of the basal leaf blades and bases and by the size of the fruiting heads and achenes. Handel-Mazzettii (1939) illustrated eight basal leaves as "var. stracheyanus." The leaf shape shown in Fig. 4(8), however, do not agree with his description "Folia basalia ambitu late ovata vel reniformia." This individual should be ascribable to var. sericeus.

This variety and the typical variety do not occur sympatrically in the Makalu region. Var. sericeus is restricted to rather xeric alpine meadows, while var. pulchellus prefers moist habitats. The two are therefore segregated ecologically.

Acknowledgments

I wish to thank Drs. H. Ohba and D. E. Boufford for critically reading the manuscript and correcting the Latin descriptions. I am grateful to Drs. T. Tateoka, H. Kanai, T. Yamazaki and K. Kosuge, for their kind suggestions and encouragement. My greatest thanks are due to the members of the Botanical Expedition to the Himalaya in 1988. Thanks are also given to the curators of the Herbaria (TI, KATH, CAL and TNS) for their kindness in permitting me to examine specimens.

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