Bull. Natn. Sci. M us., Tokyo, Ser. B, 26( 3), pp. 75- 93 , September 22, 2000

Saxifraga brachypoda D. Don and its Allies Collected in the
Sino-Himalayan Region (Saxifragaceae)
Shinobu Akiyama 1, Hideaki Ohba 2 and Su-Kung Wu 3
1

Department of Botany, National Science Museum,4- 1- 1 Amakubo, Tsukuba, Ibarak i 305- 0005, Japan2

E-mai l: ak iyama@ kahaku .go.jp
Department of Botany, University Museum, Uni versity of Tokyo
7- 3- 1 Hongo, Bunkyo-ku, Tokyo 11 3- 0033, Japan
E-mail : ohba@ um .u-tokyo.ac.Jp
3
Herbarium. Kunming Institute of Botany, Academia Sinica,
Kunming, Yunnan 650204, Ch ina

Abstract Saxifi~aga brachvpoda D. Don , S. wallichiana Sternb. and S. wardii W.W.
Sm. a re distinguished by the shape of the petals and the leaves and the presence (orabsence) of axillary buds. Two new combinations are proposed in S. brachypoda,
var. eglandulosa and var. gouldii .

Key words:

Saxifi~aga , Himalaya, Yunnan

Introduction
The species diversity in sections and subsections Ciliatae, Serpyllifoliae, Rosulares and Flagellares in the genus Saxifraga is one of the remarkable features of the
Sino-Himalayan alpine flora . Since Don ( 1821) published 'A Monograph of theGenus Saxifraga' , a great number of species have been described and revised fromthis region (Seringe, 1830; Sternberg, 181 0; 1822; 183 I; Franchet, 1889- 1890; Engler and lrmscher, 1912; 1916 & 1919; Smith, 1913 ; H. Smith, 1924; 1958; 1960;
Marquand, 1929; Handei-Mazzetti , 1931 ; Fischer, 1940; Grierson, 1987; Pan, 1991;
1992; etc.).
To understand chromosome evolution and morphological diversity in Saxifragain the Sino-Himalayan alpine flora, we conducted field research in the Himalaya andSW China since 1983 (e.g. Wakabayashi & Ohba 1988).
Saxifi··aga brachypoda D. Don and its allies are characterized by their caespitose,
simple stems which are leafy throughout and often with scaly buds in the axils, theso litary (or sometimes a few) yellow flower, and the spinulose-dentate (often with agland apically) leaves. But the specific delimitation of this group is rather dificult.
Saxiji-aga brachypoda D. Don was described in 1821 on the basis of Wallich 's collection from Nepal, and in 1830 S. fimbria fa Ser. (non D. Don) and S. g landulosa Ser.

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Shinobu Akiyama et al.

were described also based on Wallich's collections. Sternberg (1831) proposed the
new name S. wallichiana for S. fimbriata Ser. Later S. wardii W. W. Sm. (1913), S.
megalantha Marquand (1929), S. gouldii C .E.C. Fisch. ( 1939), and S. gouldii var. eglandulosa H. Sm. (1960) were described from Yunnan, Tibet and Bhutan.
Recently Grierson ( 1987) treated S. wallichiana and S. gouldii var. eglandulosa
as synonyms of S. brachypoda , and S. wardii as a distinct species. In contrast Pan( 1992) treated S. wallichiana as a distinct species and S. gouldii var. eglandulosa as asynonym of S. wardii. The specific delimitation of these species is still controversial.
To clarify species delimitation an investigation of the morphological characters of S.
brachypoda , S. wallichiana, S. wardii, and S. gouldii (incl. var. eglandulosa) areneeded.
In 1999 two of the authors (S. A. and S.-K. W.) collected S. wardii at Mt. Meili
Xueshan, NW of Deqin ('Atuntsi '), near the type locality, which allowed us to compare the morphology of S. wardii in detail with S. brachypoda, S. wallichiana and S.
gouldii var. eglandulosa.

Observations
Stems. Saxifraga brachypoda and its allies are characterized by the caespitose,
simple stems that are leafy throughout. The stems are 4- 19 (- 26) cm long in S.
brachypoda, 8- 25 cm long in S. wallichiana , (7- ) I 0- 15 cm long in S. gouldii var.
gouldii and S. gouldii var. eglandulosa, and 5- 10 cm long in S. wardii. In some plantsof S. brachypoda collected in Bhutan, Sikkim, Darjeeling and E. Nepal stems withaxillary buds and/or axillary branches only on the basal parts were creeping.
Axillary buds. One to a few axillary buds are in the cauline leaf axils of S.
brachypoda, several to ea. I 0 or more are in the leaf axils of S. wallichiana . Saxifraga
wardii and S. gouldii var. eglandulosa have no axillary buds in the axils of cauline
leaves, but have them on the basal part of the stem . Some plants of S. brachypoda
collected in Bhutan, Sikkim, Darjeeling and E. Nepal also have no axillary buds inthe axils of cauline leaves, but have them on the basal part of the stem.
Leaves. The leaves of Saxifraga brachypoda and its allies are spinulose (oftenwith an apical gland on each spine). The leaves on the apical part of the stem arelanceolate, acuminate at the apex, subcordate to truncate at the base, and sparsely
spinulose on the margins and with or without glands inS. brachypoda (Fig. I a), narrowly obovate, acuminate at the apex, subcordate to truncate at the base, and moderately spinulose on the margins with or without glands in S. wallichiana (Fig. 1 d &e) , lanceolate, acute at the apex, subcordate to truncate at the base, and regularly
spinulose on the margins and with or without glands in S. gouldii var. eglandulosa,and ellipti c, acute at the apex, truncate to subcordate at the base, distinctly cartilaginous and regularly spinulose on the margins with or without glands in S. wardii (Fig.
I f). In some plants of S. brachypoda from Bhutan, Sikkim , Darjeeling and E. Nepal

Sax i/i'Ciga brachypoda and its Allies in the Sino-Himal aya

77

without axillary buds on the middle part of the stem the leaves are linear to linearlanceolate, acute at the apex, slightly attenuate at the base and sparsely spinulose on
the margins and with or without glands (Fig. I b & c). The margins are distinctly cartilaginous in S. vvardii, while marginal sipules are cartilaginous in S. brachypoda, S.
wa/lichiana and S. gouldii var. eglandu/osa.
The leaves on the middle part ofthe stem are largest, 7- 12 (- 15) mm long and(l - )1.5- 2.5(- 3)mm wide, and usually 3-veined inS. brachypoda, (6- )8- !Smm longand 2.5-4.5(- 5) mm wide, and with more than 7 veins in S. wallichiana, ea. I 0 mm
long and ea. 1.5 mm wide, and 3-veined in S. gouldii var. eglandulosa, and 9- 11 mmlong and 3- 5 mm wide in S. wardii. In some plants of S. brachypoda from Bhutan ,Darjeeling and E. Nepal without axillary buds on the middle part of the stem theleaves are 6- 12 mm long and 1- 1.5(- 2) mm wide, attenuate at the base and 3-veined .
Uppermost part of the stem. The uppermost part of the stem is densely glandular hairy. The length between the ultimate leaf and the flower is 8- 19(- 25) mm in
S. brachypoda, 3- 9 mm inS. wallichiana, 3- 6 mm inS. gouldii var. eglandulosa.
Sepals. The sepals are (3- )4- 5 mm long, (2- )2.5- 3.5 mm wide, ovate , glandular
hairy on the margins, and sparsely to moderately glandular hairy or nearly glabrouson the outer surface in S. brachypoda (including plants from Bhutan, Sikkim, Darjeeling and E. Nepal) (Fig. I g- j), (3- )3.5-4.5 mm long, 2- 3 mm wide, ovate to narrowly ovate, sparsely glandular hairy on the margins , and nearly glabrous or sparsely
glandular hairy basally in S. wallichiana (Fig . I k- n) , 4- 5 mm long, ea. 3 mm wide,
ovate to narrowly ovate, densely glandular hairy on the margins and outer surface in
S. gou/dii var. eglandulosa , and ea. 7.5 mm long, ea. 3.5 mm wide , narrowly ovate,
densely glandular hairy on the margins and outer surface inS. wardii (Fig. I o).
Petals. The petals are 5- 8 mm long, 3.5-5 mm wide, elliptic to ovate, obtuse at
the apex , slightly attenuate at the base, very finely glandular and laciniate on the margins, glabrous on both surfaces, and without warts inS. brachypoda (including plantsfrom Bhutan, Sikkim, Darjeeling and E. Nepal) (Fig. I p- s) , 6.5- 8 mm long, 3.5-4mm wide, elliptic to obovate, obtuse at the apex, clawed or attenuate at the base,glabrous on both sides, and with 2 warts in S. wallichiana (Fig. I t- w) . The margins
are very finely laciniate in Nepalese plants (Fig. I t & u) but entire in Chinese plants(Fig. I v & w). The petals are 8- 10 mm long, 3- 5 mm wide, elliptic, obtuse at theapex , attenuate at the base, very finely to distinctly laciniate on the margins, glabrouson both sides, and without warts in S. gouldii var. eglandulosa, and ea. 11 mm long,ea. 7 mm wide, obovate, slightly retuse at the apex , attenuate at the base, distinctly
glandular (brown) on the margins , glabrous on the inner surface, sparsely glandular
hairy on the outer surface basally, and without warts inS. wardii (Fig. I x).
Stamens. The stamens are longer than or nearly equal to the pistils, and theanthers are not shriveled (with pollen) in S. brachypoda, S. wallichiana, S. gouldii
var. eglandulosa and S. wardii. In some flowers of S. brachypoda and S. wallichiana
the stamens are shorter than the pistils (nearly half) , and the anthers are shriveled and78

Shinobu Akiyama et al.

a

b .

c

Saxifi·aga brachypoda and its Allies in the Sino-Himalaya

79

without pollen.

Discussion
Engler and Irmscher ( 1916) described S. brachypoda as having the minutely
glandular-ciliate petal margins, but Fischer ( 1939) wrote that 'Don in the original description in Trans. Linn. Soc. 13, 378 makes no mention of such glands, nor can I
find any in the Wallich sheets in the Kew Herbarium .. .' Later, the petals of S. brachypoda were drawn to be entire (Pan, 1992). From this study it is clear that the petals of
S. brachypoda are very finely glandular or laciniate in Nepalese plants (Fig. I p & q).
It is uncertain that this character is specific or shows some infraspecific variation, as
in S. wallichiana (mentioned below) because we have inadequate materials from
China to resolve this issue.
Saxifraga wallichiana is distinguished from S. brachypoda by the presence ofwarts, as pointed by Pan (1992). Both Chinese and Nepalese S. wallichiana have thepetals with two warts, but the margins of the petals are different. In Chinese plants
the margins are entire (Fig. I v & w) , while in Nepalese plants they are very finely
laciniate (Fig. I t & u) . In other characters, the Chinese and Nepalese plants cannot
be distinguished. Grierson 's treatment of S. wallichiana as a synonym of S. brachypoda is not acceptable and these two species should be treated as distinct.
When Smith (1960) described S. gouldii var. eglandulosa he mentioned that
'The new variety agrees perfectly with the typical one except that the upper part ofthe petal is finely laciniate instead of bearing marginal glands .... Yar. eglandulosa
may thus be geographically distinct; or it may represent the normal condition of thespecies, and var. gouldii a rare modification.' We had an opportunity to examine thetype specimen and the para type of S. gouldii var. eglandulosa. The size of the plants
and the shape of the leaves, the sepals and the petals of S. gouldii var. eglandulosa are
indistinguishable from those of S. brachypoda, but the former can be distinguished by
the absence of axillary buds in the middle and apical part of the stems. As mentioned
in the introduction Grierson treated S. gouldii (incl. var. eglandulosa) as a synonymof S. brachypoda, while Pan teated it as a synonym of S. wardii. Consquently it is reasonable to treat S. gouldii var. eglandulosa as conspecific with S. brachypoda, but asa distinct variety, var. eglandulosa . Some plants of S. brachypoda collected in
Bhutan, Darjeeling and E. Nepal have linear to linear-lanceolate leaves and are charFig. I . Saxifi·aga brachypoda (a, g, h, p & q), S. brachypoda var. eglandulosa (b , c, i, j , r & s ), S.
wallich.iana (cl, e, k- n & t- w) and S. wardii (f, o & x). a-f: Cau lin e leaves near stem apex. go: Sepals. p- x: Petals. a, g & p: Nepa l (Wakabayashi et al. 9720272). h & q: Nepal (Miyamoto et al. 9440 121 ). b, i & r: Nepal (Ohashi et al. 775509). c, j & s: Darjeeling (Hara, 17 Sept.
1964). cl, k & t: Nepal (M iyamoto et al. 9584152). I & u: Nepal (Wakabayashi et al.
9720186). e, m & v: Ch ina (Wu et al. 754). n & w: China (Kato et al. 1849). f, o & x: China(Wu et al. 103526). All X5.

80

Shinobu Aki yama et al.

acterized by the absence of axillary buds on the middle parts as in var. eglandulosa .
These cannot be distinguished from the latter.
Saxifraga gouldii, characterized by the petals ' decorated with a single row ofsmall , sessile or shortly stipitate black glands which are continued for a varying distance along the sides' (Fischer, 1939), is known from only Bhutan and Tibet. According to the orginal description , S. gouldii has stems up to 26 cm long and 'broadly ensiform ' leaves (6- 15mm long, 2- 3mm wide). We examined the type and paratype
specimens and noted that with the exception of the marginal black glands of thepetals S. gouldii does not differ from S. brachypoda var. eglandulosa. It is therefore
reasonable to treatS. gouldii as another variety of S. brachypoda.
Saxifraga wardii is characterized by the glandular margins of the petals and theabsence of axillary buds in the cauline leaf axils as mentioned by Grierson ( 1987)
and Pan ( 1992). This species is also characterized by the short stems (5- I 0 cm long)and the size and shape of the leaves (9- 11 mm long, 3- 5 mm wide , elliptic, acute at
the apex , truncate to subcordate at the base, distinctly cartilaginous and regularly
spinulose on the margins and with or without glands). The treatment by Grierson( 1987) and Pan ( 1992) of S. wardii as a distinct species, is acceptable.
Taxonomic treatment
Saxifraga brachypoda D. Don in Trans. Linn. Soc. 13: 378 (1821 ); Prodr. Fl.
Nepal.: 209 ( 1825). Clarke in Hook . f. , Fl. Brit. lnd. 2: 396 ( 1878). Engler & lrmscher in Notes Roy. Bot. Gard. Edingurgh 5(24): 140 (1912); in Bot. Jahrb. Syst. 48 : 591( 1912); in Engl. , Pftanzenr.IV-117 (Heft 67): 136 (1916). Hand.-Mazz., Symb. Sin .
7(2): 422 ( 1931). Hara, FI. E. Himal.: 116 (1966); in Hara & Williams , Enum . Flow.
PI. Nepal 2: 151 (1979). J. T. Pan in Acta Phytotax. Sin . 16(2): 14 (1978); in Acta
Bioi. Plateau Sin. 1: 28 ( 1983); in Wu (ed.), Index Fl. Yunnan . I: 234 ( 1984); in Wu(ed.), Fl. Xizang. 2 : 466, fig. 154 18- 23 ( 1985); in Fl. Reipubl. Popularis Sin. 34(2):145 , fig . 36 I 0- 15 ( 1992). Polunin & Stainton, Fl. Himalaya : 132, 46 7 ( 1984 ). Grierson in Grierson & Long, Fl. Bhutan 1(3): 503 (1987). Stainton, Fl. Himalaya Sup.:20, PI. 38 (1988). Ohba & Akiyama, Alp. Fl. Jaljale Him. : 30 (1992). Type: Nepal(Wallich, Herb. Lambert [not found, see Fischer ( 1939)]). [Figs. I a, g, h, p, q, and 2]
Saxifraga glandulosa Wall. [Numer. List: 442 ( 1829), nom. nud.] ex Ser. in DC. ,Prodr. 4: 45 (1830). Types : Nepal. Gossaingsthan [Gosainkund] (Wallich 442 , Herb.
DC.). Kumaon [Kamoon] (Wallich 442A , Herb. DC.) .
Specimens examined. China. Yunnan . Tali Range, lat. 25 °40'N, ll - 12000ft.
(Forrest 11688 , Sept. 1913 , TI); C hih-tse-lo, 3000m (Tsai 54153 , 4 Sept. 1933 , A) ;Mekong-Salwin Divide, Sewalongba, 3400 m (Yu 22515 , 26 Aug. 1938, A). Tibet.
Changi, 12000 ft. (Cutting & Vernay 119B, 24 Sept. 1935 , A) . Bhutan . Me La, 14000
ft. (Ludlow & Sherriff 450 , 8 Aug . 1933 , TI); Chomolhari lhakan , 4150 m (Nishioka ,6 Sept. 1966, Tl). Nepal. Gossaingsthan [Gosainkund] (Wallich 442 , Tl , isosyntype

Saxifinga brachypoda and its Allies in the Sino-Himalaya

Iv

Fig. 2.

81

It:

Saxifi-aga brachypoda D. Don. Nepal (Wakabayashi et al. 9720190, Tl ).

of Saxifraga glandulosa Wall. ex Ser.). Lamrak, 10000- 14000ft. (Dhwoj 167, in
1929, Tl); Jata Pokhin, 14000- 15000 ft. (Dhwoj 0624, in 1930, T I); Gosainkund,Thale Patil- Rock Shelter I, 3400 m (Kanai & Mall a 674 759, 24 Aug. 1969, TT); E.
Nepal, Piling La, 4350 m (Ohashi et al. 773679, 11 Aug. 1977, Tl); 4400 m (Ohashi

82

Shinobu Akiyama et al.

et al. 772285, 11 Aug. 1977, Tl); E. Nepal , Topke Gola-Jalang Chhyongo, 36004300 m (Ohashi et al. 772579, 20 Aug. 1977, TI); Janakpur Zone, Ramechhap Distr. ,Dubikharka- Baula Pokhari, 3 720-4000 m (Ohba et al. 8520044, 9 July 1985, Tl);
Janakpur Zone , Ramechhap Distr. , Thare Og- Thare Teng- Thare Og, 4150-4260 m(Ohba et al. 8580446, 24 July 1985, TI); Sagarmatha Zone, Solukhumbu Distr.,
Sete- Taktor, 2550- 3400 m (Ohba et al. 8530930 & 8580973 , 19 Aug. 1985, Tl);
Sagarmatha Zone, Solukhumbu Distr., Mopung- Thosa Kharka, 2800- 3670 m (Ohbaet al. 8581071 , 21 Aug. 1985, TI); Sagarmatha Zone, Solukhumbu Distr. , Thosa
Kharka- Beni Kharka (Dambuk), 4000 m (Ohba et al. 8520296, 22 Aug. 1985, Tl) ;3670-4350 m (Ohba et al. 8571945, 22 Aug. 1985, TI); Sagarmatha Zone,
Solukhumbu Distr., Beni Kharka- Tschokarma- Beni Kharka, 3970-4950 m (Ohba &Wakabayashi 8531050, 24 Aug. 1985, Tl); Sagarmatha Zone, Solukhumbu Distr. ,Beni Kharka- below DudhKund, 3970-4450 m (Ohba et al. 853110 I, 26 Aug. 1985,
Tl); Sagarmatha Zone, Solukhumbu Distr. , below DudhKund- Beni Kharka ,4450- 3970 m (Ohba & Wu 8531179, 30 Aug. 1985, TI); Sagarmatha Zone,
Solukhumbu Distr. , Pike Bhanjyang- Pike Dongshar, 3500- 4000 m (Ohba et al.
8531330, 6 Sept. 1985, TI) ; Koshi Zone, Sankhuwa Sabha Distr. , Milke Danda,
Chhippon- Gidde, 2980- 3500 m (Ohba et al. 9110117 , 19 July 1991 , Tl); Koshi
Zone, Sankhuwa Sabha Distr. , Jaljale Hima1, around Banduke, ea. 4150 m (Ohba et
al. 9110213, 25 July- 3 Aug. 1991 , Tl); Koshi Zone, Sankhuwa Sabha Distr. , Jaljal e
Hi mal , Banduke- Jomle, 4150-4000 m (Ohba et al. 9120213 , 4 Aug. 1991 , TI); Koshi
Zone, Sankhuwa Sabha Distr., Jaljale Himal , Jomle- Goja (Ohba et al. 9110307 , 5Aug. 1991 , TI); Koshi Zone, Sankhuwa Sabha Distr. , Jaljale Hi mal , Goja- Shuwan
Kharka (near Panch Pokhari), 4130-4300 m (Ohba et al. 9130264b, 6 Aug. 1991 ,Tl); Koshi Zone, Sankhuwa Sabha Distr., Jaljale Himal, Bomrang- Singoa Kharka,
4140-4630 m (Ohba et al. 9110426, 10 Aug. 1991 , TI); Koshi Zone, Sankhuwa
Sabha Distr., Jaljale Hi mal , Singoa Kharka- Pahakhola, 4140- 2400 m (Ohba et al.
9120382, 11 Aug. 1991 , TI ); Bagmati Zone, Rasuwa Distr., Paldol Base Camp- a
Kharka, 4200 m (Miyamoto et al. 9420150, 3 Aug. 1994, TI); Bagmati Zone, Rasuwa
Distr. , Ganesh Base Camp- a Kharka, 4190m (Miyamoto et al. 9440121 , 10 Aug.
1994, TI); Sagarmatha Zone, Solukhumbu Distr., around Dudh Kund, 4500 m(Miyamoto et al. 9592436, 25 & 26 Aug. 1995 , Tl); Sagarmatha Zone, Solukhumbu
Distr., around Tangnang, 4400 m (Wakabayashi et al. 9720190, 12 Aug. 1997, TI) ;Sagarmatha Zone, Solukhumbu Distr., Dudh Kund- Mosom Kharka , 4540 m (Wakabayashi et al. 9720272 , 25 Aug. 1997, TI). Kumaon [Kamoon] (Wallich 442A, TI ,isosyntype of Saxifraga glandulosa Wall. ex Ser.).
Note. Saxifraga brachypoda , which ranges across the Hima1aya from Tibet to
Yunnan , is common in Nepal but rather rare in Yunnan.
var. eglandulosa (H. Sm.) S. Akiyama, H. Ohba & S.K. Wu, comb. nov.
[Figs. 1b, c, i, j , r, s, 3, and 4]
Saxifraga gouldii C.E.C. Fisch . var. eglandulosa H. Sm . in Bull. Brit. Mus. (Nat.

Saxifi·aga brachypoda and its Allies in the Sino-Himalaya

Fig. 3. SaxijiT1ga brachypoda D. Don. var. eglandulosa (H. Sm.) S. Akiyama, H. Ohba & S.K.
Wu. Bhutan (Ludlow, Sherriff & Hicks 19731 , BM , holotype of S. gouldii C. E. C. Fisch. var.
eglandulosa l-1 . Sm.).

83

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Shinobu Akiyama et al.

I£

jlllljllllj

I WJ

\

I

y

Fig. 4. Saxifi·aga brachypoda D. Don. var. eglandulosa (H. Sm.) S. Akiyama, H. Ohba & S.K.
Wu. Darjeeling (Hara, 17 Sept. 1964, Tl).

Sax ifi'aga brachypoda and its Allies in the Sino-Himalaya

85

Hist.), Bot. 2(9) : 249 ( 1960). Type: Bhutan. Tolegang, Tsampa, 4050-4200 m (Ludlow, Sherriff & Hicks 19731 , holo-BM , iso-CAL).
Specimens examined. Bhutan. Tolegang, Tsampa, 4050-4200 m [ 13500- 14000
ft.] (Ludlow, Sherriff & Hicks 19731 , I 0 Sept. 1949, BM , holotype of S. gouldii var.
eglandulosa H. Sm.; CAL, isotype); Bumthang Chu, Ju La, 4200m [14000 ft.] (Ludlow, Sherriff & Hicks 16915a, 21 July 1949, BM , paratype of S. gouldii var. eglandulosa H. Sm.); Between Me La and Cho La, 4050 m [ 13500 ft.] (Ludlow, Sherriff &Hicks 21412, 21 Aug. 1949, A); Samtengang- Ritang, 1900- 2600m (Hara et al.
3417, 11 Apr. 1967, TI) . Sikkim . 12000ft. (Ribu in 1908, TI). Darjeeling. Sandakphu- Gatribas (Hara s.n ., 17 Sept. 1964, TI); Below Sandakphu, 3400 m (Hara s.n., 17
Sept. 1964, TI); Sandakphu, 3600 m (Hara s.n ., 17 Sept. 1964, TI). Nepal. E. Nepal ,Tinjuray, 2900 m (Hara et al. 6303270, 27 Oct. 1963, TI, TNS); Hati Sar- Minchin
Dhap, 2900 m (Hara et al. 6303271 , 28 Oct. 1963 , Tl); Minchin Dhap- Mul Pokhari(Hara et al. 630272, 29 Oct. 1963, Tl); Batasay- Bhuspate Danra (Hara et al.
6303273 , I Dec. 1963 , TI , TNS); Cahuke- Tute (Ohashi et al. 774161 , 1 Sept. 1977,
KATH, TI); Jhangdang Kharka- Thulo Tingsang, 3 150- 3200 m (Kanai et al. 672940,
24 Sept. 1970, KATH , Tl); Kuri- Charikot, 3250- 1950 m (Kanai et al. 674659, 29
Sept. 1970, KATH, TI) ; Above Sauwala Khola, 12000 ft. (Stainton, Sykes &Williams 4418, 15 Sept. 1954, TI).
Note . Variety eglandulosa is usually caespitose but sometimes the basal part ofthe stems are creeping. This variety is restricted to eastern Nepal, Sikkim, Darjeelingand Bhutan.
var. gouldii (C.E.C. Fischer) S. Akiyama , H. Ohba & S.K. Wu, comb. et stat.
nov.
[Fig. 5]
Saxifraga gouldii C. E. C. Fischer in Kew Bull. 1939: 664 ( 1940). Type: Bhutan.
Chira, 5100 m (Gould 1310, holo-K).
Specimens examined. Bhutan . Chira, 5100 m [ 14200 ft.] (Gould 1310, Aug .
1938, K, holotype of S. gouldii C.E.C. Fischer). Tibet. Gautsa to Phari , 4300-5200 m[12000- 14300ft.] (Gould 1467, 13 Aug. 1938, K, paratype of S. gouldii C.E.C. Fischer).
Note. Variety gouldii is characterized by the glandu lar margins of the petals. 1t is
restricted to Bhutan and Tibet.
Saxifraga wallichiana Stern b. , Rev. Saxifr. Suppl. 2: 21 , t. 22 ( 1831 ). J. T. Pan
in Acta Bioi. Plateau Sin. I : 28 (1983); in Wu (ed.) , Index Fl. Yunnan. 1:234 (1984);
in Wu (ed.), F1. Xizang. 2: 466, fig . !54 24- 29 (1985); in Fl. Reipubl. Popularis Sin.
34(2): 14 7, fig. 3 7 1- 6 ( 1992). Type: Same as Saxifragafimbriata Wall. ex Ser.
[Figs. Id, e, k- n, t- w, and 6]
Saxifragafimbriata Wall. [Numer. List: 443 (1829) , nom . nud.] ex Ser. in DC.
Prodr. 4 : 45 ( 1830), non D. Don ( 1821 ). Types : Nepal. Gossaingsthan [Gosainkund](Wallich 443 , Herb. DC.). Kumaon [Kamoon] (Wallich 443A, Herb. DC.).
Saxifraga brachypoda D. Don var.fi.mbriata (Ser.) Engl. & 1rmsch. in Bot. Jahrb.

86

Shinobu Akiyama et al.

;~..
Fig. 5. Saxifraga brachypoda D. Don. va r. gou/dii (C.E.C. Fischer) S. Akiyama, H. Ohba & S.K.
Wu. Bhutan. Ch ira, 5100m [14200ft.] (Gou ld 1310, Aug. 1938, K, holotype of S. gouldii
C. E.C. Fischer) (upper three plants).

Saxi/ivga brachvpoda and its A llies in the Si no- Hima laya

87

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£ 1'

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llllll llld

I

2

3

I

I I I I I

Fig. 6.

4

5

S. wallichiana Stern b. Nepa l (Wakabayashi et al. 9720 186, TI).

Syst. 48 : 591 ( 1912); in Engl. , Pflanzenr. IV- 117 (Heft. 67): 137 ( 19 16). J. T. Pan in
Acta Phytotax. Sin . 16(2): 14 (1978). Hara in Hara & Williams, Enum. Flow. PI.
Nepal2: 151 ( 1979).
Specimens examined. China. Yunnan. N.W. Yunnan, the eastern flank of the88

Shinobu Akiyama et al.

Lichiang Range, Lat. 27° !5 'N, II000- 12000ft. (Forrest 2867, Sept. 1906, TI); E.
flank of the Lichiang Range (Forrest 6627, in 1910, Tl); Chien-chuan-Mekong di vide, Lat. 26°30'N, Long. 99°44'E, 11000 ft. (Forrest 22211, Aug. 1922, TJ); Li jiang, Yulongshan (Yulong Xueshan), 3650-4000 m (Ohba et al. 517, 17- 26 Sept.
1987, KUN , TI); Lijiang, Yulongshan (Yulong Xueshan), 3700 m (Ohba et al. I 004,
17- 26 Sept. 1987, KUN , Tl); Lijiang County, Qilanbiao, Yulong Xueshan,
3500- 3900 m (Kato et al. 1849, I Aug. 1988, KUN , TI); Deqe : around Daxue Shan,
4050 m (Wu et al. 781, 29 Aug. 1996, KUN , TT). Sichuan. Xiangcheng Xian, around
Da Xue Shan (near Yunnan-Sichuan border), 3970-4420 m (Wu et al. 1115, 25 Sept.
1994, KUN , TI); Xiangcheng Xian, around Da Xue Shan (near Yunnan-Sichuan border) , 3970-4420 m (Wu et al. 1122, 25 Sept. 1994, KUN, TI); Daocheng, Haizi (nearDaocheng), 4000 m (Wu et aL. 698, 17 Aug. 1996, KUN , TI); Daocheng, around
Gongga Shan (Zhong Niuan, a pasture), 3960 m (Wu et al. 728, 21 Aug. 1996, KUN,Tl); Daocheng, around Gongga Shan, 4170 m (Wu et al. 754, 23 Aug. 1996, KUN ,TI); Daocheng, around Gongga Shan, 3920-4910 m (Wu et al. 2350, 24 Aug. 1996,
KUN, Tl). Sikkim. Tauqu, 13500 ft. (Smith & Cave 2559, 15 Aug. 1909, Tl). Nepal.
Karnali Zone, Marghor Lagna, 3600 m (Rajbhandari & Roy 4118, 29 July 1979, Tl);
Singum Gompa- Gosainkund, 3200-4200 m (Kanai et al. 723945 , 23 Aug. 1972, Tl);
Rolwaling Khola, Thandingma- Beding, 3200- 3600 m (Ohba et al. 8331759, 2 Sept.
1983, TI); Rolwaling Khola , Beding- Na, 3600-4050 m (Ohba et al. 8351292, 4 Sept.
1983 , Tl ); Janakpur Zone, Ramechhap Distr. , around Thare Og, 4150 m (Ohba et al.
8520170, 23 July 1985, Tl) ; Janakpur Zone, Ramechhap Distr., Thare Og- Phedi
Kharka-Thare Og, ea. 4150 m (Ohba 8530517, 26 July 1985, Tl); Janakpur Zone,
Ramechhap Distr., Thare Og- Neju, 3651-4877 m (Ohba et al. 8570928, 30 July
1985, TI); Janakpur Zone, Ramechhap Distr., around Neju, 3651 m (Ohba et al.
8520236, 31 July and I Aug. 1985, Tl); Janakpur Zone, Ramechhap Distr. , Nejuabove Luk Kharka- Neju, 3651-4600 m (Ohba 8530669, I Aug. 1985, TI); Janakpur
Zone, Ramechhap Distr., Neju- Choarma, 3651 - 2760 m (Ohba et al. 8530694 &
8580672, 2 Aug. 1985, Tl , TNS ; 8571122, 2 Aug. 1985, TI); Bagmati Zone, Rasuwa
Distr. , Yure Kharka- Tinbu Kharka , 3700 m (Miyamoto et al. 9420040, 26 July 1994,
TI); Sagarmatha Zone, Solukhumbu Distr. , Rato Odara- Chhomalang Base Camp,
4550 m (Miyamoto et al. 9584152, 11 Aug. 1995, Tl); Sagarmatha Zone, Solukhumbu Distr. , Chhomalang Base Camp- Rato Odara, 4500 m (Miyamoto et al. 9592319,
18 Aug. 1995, Tl); Dhawalagiri Zone, Mustang Distr., Chabarbu (Thorung Phedi)Jharkot, 4060 m (Mikage & Yonekura 9552444, 24 Sept. 1995, Tl); Sagannatha
Zone, Solukhumbu Distr. , around Tangnag, 4300 m (Wakabayashi et al. 9720186, 12
Aug. 1997, TI); Sagarmatha Zone, Solukhumbu Distr. , Khare- Tangnag, 4320 m(Wakabayashi et al. 18 Aug. 1997, TI). Garhwal. Valley of Flowers (Kobatake 96, 7
Aug. 1988, Tl).
Note. Saxifraga wallichiana occurs in the Himalaya in India, Nepal , Sikkim,
Tibet and Yunnan and is common in Nepal and Yunnan .

89Saxifi·aga brachypoda and its Allies in the Sino-Himalaya

II

I J...tJ

1/i'ttll.'' "'HERB. HORT. REG. BOT. EDIN.

,..;,. f· .K~.ll." ~Fig. 7. Saxifi'aga wardii W.W. Sm. China. N.W. Yunnan, near Atuntsi [Deqin], 14000 ft. (F.
Kingdon Ward 46, Sept. 1911, E, holotype of S wardii W.W. Sm.).

Shinobu Akiyama et al.

90r.,_~ 1 ~ · r,

(1,Oct.

~H~~A-~~... . ...

~' ·"··'j-):;{\;l%::At."',"')_:. ..,.

~ - ""·~

\

l!mDJ

;:f
,-f;.gi V/"1 l d,£,.. , 'f'tk.r,.<t(h,. (~-v

: ;--'j;M i-,AA-U <.""
lt<

6~'4""'tjJ

cf://_., NC.; f?'lFig. 8. Saxiji-aga wardii W.W. Sm. Tibet (F. Kingdon Ward 5979, K, holotype of S. megarantha
Marquand).

Saxiji-aga brachypoda and its Allies in the Sino-Himalaya

Fig. 9.

Saxifi-aga wardii W.W. Sm. China (Wu et al. 103526, TNS).

91

92

Shinobu Aki yama et al.

Saxifraga wardii W.W. Sm. in Notes Roy. Bot. Gard. Edinburgh 8: 134 (1913).
J. T. Pan in Acta Bioi. Plateau Sin. 1: 29 (1983); in Wu (ed.), Index Fl. Yunnan. 1:
246 ( 1984); in Wu (ed.), Fl. Xizang. 2: 468, fig. 154 7-12 (1985); in Fl. Reipubl.
Popularis Sin. 34(2) : 147, fig . 37 1-6 (1992). Grierson in Grierson & Long, Fl.
Bhutan 1(3): 505 ( 1987). Type: China. N. W. Yunnan, near Atuntsi [Deqin ], 14000 ft .
(F. Kingdon Ward 46, holo-E).
[Figs. If, o, x, and 7- 9]
Saxifraga megalantha Marquand in J. Linn. Soc. Bot. 48: 179 ( 1929). Type: S. E.
Tibet. Nam La, eastern extremity of the Himalaya, 4200 m (F. Kingdon Ward 5979,
holo-K, iso-CAL & E).
Specimens examined. China. Yunnan. (Yil 9295, A) ; N.W. Yunnan, near Atuntsi[Deqin] , 14000ft. (F. Kingdon Ward 46, Sept. 1911 , E, holotype of S. wardii W.W.
Sm.); A-tun-tze [Deqin] , Bai-mar-shan, 3500 m (Wang 69568, Sept. 1935); A-tun-tze[Deqin] , Huann-fu-ping, 3500 m (Wang 69053 , Aug. 1935, A); Mountains of Moting,northeast of the Yangtze-Mekong watershed, 14000 ft. (Rock I 0319, June 1923, A);
Deqin. Mt. Meili Xueshan, 4400 m (Wu et al. I 03526, 20 Aug . 1999, KUN , Tl ,TNS). Tibet. S.E. Tibet. Nam La, eastern extremity of the Himal aya, 4200m [14000
ft.] (F. Kingdon Ward 5979, K, holotype of S. megalantha Marquand; CAL & E, isotypes).
Note. Saxifraga wardii is ristricted to Tibet and Yunnan and is rather rare.
Acknowledgement
We are indebted to Dr. David E. Boufford, the Harvard University Herbaria, forreading the manuscript. We thank the curator of the following herbaria, A, BM, CAL,
E, K, KUN , TI and TNS) for allowing us to examine specimens in their care. We arealso grateful to Drs. Yu'ichi Kadota, National Science Musum , and Futoshi Miyamoto , Tokyo Agricultural University, who kindly gave us the photographs of the specimens of Saxifraga. This study was supported by a Grant-in-A id for Scientific Research (A) from the Japan Society for the Promotion of Science, no . 11691178 in
1999 and 2000 (to H. 0.).

References
Don, D. 1821. A monograph of the genus Saxij;"aga. Trans. Linn. Soc. 13: 341-452.
Eng ler, A. and E. lrm sc her. 1912. Plantae chinenses forrestianae. Enumeration and description of spec iesof Saxifraga and Be1genia. No tes Roy. Bot. Card. Edinbwgh 5(24): 123- 148, pis. 87- 102.
Engler, A. and E. lrmscher. 1916 & 1919. Saxiji"Gga . In: A. Engler (ed.), Das Pflanzenreich . Heft. 67( 19 16) and 69 ( 1919). Wilhelm-Engelmann, Berlin.
Fisc her, C. E. C. 1940. LXX- New plants from Bhutan and Tibet. Kew Bull. 1939: 663-665.
Fra nchet, A. R. 1889- 1890. Plantae Delavayanae [Saxifi·aga, pp. 232- 240] . Paul Klincksieck, Paris.
Grierso n, A. J. C. 1987. 6. Saxifi-aga. In Grierson , A. J. C. and D. G. Long (eds.), Flora of Bhutan 1( 3 ):492- 5 15. Royal Botanic Garden, Edinburgh.
Handel-Mazzetti, H. 1931. Symbolae Si nicae. 7(2). Verlag von Juliu s Springer, Wien. [Saxifi·aga, pp.

Saxi(raga brachypoda a nd its Allies in the Sino-Himalaya

93

4 16-427].
Marquand, M.A. 1929. Th e botanical collec tion made by Capta in F. Kingdon Ward in the eastern Himalaya a nd Tibet in 1924- 25 [Saxifi·aga]. J Linn. Soc. Bot. 48 : 176- 183.
Pan, J. -T. 1991. A study on the genu s Saxifi·aga L. from Ch ina. Acta Phy totax. Sin. 29( I ): 1- 24.
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Smith, H. 1924. Pl antae s in enses. I. The ge nus Saxiji'Gga. Acta Horti Gothob. 1: 1- 22, t. 1- 10.
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Smith, H. 1960. Saxifi·aga o f the H ima laya . 11 . Some new spec ies. Bull. Brit. M us. (Nat. His!.) Bot. 2(9):229- 260, pi. 13- 2 1.
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