Bull. Natl. Mus. Nat. Sci., Ser. B, 33(3 · 4), pp. 91–103, December 21, 2007

Inquiries about taxonomy of the Sino-Himalayan Saxifraga(Saxifragaceae)
Shinobu Akiyama1 and Hideaki Ohba2
1

Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba 305–0005, Japan
E-mail: akiyama@kahaku.go.jp
2
Department of Botany, University Museum, University of Tokyo,
Hongo 7–3–1, Bunkyo-ku, Tokyo 113–0033, Japan

Abstract The ranges of distribution for Saxifraga diapensia Harry Sm., S. dhwojii (Harry Sm.)S. Akiyama et H. Ohba, S. implicans Harry Sm. and S. jacquemontiana Decne. are presented. The
lectotype of Saxifraga diapensia Harry Sm. is here selected. Notes on Saxifraga sphaeradena
Harry Sm. and S. implicans Harry Sm. are provided. Saxifraga sphaeradena subsp. dhwojii Harry
Sm. is regarded as a distinct species, S. dhwojii (Harry Sm.) S. Akiyama et H. Ohba.
Key words : China, Himalaya, Saxifraga, Sino-Himalaya, Tibet

Saxifraga is one of the most diversified generain the Sino-Himalayan region, with more than
200 species. The Sino-Himalayan region is thecentre of diversification of sections Ciliatae andPorphyrion. The taxonomy of section Ciliatae isstill controversial.
We have examined specimens of Saxifraga
section Ciliatae from the Sino-Himalayan region
preserved in various herbaria worldwide since1991. From our observations we were able to determine the range of distribution of several socalled rare species, such as Saxifraga diapensia
Harry Sm. and S. jacquemontiana Decne.
Pan (1991, 1992) classified Saxifraga diapensia along with S. hookeri Engl. et Irmsch., S.
sphaeradena Harry Sm., S. parnassifolia D. Don
and S. bulleyana Engl. et Irmasch. in ser. Bulleyane of sect. Ciliatae. Pan et al. (2001) later
broke down the series without comment. We
have discuss the morphological differences andthe relationships of the species classified in ser.
Bulleyane by Pan (1991).
Saxifraga sphaeradena Harry Sm. and S. implicans Harry Sm., which were described on thebasis of Ludlow and Sherriff’s collections fromSE Tibet, have not been examined.

The delimitation of sect. Ciliatae
ser. Bulleyane
Pan (1991) established ser. Bulleyane in sect.
Ciliatae to include Saxifraga bulleyana Engl. et
Irmasch., S. hookeri Engl. et Irmsch., S. sphaeradena Harry Sm., S. parnassifolia D. Don and S.
diapensia Harry Sm. Pan (1992) followed this in
the Flora Reipublicae Popularis Sinicae. Pan etal., in the Flora of China, (2001) did not use any
subdivisions under sections in Saxifraga. Series
Bulleyana has not been critically studied.
Saxifraga bulleyana has numerous, not glaucous leaves spaced at close intervals, of which
the basal ones are caducous, and both the proximal and distal ones are nearly equal to the median ones in shape and size. The pedicels have both
short glandular hairs and brown curled hairs.
These features differ greatly from all otherspecies in the series, but approach those of S. auriculata Engl. et Irmsch., S. brachyphylla
Franch., S. hypericoides Franch., S. yezhiensis C.
Y. Wu and allied species.
Saxifraga diapensia Harry Sm. is unique inhaving leaves densely crowned at the base andpersistent until fruiting and in having solitary

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Shinobu Akiyama and Hideaki Ohba

flowers. Except for these two species, S. bulleyana and S. diapensia, all other species are
similar and morphologically approach S. diversifolia classified in sect. Ciliatae ser. Caveanae J.
T. Pan, S. implicans in sect. Ciliatae ser. Stellariifoliae (Engl. et Irmsch.) J. T. Pan, and S. dianxibeiensis in sect. Ciliatae ser. Heterocladoides J.
T. Pan.
1) Saxifraga diapensia Harry Sm.
Saxifraga diapensia (Fig. 1) has dense blackish brown tipped glandular hairs on pedicel andmargins of the cauline leaves and is characterized
by abaxially gibbous sepals and the relatively
long pedicel, and belongs to S. diversifolia
group. Saxifraga diapensia lacks brownish curledhairs.
Saxifraga diapensia was described by Harry
Smith based on specimens he collected in northern and northwestern Sichuan in 1924. Although
Smith compared his collections with Saxifraga
nigroglandulosa Engl. et Irmsch., S. caveana W.
W. Sm. described from Sikkim is more similar inhaving nearly leafless flowering stems. Also Saxifraga caveana shares S. diapensia with gibbous
sepals and flowering stems with a solitary flower.
Hara (1978) and Grierson (1987) united S. diapensia with S. caveana, but Pan et al. (2001) distinguished S. diapensia from S. caveana by thepetals with two callosities. Specimens collectedby Wu et al. 10367 and 103705 of S. diapensia in
Yunnan have ecallous petals. Smith (1924) illustrated an ecallous petal taken from Smith 3331,
which we select here as the lectotype of S. diapensia. A remarkable point is that S. diapensia
has ascending or suberect sepals while S.
caveana has reflexed sepals. Saxifraga diapensia
is distributed in Sichuan, Yunnan and Tibet,
while S. caveana occurs in S. and SE. Tibet,
Bhutan, Sikkim, and Nepal. At present we refrainfrom making taxonomic changes until further information is available. The more than 30 specimens of S. diapensia we examined show therange to be northeast of the Himalaya and theHengduan Mountains (Fig. 2).

Saxifraga diapensia Harry Sm. in Acta Hort.
Gothob. 1: 10 (1924). Pan in Fl. Reip. Pop. Sin.
34(2): 139 (1992). Pan et al. (2001) in Fl. China
8: 306 (2001).
[Figs. 1, 2]
Lectotype (here selected): N. Sichuan (North
Sze-ch’uan): Dongrergo, between snow patches,
4800–5000 m. Harry Smith 3331 (UPS).
Specimens examined: Tibet. Tze la, 15000 ft.,alpine turf and rocky slopes (Kingdon-Ward12246, 22 Aug. 1935, BM). S. Tibet. Truka La,
Mago, 15000 ft., in clumps amongst rocks etboulders on summit of pass (Ludlow and Sherriff
815, 5 Aug. 1934, BM). Kongbo, Dryang la,
14000 ft., on cliffs (Ludlow, Sherriff and Elliot
14281, 8 Aug. 1947, BM). Kongbo La, Chagul
Chu, 14000 ft., in large clump on rocky ledges(Ludlow and Sherriff 2398, 19 July 1936, BM).
SE. Tibet. Pero La, Tsangpo valley, 29°30N
95°0E, 15500 ft. (Ludlow, Sherriff and Taylor
5191, 9 July 1938, BM). Kongbo, Budi Tsepo La(Ludlow, Sherriff and Elliot 14417, 21 Aug.
1947, BM). Bimbi La, Tsari Chu, Tsari, 28°45N
93°26E, 14000 ft. (Ludlow Sherriff and Taylor
6327, 14 Oct. 1938, BM). Kongbo, Mira La,
Nyang Chu, 29°30N 94°15E, 15500 ft., on
open bloch boulder scree, in moss (Ludlow, Sherriff and Taylow 6071, 14 Aug. 1938, BM). Kongbo Prov., Mira La, Nyang Chu, Puchu, 29°30N
94°15E, 16000 ft. (Ludlow, Sherriff and Taylor6079, 16 Aug. 1938, BM). Kongbo Prov., Kulu
Phu Chu, near Paka, 29°16N 94°26E, 15–
16000 ft. (Ludlow, Sherriff and Taylor 6516, 23
Sept. 1938, BM). Kongbo Prov., Kucha La, Kulu
Phu Chu, near Paka, 29°15N 94°25E, on open
cliff edges (Ludlow, Sherriff and Taylor 5935, 25
July 1938, BM). Kongbo Prov., Pasum La, Drukla Chu, 30°28 N93°22E, 17000 ft., amongst
boulders (Ludlow, Sherriff and Taylor 6916,
BM). Kongbo, Lochen, 15500 ft. (Ludlow, Sherriff and Elliot 15674, 2 Sept. 1947, BM). Kongbo, Pungkar, 14500 ft., on rocks (Ludlow, Sherriff and Elliot 15539, 1 Aug. 1947, BM). Kongbo, Dosbong La, 13500 ft., on cliff (Ludlow,
Sherriff and Elliot 14368, 16 Aug. 1947, BM).
Kongbo. Dryang La, 14000 ft., on cliff (Ludow,
Sherriff and Elliot 14281, 8 Aug. 1947, BM).

Inquiries about taxonomy of the Sino-Himalayan Saxifraga

Fig. 1.

93

Saxifraga diapensia Harry Sm. (Wu et al. 103521, TNS).

Nambu La, Pome, 30°0N 94°30E, 13500 ft., onboulders (Ludlow, Sherriff and Taylor 6943,
BM). Sichuan (Sikang). Taofu distr., Mt. Yara,

north distr. in glacea granites, alt. ca. 4850 m (H.
Smith 11569, 29 Aug. 1934, BM). Between Taining (Ngata) and Maoniu (Ndröme), Zungkong

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Shinobu Akiyama and Hideaki Ohba

Fig. 2.

Geographic range of Saxifraga diapensia Harry Sm.

La, in fissuris rupium, alt. 4500–4700 m (H.
Smith 12563, 30 Sept. 1934, BM). Kangting(Tachienlu) distr., Tapaoshan, alt. 46–4800 m, infissuris rupium (H. Smith 11456, 22 Aug. 1934,
A, BM). Ngaala, Tsa-wa-rung, alt. 3800 m (C. W.
Wang 66076, Aug. 1935, A); loc. cit. 3850 m (C.
W. Wang 77036A, Aug. 1935, A). SW. Szechuan,
Mount Konka, Risonquemba, Konkaling, alt.
3960–5335 m (J. F. Rock 16835, Aug. 1928,
BM). Yunnan. Chungtien area (Forrest 30840,
BM). Doker-la, Mekong-Salween divide,
28°20N, 12600 ft., open stony pasture (Forrest
14573, Aug. 1917, BM). NW. Yunnan. Mountains of Moting, northeast of the YangtzeMekong watershed, 14000 ft. (J. F. Rock 10325,
June 1923, BM). Mount Habashan, north of
Ndaku, north of the Likiang Snow Range,
Yangtze drainage basin, 14000 ft. (J. F. Rock
9701, July 1923, A). N. flank of Haba Snow
Range, on open hill (K. M. Feng 2169, 24 Aug.
1939, A). Zhongdian Xian, Haba Xueshan, Haba
Hai, alt. 4400 m (Wu et al. 103671, 10 Aug.
1999, KUN, TI). Deqin Xian, Meili Xueshan,
Meilishi – Suola (Wu et al. 103521, 20 Aug.
1999, KUN, TI, TNS). Deqin Xian, Meili Xueshan, Suola, alt. 4720 m (Wu et al. 103705, 20

Aug. 1999, KUN, TI). Doker-la, A-tun-tze, alt.
3500 m (C. W. Wang 64938, 3–5 Aug. 1935, A).
Huann-fu-ping, A-tun-tze, alt. 3700 m (C. W.
Wang 68890, Aug. 1935, A). In rupestribus humosis mont. in reg. Gunghisn, alt. 4000 m (C.
Schneider 3024, Sept. 1914, A). Mekong-Salvin
Divide, Sila, mt. rocky slope, alt. 4200 m (T. T.
Yü 22283, 11 Aug. 1938, A); loc. cit., mt. rocky
slope, alt. 4000 m (T. T. Yü 22371, 16 Aug.
1938, A). Sine loc. (T. T. Yü 9340, A).
2) Saxifraga sphaeradena Harry Sm. andsubsp. dhwojii Harry Sm.
Smith (1960) described Saxifraga sphaeradena
based on the collections from Lukuthang, Magoin S. Tibet, near the border with Bhutan andcompared it with S. subamplexicaulis Engl. et
Irmsch. Pan (1991, 1992) and Pan et al. (2001)
reported S. sphaeradena from S. Tibet.
Saxifraga sphaeradena resembles S. implicans,
S. parnassifolia, S. hookeri and S. diversifolia in
habitat, shape of leaves, and other features. Fromthese, however, S. sphaeradena differs in having
ovate-cordate leaves, somewhat glaucous on thelower surface.
Smith (1960) described S. sphaeradena subsp.

Inquiries about taxonomy of the Sino-Himalayan Saxifraga

dhwojii from Jata Pokhni (probably an error of“Pokhari”), Nepal, based on Lall Dhwoj 0625
(BM). Subspecies dhwojii was distinguishedfrom subsp. sphaeradena by the clawed petals
with 2- or 4-callosities (Petala . . . in unguem distinctum 1 mm vel ultra longum contracta, elevatim 2- vel 4-callosa). All the specimens we examined have widely oblong or ovate clawed
petals with basal callosities (Figs. 3a, b) and differ from that of subsp. sphaeradena, which has

95

obovate, sessile, ecallous petals. The shape andpresence of callosities of the petals are constantand stable, and the differences between subsp.
shaeradena and subsp. dhwojii are sufficient toseparate them as different species. Thus, we regard subsp. dhwojii as a distinct species, S.
dhwojii (Harry Sm.) S. Akiyama et H. Ohba
(Fig. 4). Pan (1992) reported subsp. sphaeradena
to have petals with 4 callosities. Pan’s subsp.
sphaeradena is therefore considered to be S.

Fig. 3. Petals and sepals of Saxifraga dhwojii (a, b), S. parnassifolia (c), S. diversifolia (d, e) and S. implicans(f–h). a: F. Miyamoto et al. 9420151. b: F. Miyamoto et al. 9584275. c: H. Ohba et al. 8320716. d: H. Ohba
et al. 8351226. e: F. Miyamoto et al. 9592492. f: Wu et al. 103685. g: Wu et al. 764. h: Wu et al. 103679. All
5.

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Shinobu Akiyama and Hideaki Ohba

Fig. 4.

Saxifraga dhwojii (Harry Sm.) S. Akiyama et H. Ohba (F. Miyamoto et al. 9420051, TI).

dhwojii.
Saxifraga dhwojii is apparently a common
species on scree and gravelly slopes in alpine

meadows especially in E Nepal eastward from86°15E (Fig. 5). It is distinct and no transitional
or intermediate forms with other species haveInquiries about taxonomy of the Sino-Himalayan Saxifraga

Fig. 5.

97

Geographic range of Saxifraga dhwojii (Harry Sm.) S. Akiyama et H. Ohba

been found.
Saxifraga sphaeradena Harry Sm. in Bull.
Brit. Mus. Nat. Hist. 2(9): 235 (1960), as subsp.
sphaeradena. H. Hara in H. Hara et Williams,
Enum. Flow. Pl. Nepal 2: 154 (1978), as subsp.
sphaeradena. Grierson in Fl. Bhutan 1(3): 511(1987). Pan et al. in Fl. China 8: 297 (2001), as
subsp. sphaeradena.
[Figs. 3a, 3b, 4, 5]
Type: Tibet. SE Tibet, Lukuthang, Mugo, alt.
4050 m. Ludlow and Sherriff 807 (BM – holo).
Saxifraga dhwojii (Harry Sm.) S. Akiyama et
H. Ohba, stat. nov.
Saxifraga sphaeradena Harry Sm. subsp.
dhwojii Harry Sm. in Bull. Brit. Mus. Nat. Hist.
2(9): 236 (1960). H. Hara in H. Hara et Williams,
Enum. Flow. Pl. Nepal 2: 154 (1978). Grierson in
Fl. Bhutan 1(3): 507 (1987), in syn. of Saxifraga
parnassifolia D. Don. Pan in Fl. Reip. Pop. Sin.
34(2): 140 (1992). Pan et al. (2001) in Fl. China
8: 297 (2001).
Saxifraga sphaeradena Harry Sm. subsp.
sphaeradena auct. non Harry Sm: Pan in Fl.
Reip. Pop. Sin. 34(2): 140 (1992), excl. type.
Type: Nepal. Jata Pokhni, alt. 4200–4500 m.

Lall Dhwoj 0625 (BM – holo).
Specimens examined. Nepal. Dolakha Distr.,
Bitta Kharka – a valley – Patale Pokhari,
27°50N 86°23E (H. Ohba et al. 8331970 and
8351411, 12 Sept. 1983, TI). Dolakha Distr.,
Kyalche – Dongang – Thandingma, 27°54N
86°19E (H. Ohba et al. 8331720 and 8351202, 1
Spet. 1983, TI). Dolakha Distr., Na – Yalung
Kharka, 4500 m (H. Ohba et al. 8320655, 7 Sept.
1983, TI). Dolakha Distr., Pam Lhang – Bitta
Kharka, 27°50N 86°24E, alt. 4100 m (H. Ohba
et al. 8320691, 10 Sept. 1983, TI). Dolakha
Distr., Thandingma – Nimare – Beding, 27°54N
86°22E (H. Ohba et al. 8331774, 2 Sept. 1983,
TI). Manang Distr., Yak Kharkaa – Churi
Lattar – Thorung Phedi, 28°43N 83°58E, alt.
4060 m (M. Mikage et al. 9450144, 18 Aug.
1994, TI). Ramechhap Distr., around Neju,
27°44N 86°31E, alt. 3651 m (H. Ohba et al.
8520229 and 8570987, 31 July 1985, TI).
Ramechhap Distr., around Thare Og, 27°45N
86°28E (H. Ohba et al. 8520169, 8520174, and8580478, 25 July 1985, TI). Ramechhap Distr.,
Jata Pokhari – east side of Jata Pokhari – Botase
Kharka – Jata Pokhari, 27°43N 86°25E (M.
Wakabayashi 8520098, 16 July 1985, TI).

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Shinobu Akiyama and Hideaki Ohba

Ramechhap Distr., Thare Og – Thare Teng –Thare Og, 27°45N 86°28E (H. Ohba et al.
8580452, 24 July 1985, TI). Rasuwa Distr.,around Seto Kund, 28°16N 85°08E, alt. 3930 m(F. Miyamoto et al. 9420219, 10 Aug. 1994, TI).
Rasuwa Distr., around Tinbu Kharka, 28°11N
85°13E, alt. 3730 m (F. Miyamoto et al.
9420051, 27 July 1994, TI). Rasuwa Distr.,
Khodang
Danda – Gul
Bhanjyang – Latsu,
27°57N 85°29E (H. Hara et al. 729361, 29
Aug. 1972, TI). Rasuwa Distr., Paldol BaseCamp – a Kharka, 28°14N 85°12E, alt. 4300 m(F. Miyamoto et al. 9420151, 3 Aug. 1994, TI).
Rasuwa Distr., Singum Gompp – Gosainkund,
28°06N 85°21E (H. Hara et al. 723959, 23
Aug. 1972, TI). Rasuwa Distr., Thale Patil – Hile
Dhap – Mangen, 28°01N 85°30E (H. Hara etal. 723960, 27 Aug. 1972, TI). Sankhuwasawa
Distr., Goja (Goja Kharka) – Shuwan Kharka(near Panch Pokhari), 27°35 87°32 (H. Ohba et
al. 9110322, 6 Aug. 1991, TI). Sankhuwasawa
Distr., Jomle – Goja (Goja Kharka), 27°34N
87°30E (H. Ohba et al. 9110318 and 9110918, 5
Aug. 1991, TI). Sankhuwasawa Distr., Shuwan
Kharka (near Panch Pokhari) – Topke Gola,
27°37N 87°34E (H. Ohba et al. 9110363, 7
Aug. 1991, TI). Sankhuwasawa Distr., Shuwan
Kharka (near Panch Pokhari) – Topke Gola,
27°37N 87°34E (H. Ohba et al. 9120287, 7
Aug. 1991, TI). Solukhumbu Distr., around Beni
Kharka (Dambuk), 27°42 86°35 (H. Ohba et al.
8520304, 23 Aug. 1985, TI). Solukhumbu Distr.,around Pike Dongshar, 27°30N 86°27E (H.
Ohba et al. 8572429, 7 Sept. 1985, TI).
Solukhumbu Distr., around Tangna, 27°44N
86°51E, alt. 4500 m (F. Miyamoto et al.
9584226, 21 Aug. 1995, TI). Solukhumbu Distr.,
Beni – Lekhani – Tambakhani – Loding Khola –Loding, 27°32N 86°34E (H. Ohba et al.
8531305, 4 Sept. 19850, TI). Solukhumbu Distr.,
Beni Kharka – Sarkari Pati, 27°40N 86°35E(H. Ohba et al. 8531249, 8572159, 8572186, and8581286, 2 Sept. 1985, TI). Solukhumbu Distr.,
Beni Kharka (Dambuk) – Dikitscho – below
DudhKund, 27°40N 86°35E (H. Ohba et al.
8531093, 26 Aug. 1985, TI). Sokukhumbu Distr.,

Beni Kharka (Dambuk) – Dikitscho – upper
DudhKund, 27°40N 86°35E, alt. 4120 m (M.
Wakabashi et al. 8541190, 26 Aug. 1985, TI).
Solukhumbu Distr., Beni Kharka (Dambuk) –Tschokarma – Beni Kharka, 27°40N 86°35E(H. Ohba and M. Wakabayashi 8531054, 24 Aug.
1985, TI). Solukhumbu Distr., Dudh Kund –Hinku Khola – Thasing Dingma, 27°42N
86°50E, alt. 3770 m (F. Miyamoto et al.
9584275, 27 Aug. 1995, TI). Solukhumbu Distr.,
Horsola Kharka – Tangnag, 27°41N 86°49E,
alt. 3980 m (M. Wakabayashi et al. 9720182, 11
Aug. 1997, TI). Solukhumbu Distr., Loding –Pikebuk – Pike Bhanjyang, 27°32N 86°32E (H.
Ohba et al. 8520395, 8531315, and 8572316, 5
Sept. 1985, TI). Solukhumbu Distr., Mopung –Thosa Kharka, 27°40N 86°35E (H. Ohba et al.
8581058, 21 Aug. 1985, TI). Solukhumbu Distr.,
Mosom Kharka – Bhedi Kharka, 27°40N
86°49E, alt. 3740 m (M. Wakabayashi et al.
9720264, 22 Aug. 1997, TI). Solukhumbu Distr.,
Mosom
Kharka – Rangdu
Kharka – Hinku
Khola – Tashing Dingma, 27°40N 86°49E, alt.
3300 m (M. Wakabayashi et al. 9720288, 26
Aug. 1997, TI). Solukhumbu Distr., Pike Bhanjyang – Pike – Pike Dongshar, 27°32N 86°29E,
alt. 3700 m (H. Ohba et al. 8520398, 8531331,
8572383bis, 8581395, 8581400, 8591400, 6
Sept. 1985, TI). Solukhumbu Distr., Pike Dongshar – Ngawur – Goligaon – Namikhil, 27°30N
86°27E (H. Ohba et al. 8531364, 9 Sept. 1985,
TI). Solukhumbu Distr., Sete – Kata Bisana –Goyem – Lamjura Bhanjyang – Taktor, 27°34N
86°26E (H. Ohba et al. 8581016, 19 Aug. 1985,
TI). Solukhumbu Distr., Tangnag – Horsola
Kharka – Mosom Kharka, 27°43N 86°50E, alt.
4020 m (M. Wakabayashi et al. 9720248, 21
Aug. 1997, TI). Solukhumbu Distr., Thosa
Kharka – Khonglemo Deorali – Beni Kharka(Dumbuk), 27°36N 86°34E, alt. 3700 m (H.
Ohba et al. 8530986, and 8581108, 22 Aug.
1985, TI). Taplejung Distr., Bomrang (Bomrang
Kharka) – Pamphule Deurali – Singoa Kharka,
27°39N 87°33E (H. Ohba et al. 9110472, 10
Aug. 1991, TI).

Inquiries about taxonomy of the Sino-Himalayan Saxifraga

3) Saxifraga implicans Harry Sm.
Saxifraga implicans Harry Sm. (Fig. 6) isclosely related to S. hookeri Engl. et Irmasch.,

Fig. 6.

99

but distinguished by the reflexed sepals at flowering and the clawed petals with 6–8 or no callosities. In these two species the persistence of basal

Saxifraga implicans Harrry Sm. (Wu et al. 103685, TI).

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Shinobu Akiyama and Hideaki Ohba

Fig. 7.

Geographic range of Saxifraga implicans Harry Sm.

leaves is not stable. Saxifraga implicans approaches S. diversifolia also, but clearly differs inhaving more median and distal leaves, narrowly
ovate basal leaves with truncate base and subacute apex, and the clawed petals usually with callosities (Figs. 3f–h). Saxifraga parnassifolia D.
Don (Fig. 3c) differs from S. implicans, and also
S. hookeri and S. diversifolia (Figs. 2d, 2e) by thepetals with 2 or 4 thorn-like outgrowths near thebase and the sepals with veins always confluent
into verruca below the apex. Saxifraga implicans
can be distinguishable from S. subaequifoliata
Irmsch. by the extremely smaller distal cauline
leaves and inflorescences with less than 10 flowers. As a conclusion Saxifraga implicans is considered to be distinct species in the Ciliatae, distributed in Sichuan, Yunnan and Tibet (Fig. 7).
Saxifraga implicans Harry Sm. in Bull Brit.
Mus. Nat. Hist. 2(9): 231 (1960). Pan in Fl. Reip.
Pop. Sin. 34(2): 116 (1992), with var. weixiensis
C. Y. Wu. Pan et al. (2001) in Fl. China 8: 298(2001).
[Figs. 3f–h, 6, 7]
Type: Tibet. SE Tibet, Kongbo, Nyoto Sama,
13000 ft., in open forest amon rocks. Ludlow,
Sherriff et Elliot 15582, 10 Aug. 1947. (BM –holo).
Specimens examined. Tibet. Shinden Gompa,
Nagong river, 12–13000 ft., on granite cliffs inthe shade (Kingdon-Ward 10791, 26 Aug. 1933,
BM). SE. Tibet. Kongbo, Doré Nye Chu, 13000
ft., on dry ground (Ludlow, Sherriff and Elliot
15570, 6 Aug. 1947, A, BM). Kongbo Prov.,
Pangkar, Drukla Chu, 11500 ft., in burch forest in
deep shad (north face) (Ludlow, Sherriff et Taylor 6875, 21 Aug. 1938, BM). Northern slope of
Mt. Kenichunpo, north of Sikitung, upper Salwin
River, 12500 ft., on rocks, alpine region (J. F.
Rock 22144, May – June 1932, BM). Prov.
Tsarung, northern slope of Mt. Kenishunpo,
north of Sikitung, Upper Salwin River, 11–12000
ft., among rocks (J. F. Rock 22144, 22151,
May – June 1932, A). Burma-Tibet Frontier.
Sources of the Irrawaddy, Adung Valley, 28°20N
97°40E, 12000 ft. (F. Kingdon-Ward 9870, 28
July 1931, BM). E. Tibet and SW. China. (Forrest 30583, BM). Sichuan (Sikang or Szechuan).
Chi-na-tung, Tsa-wa-rung, 2800 m (C. W. Wang
65277, Aug. 1935, A). Daocheng, 4010 m (Wu etal. 722, 19 Aug. 1996, TI). Daocheng, around
Gongga Shan, 3920 m (Wu et al. 727, 21 Aug.
1996, TI). Huei-li Hsien (T. T. Yü 1592, 17 Sept.

Inquiries about taxonomy of the Sino-Himalayan Saxifraga

Fig. 8.

101

Saxifraga jacquemontiana Decne. (H. Ohba et al. 8331928, TI).

1932, A). Muli, Mt. Mitzuga, 4300 m (J. F. Rock
48131, Sept. 1929, A). Yien-pein Hsien, 3200 m(T. T. Yü 1714, 16 Oct. 1932, A). Yunnan.

Chungtien, Shianrentung, 3600 m, under thickets(T. T. Yü 13750, 18 Oct. 1937, A); sine loc. (T. T.
Yü 15517, A). Chungien, Tehgoh, 3200 m (T. T.

102

Shinobu Akiyama and Hideaki Ohba

Yü 13818, Sept. 1935, A). S. Chungtien, Wu-tso
on Yangtze bank, 3000 m (K. M. Feng 3327, 12
Nov. 1939, A). A tun-tze, Bai-mar-shan, 3500 m(C. W. Wang 69683, A). Ma-on-shan, W end of
Likiang valley (R. C. Ching 21388, 31 Aug.
1939, A). E. Likiang, Tai-ngo-koo (R. C. Ching
21642, 19 Sept. 1939, A). Likiang Snow Range(R. C. Ching 30588, 3 Sept. 1939, A). In prato ...
in mont. ivosis prope Lichiang, 3600 m (C.
Schneider 3023, 1 Sept. 1914, A). Lijiang, eastside of Mt. Yu-long-shui-shan, between Mahuang-ba and Wu-dou-di, 3345–3750 m (J. Murata et al. 603, 17 Aug. 1990, A).
Saxifraga jacquemontiana Decne.
Saxifraga jacquemontiana, described by Decaisne on the basis of a Jacquemont collection
from Kashmir in 1844, is a small cushion-forming species of the Himalaya and Tibet (Fig. 8).
The relationships of S. jacquemontiana are still
unclear, but S. paiquensis J. T. Pan and S.
jainzhuglaensis J. T. Pan are regarded as allied
taxa. Saxifraga paiquensis, described from SE.
Tibet, is closely similar to S. jacquemontiana in
the glandular ciliate margins of the basal leaves.
Saxifraga paiquensis was distinguished from S.

Fig. 9.

jacquemontiana in having leaves with a glandular (not aristate) apex and short glandular hairybasal leaves. These differences, however, probably fall within the range of variation of a single
species. Saxifraga jainzhuglaensis, also described from Tibet, apparently differs from S.
jacquemontiana in having widely spreading
sepals and clawless petals without callosities.
Pan (1991, 1992) classified Saxifraga jacquemontiana in the monotypic ser. Jacquemontianae
J. T. Pan. in sect. Ligulatae Haw. Pan attachedgreat significance to the cushion-like habit, theleafy shoots with a woody base, the floweringstems 3–4 mm long and embedded among the rosulate leaves, and the solitary flowers.
Saxifraga jacquemontiana has a rather wide
distribution, from the western Himalaya (Kashmir) to eastern Himalaya (Bhutan), and southeastern Tibet (Fig. 9).
Saxifraga jacquemontiana Decne. in Jacquemont, Voy. Inde 4 (Bot.): 68 (1844). Clarke in
Hook. f., Fl. Brit. Ind. 2: 395 (1878). Engl. et
Irmsch. in Engl., Pflanzenr. IV–117 (Heft 67):156 (1916). Marquand in J. Linn. Soc. Bot. 48:
178 (1929). H. Hara in H. Hara et Williams,

Geographic range of Saxifraga jacquemontiana Decne.

Inquiries about taxonomy of the Sino-Himalayan Saxifraga

Enum. Flow. Pl. Nepal 2: 153 (1978). Grierson in
Fl. Bhutan 1(3): 514 (1987). Pan in Fl. Reip. Pop.
Sin. 34(2): 208 (1992). Pan et al. in Fl. China 8:322 (2001).
[Figs. 8, 9]Specimens examined: Kashmir. W. Kashmir.
Kishenganga Valley and the road to Nanga Parbat via the Gangabal Lakes, Zojibal Pass, top,
13500 ft. (R. R. and I. D. Stewart 18196, 10 Aug.
1939, A). Kishenganga Valley and the road to
Nanga Parbat via the Gangabal Lakes, Burzil
Pass, top, 13800 ft. (R. R. and I. D. Stewart
19083, 27 Aug. 1939, A). Gilgit Road, Deosai
Plains, Baltistan and Ladak, Thalle La, Baltistan,
15–16000 ft. (R. R. Stewart 20725, 15 Aug.
1940, A). Gilgit Road, Deosai Plains, Baltistan
and Ladak, Burji La, Deosai side, 15000 ft. (R.
R. Stewart 20153, 31 July 1940, A). Baltistan,
Lal Pir, 135000 ft. (W. Koelz 9516, 10 Aug.
1936, A). Punjab. Simla Hill States, Rupin Pass,
Dhasla Dhar Range (31°22N 78 °09E), 14000
ft., on open rocky hillsides (Sherriff 7503, 2 Aug.
1939, A). Garhwal. Chor Noti, 16000 ft. (Strachey and Winterbottom 13, pro parte, A). Nepal.
Arun-Tamur Watershed, Lumba Sumba Himal,
N.N. of Walunchung, 16000 ft., on rocy grandgravel (Stainton 1027, 17 July 1956, A, BM).
Dolakha Distr., Yalung Kharka – Yalung
La – Pam Lhang, 27°50N 86°25E (H. Ohba et
al. 8320664, 8320683, 8331928, and 8351379, 8
Sept. 1983, TI). Solukhumbu Distr., around
Khare, 27°44 N 86°53E, alt. 4680 m (M. Wakabayashi et al. 9720218, 16 Aug. 1997, TI).
Solukhumbu Distr., Panch Pokhari – Seto
Pokhari, 27°51 86°56, alt. 5230 m (F. Miyamoto et al. 9584185, 16 Aug. 1995, TI). Sikkim.
Regio Alp., 16–18000 ped. (J. D. Hooker s. n.,
A). North Distr., Lakes on SE side of Sebu La,
sandy flats on lake shore, 27°55N 88°39E, alt.
5040 m (D. G. Long and H. J. Noltie 399, 22 July
1996, E, TI). West Distr., Goecha La, near summit, 27°36N 88°11E, alt. 4870 m (D. G. Longet al. 652, 22 July 1992, E, TI). Bhutan. Mar-103

lung, 14500 ft. (Ludlow, Sherriff and Hicks
19362, 6 July 1949, BM). Tibet. Lochen La,
16–17000 ft. (F. Kingdon-Ward 12234, 20 Aug.
1935, BM). SE. Tibet. Kongbo Prov., Budi
Tsepo La, 13000 ft., in mats on rocks (Ludlow,
Sherriff and Elliot 14425a, 21 Aug. 1947, BM).
Budi Tsepo La, 135000 ft., on rocks (Ludlow,
Sherriff and Elliot 15270a, 18 June 1947, BM).
Acknowledgements
We express our appreciation to the curators of
the herbaria cited above for permitting us to
study their valuable collections. We are indebted
to Dr. David E. Boufford, the Harvard University
Herbaria, for reading the manuscript. This study
was supported by a Grant-in-Aid for Scientific
Research (A) from the Japan Society for the Promotion of Science (no. 18255004 to HO).
References (other than thosein the systematic treatment)
Grierson, A. J. C. 1987. Saxifraga. In: Grierson, A. J. C.
and Long, D. G., Flora of Bhutan 1(3), pp. 492–515.
Royal Botanic Garden, Edinburgh.
Hara, H. 1979. Saxifraga. In: Hara, H. and Williams, L.
H. J. (eds.), An Enumeration of the Flowering Plants of
Nepal 2, pp. 151–156. Trustees of British Museum(Natural History), London.
Pan, J.-T. 1991. A study on the genus Saxifraga L. from
China. Acta Phytotaxonomica Sinica 29(1): 1–24.
Pan, J.-T. 1992. Saxifraga. In: Pan, J.-T. (ed.), Flora Republicae Popularis Sinicae 34(2), pp. 35–231. Science
Press, Beijing.
Pan, J.-T. Gornall, R. and Ohba, H. 2001. Saxifraga. In:
Wu, Z. and Raven, P. (eds.), Flora of China 8, pp.
280–344. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis.
Smith, H. 1924. Plantae Sinenses, A dre. H. Smith annis
1921–22 lectae. 1. The genus Saxifraga. Acta Horti
Gothoborgensis 1: 1–22, pls. I–X.
Smith, H. 1960. Saxifraga of the Himalaya II. Some new
species. Bulletin of British Museum (Natural History),
Botany 2(9): 229–260, pls. 13–21.