Taxonomic Studies on the Sino-Himalayan Saxifraga (1) : Section Ciliatae Series Lychnitideae, and Saxifraga excellens Harry Sm. and Resembling Species

2007·0.67 MB·English

by  Shinobu Akiyama

植物研究雑誌
J.Jpn.Bot.
82:284–295(2007)
Taxonomic Studies on the Sino-Himalayan Saxifraga (1):
Section Ciliatae Series Lychnitideae, and
Saxifraga excellens Harry Sm. and Resembling Species
Shinobu AKIYAMAa and Hideaki OHBAb
aDepartmentofBotany,NationalMuseumofNatureandScience,
4–1–1,Amakubo,Tsukuba,Ibaraki,305-0005JAPAN;
E-mail:[email protected]
bDepartmentofBotany,theUniversityMuseum,theUniversityofTokyo,
7–3–1,Hongo,Bunkyo-ku,Tokyo,113-0033JAPAN
(ReceivedonApril28,2007)
The genus Saxifraga is highly diverse in the Sino-Himalayan region, and our taxo-
nomic knowledge of the group is still insufficient. This is the first of a series of papers
concerning the taxonomy of Saxifraga, and deals with 1) the delimitation of section
CiliataeseriesLychnitideae(Engl.&Irmsch.)Gornalland2)S.excellensHarrySm.and
similar species. 1) Saxifraga series Lychnitideae consists of four species, S. lychnitis
Hook.f.&Thomson,S.nigroglanduliferaN.P.Balakr.,S.pseudohirculusEngl.,andS.
viscidulaHook.f.&Thomson.AnewcombinationS.lychnitisvar.oresbia(J.Anthony)
S. Akiyama & H. Ohba is proposed. 2) Three species with reddish or reddish purple
petals of Saxifraga section Ciliatae are compared. Saxifraga bergenioides C. Marquand
differs from S. excellens Harry Sm. and S. pardanthina Hand.-Mazz. in having pedicels
and sepals with brownish curled hairs against those with short, black tipped glandular
hairsofthetwootherspecies.SaxifragapardanthinadiffersfromS.excellensmainlyby
the spotted petals with cordate base and reflexed sepals. A recent collection of S.
excellens, endemic to Nepal and known only from a few specimens, allowed us to ob-
serve and illustrate it in detail.
Key words: China, Himalaya, Saxifraga, Sino-Himalaya, Tibet.
Saxifraga is one of the most diverse Akiyama et al. 1998, 1999, 2002, Ohba and
genera in the Sino-Himalayan region. Smith Akiyama 1999), and have also examined
(1958, 1960), Hara (1979), Pan (1992) and other collections in various herbaria, A, B,
many other workers have contributed to the BM, E, GH, K, L, LE, KUN, KYO, P, PE,
taxonomy of Saxifraga of the region, but the TI, TNS and others. Ohba cooperated in the
taxonomy of the genus is still insufficient, treatment of Saxifraga during preparation of
especially in the delimitation of species the Flora of China treatment of Saxifraga
described or recorded from both the with J.-T. Pan and R. Gornall (Gornall et al.
Himalaya and the Tibet-SW China regions. 2000, Pan et al. 2001). Akiyama is preparing
Since 1983 we have studied the genus and the treatment for the forthcoming Flora of
made field observations throughout the Nepal with R. Gornall and Ohba.
entire region, including the Himalaya, Tibet In this series of papers we intend to pro-
and SW China (Sichuan and Yunnan) (Ohba vide evidence on species delimitation, varia-
1984, 2002, Ohba and Wakabayashi 1987, tion, distribution, and specimens of
—284—
October2007 JournalofJapaneseBotanyVol.82No.5 285
Saxifraga in the region. having glabrous or brownish hairy (some-
times with glandular hairs) spreading sepals
(1) Section Ciliatae series Lychnitideae at anthesis, and petals with two calloses.
Saxifraga section Ciliatae Haw. (syn. Saxifragapseudohirculusmuchmoreclosely
Hirculus (Haw.) Tausch.) shows great diver- approachesS.viscidulaandS.nigroglanduli-
sity in the Sino-Himalayan region. The sub- fera than do the other species classified in
division of the section has been proposed by series Tanguticae by Pan (1991, 1992), espe-
various workers (Hooker and Thomson cially in having short glandular hairy
1858,EnglerandIrmscher1912,1913,1916, pedicels and persistent, glandular hairy,
Engler 1930, Gornall 1987, Pan 1991, 1992, petiolate basal leaves. Saxifraga yarlung-
Pan et al. 2001, etc.). zangboensis, described from Tibet and clas-
Engler and Irmscher (1912) established sified in series Yarlungzangboenses [as
section Hirculus subsection Lychnitideae as Yaluzangbuenses], is considered to be
a group with linear-oblong petals more than conspecific with S. viscidula Hook. f. &
3 times longer than wide. They classified Thomson from E Himalaya and Tibet. Pan
three species, S. lychnitis, S. viscidula and S. (1991, 1992) did not treat S. viscidula,
pseudohirculus into this subsection (Engler because he erroneously omitted this from the
andIrmscher1913,1916).Engler(1930)fol- Chinese flora.
lowed Engler and Irmscher (1913, 1916).
Gornall (1987) regarded Engler and Saxifraga sect. Ciliatae ser. Lychni-
Irmscher’s subsection Lychnitideae as a tideae (Engl. & Irmsch.) Gornall in Bot. J.
series in the section Ciliatae. Linn.Soc.95:276(1987)–J.T.PaninActa
Pan (1991, 1992) modified Gornall’s Phytotax. Sin. 29: 17(1991), as subgen.
infrageneric system and regarded Lychni- Hirculus sect. Ligulatae subsect. Nutantes
tideae as a series in subgenus Hirculus sec- ser. Lychnitideae – Akiyama & al. in Bull.
tion Ligulatae subsection Nutantes. Pan Natn. Sci. Mus. Tokyo, ser. B, 24: 67(1998).
(1991, 1992) classified three species, S. Saxifraga sect. Hirculus subsect.
lychnitis, S. oresbia and S. nigroglandulifera Lychnitideae Engl. & Irmsch. in Notes Roy.
in series Lychnitideae but excluded S. Bot. Gard. Edinburgh 5(24): 129 (1912); in
viscidula and S. pseudohirculus. Engler and Bot. Jahrb. 48: 588 (1913); in Engl.,
Irmscher (1912, 1913, 1916), Engler (1930) Pflanzen-reich, Heft 67: 132 (1916), as
and Gornall (1987) classified S. nutans grex–Engl.inEngl.&Prantl,Pflanzen-fam.
Hook. f. & Thomson, now known as S. 2 ed. 18a: 133 (1930).
nigroglandulifera, inthe monotypic subsec- Saxifraga sect. Hirculus subsect. Nutantes
tion or series Nutantes. Pan (1991, 1992) Engl. & Irmsch. in Notes Roy. Bot. Gard.
used the name ‘Nutantes Engl. & Irmsch.’ as Edinburgh 5(24): 129, 139 (1912); in Bot.
the name for the subsection in which the se- Jahrb. 48: 590 (1913); in Engl., Pflanzen-
riesLychnitideaeisclassifiedwithotherfour reich,Heft67:134(1916),asgrex–Engl.in
series including series Tanguticae J. T. Pan. Engl. & Prantl, Pflanzen-fam. 2 ed. 18a: 133
Akiyama et al. (1998) regarded series (1930) – J. T. Pan in Acta Phytotax. Sin.
Nutantes as a synonym of series Lychni- 29: 17(1991), as subgen. Hirculus sect.
tideae, since S. nigroglandulifera shares sev- Ligulatae subsect. Nutantes, excl. ser.
eral features with the species of that series. Tanguticae, Jainzhuglaenses and Jacqumon-
Pan(1991,1992)placedS.pseudohirculus tianae.
in series Tanguticae. Series Tanguticae Saxifraga sect. Ciliatae ser. Nutantes
which is distinct from series Lychnitideae in (Engl. & Irmsch.) Gornall in Bot. J. Linn.
286 植物研究雑誌 第82巻 第5号 平成19年10月
Soc. 95: 276 (1987). Nepal 2: 153 (1979) – Pan & al., Fl. China
Saxifraga subgen. Hirculus sect. Ligulatae 8: 306 (2002).
subsect. Nutantes Engl. & Irmsch. ser. 1a) Var. lychnitis
Yarlungzangboenses J. T. Pan in Acta Type: Sikkim, regio alp., Lama Kirngra,
Phytotax. Sin. 29: 18(1991), as 14000 ped. (J. D. Hooker s. n., July 24,
Yaluzangbuenses. 1849, Herb. Hook., K–holo).
Type species: Saxifraga lychnitis Hook. f. Distr. Himalaya (Kumaon to Bhutan),
& Thomson. Tibet and SW China (Yunnan).
This series consists of four species and is Specimens examined. Kumaon: Barji kang Nap,
characterized by the linear to narrowly ob- 14700ft.(Strachey&Winterbottom14,K).
long or oblanceolate to obovate petals, the Tiri-Garhwal: Rhudughera, 15–16000 ft. (Duthie
672, June 20, 1883, K); Dudu Gadh under Srikanta,
more or less densely brownish hairy calyx
15–16000ft.(Duthie672[sic],Aug.9,1883,K)
and pedicels, and the gradually descendent
Nepal: CNepal, Yalung Kharka–Yalung La, 4800
leaves. The species in the series m, on mossy stones and moss-covered bed by/in
Lychnitideae are distinguishable by a combi- streams (Ohba & al. 8351373, Sept. 8, 1983, TI);
nation of several characters. Yalung Kharka–Pam Lhang, 4800 m (Ohba & al.
8341241,8351373,8331942,8320665[2n(cid:1)16],Sept.
8,1983,TI).ENepal,SagarmathaZone,Solukhumbu
Key to the species of Saxifraga section
Distr., Rato Odara–Chhomalang Base camp, 4500 m,
Ciliatae series Lychnitideae
on exposed place or on mossy rocks on banks by
1. Basal leaves petiolate, blade elliptic or stream (Miyamoto & al. 9580272, 9592247, Aug. 11,
ovate or oblong 1995,TI);loc.cit.,4540m,barrenplace(Miyamoto&
2. Flowers secund; leaves subglabrous, al. 9584159, Aug. 11, 1995, TI); loc. cit., 5000 m, on
exposed place (Miyamoto & al. 9592281, Aug. 11,
blade of basal leaves 1.5–4 cm long ......
1995,TI);loc.cit.,5225m,rockyplace(Miyamoto&
............................ 2) S. nigroglandulifera
al. 9596316, Aug. 11, 1995, TI); Sagarmatha Zone,
2. Flowers in many sided; leaves glandular Solukhumbu Distr., around Dig Kharka, 4470 m
pubescent,bladeofbasalleaveslessthan (Wakabayashi & al. 9720206, Aug. 14, 1997, TI);
1.5 cm long ........... 3) S. pseudohirculus Sagarmatha Zone, Solukhumbu Distr., Beni Kharka–
Tschokarme,4550m(Ohba&Wakabayashi8531220,
1. Basal leaves sessile or nearly so, spatulate
8520372 [2n(cid:1)16], Sept. 1, 1985, TI); Sagarmatha
2. Petals 5–7-nerved, ca. 7 mm long; basal
Zone, Solukhumbu Distr., around Beni Kharka, 3970
leaves lanceolate-spatulate, cauline
m (Ohba & al. 8531213, Aug. 31, 1985, TI); Khung
leaves narrowly oblong or linear ........... Khola headwaters, ca. 5200 m (Grey-Wilson &
.......................................... 4) S. viscidula Phillips622,Aug.11,1973,K).
2. Petals 3(or 4)-nerved, 8.5–11 cm long; Sikkim: Naku Chu, Llonak, 17000 ft. (Smith &
Cave1943,Aug.3,1909,K);Llonak,15000ft.(Smith
basal leaves spatulate to ovate-spatulate
& Cave 2688, Aug. 4, 1909, K); Chapopla, 16000 ft.
toovate,caulineleavesoblong-ellipticto
(Ribu&Rhomoo5242,K);JankiLa(King’scoll.s.n.,
lanceolate-oblong or elliptic ................... Aug.,1877,K);LungnakLa,16000ft.,onbank(Cave
........................................... 1) S. lychnitis 58, Aug. 29, 1947, K); Donki La and southwards to
3. Petals narrowly oblong, 8.5–9 mm long Samdong (L. R. Wager 342, Aug. 17, 1933, K); N
Distr., Phaklung, Lasha Chhu Valley, 4620 m,
..................1a)S.lychnitisvar.lychnitis
hummockymarsh(Long&NoltieEENS332,July15,
3. Petals narrowly obovate, 10–11.7 mm
1956,E,TI).
long ...........1b) S. lychnitis var. oresbia
Bhutan: Yala, Pyla, Gilela areas, 15000 ft. (Gould
1334,Aug.1,1938,K).
1) Saxifraga lychnitis Hook. f. & Tibet:S.Tibet,MelongGompa,16–17000ft.(B.J.
Thomson in J. Linn. Soc. Bot. 2: 68(1857) Gould2222,July2,1939,K).
Himalaya-Tibet border: Mount Everest Exped.,
– C. B.Clarke in Hook. f., Fl. Brit. India 2:
15000ft.(A.F.R.Wollaston79,July,K).
391 (1878) – H. Hara, Enum. Flow. Pl.
October2007 JournalofJapaneseBotanyVol.82No.5 287
(1992) – Pan & al., Fl. China 8: 306 (2002).
Type: Sichuan (Szechuan), Litang–Yaling
divide, 14000 ft. (Kingdon-Ward 4939, Sept.
1921, E–holo).
Distr.: SW Sichuan.
Specimens examined. SW Sichuan: Win-chuan,
Tsao-puh,14800ft.(S.Y.Hu2637,Aug.7,1942,A);
Daocheng, Bowa Shan, 4460 m (Wu & al. 686, Aug.
16,1996,TI,KUN);Daocheng,Haizi,4550m(Wu&
al.717,Aug.17,1996,TI,KUN).
Variety oresbia is distinguished from var.
lychnitis by the narrowly obovate petals and
lanceolateoroblongtoellipticsessileleaves.
Hu 2637 differs from the typical form in
having pedicels without glandular hairs and
ovate or elliptic petals. Wu & al. 686 and
717 more closely approach var. lychnitis.
Specimens collected in Lumbasumba Himal,
E Nepal; the interior of Topke Gola and
Thudam, around Lamni Nama, 4300 m, on
rocksbystreams(Ohashi&al.773810,Aug.
15, 1977, TI) and Lama Chunghu-Samdan,
4380 m, open mossy places by streams
Fig.1. Saxifraga lychnitis var. oresbia (Kingdon-
Ward 4939, E, holotype of Saxifraga oresbia J. (Kanai & al. 720719, P. R. Shakya 1706,
Anthony).Scaleindicates5cm. June 24, 1972, TI), are also transitional
between vars. lychnitis and oresbia.
Bhutan-Tibet border: East of Phari, ca. 16000 ft. 2) Saxifraga nigroglandulifera N. P.
(Gould1405,Aug.1,1938,K). Balakr. in J. Bombay Nat. Hist. Soc. 67: 59
Yunnan: N.W. Yunnan, Haba snow Range, open
(1970) – Pan & al., Fl. China 8: 306 (2001).
hill(K.M.Feng2174,Aug.24,1939,A).
[Fig. 2A]
The range of Saxifraga lychnitis is the
Saxifraga nutans Hook. f. & Thomson in
most extensive of the series Lychnitideae
J. Linn. Soc. Bot. 2: 69(1857), not D. Don
and covers nearly the whole range of the
(1822) et Adams (1834) – H. Hara, Enum.
series. Saxifraga lychnitis is rather common
Flow. Pl. Nepal 2: 154 (1979). Type:
in E Himalaya (especially Nepal and
Sikkim, regio temp., 10–13000 ped. (J. D.
Sikkim), and often gregarious in alpine
Hooker s. n., K–holo; A–iso).
meadows, but is rather rare in Tibet and SW
Distr. E Himalaya (Nepal and Sikkim),
China (Yunnan). No specimens from
Tibet and SW China (Yunnan and Sichuan).
Sichuan have been examined. The flowers
Specimensexamined.Nepal:Tegar,NofMustang,
arenoddingthroughoutthefloweringseason.
1500ft.(Stainton,Sykes&Williams2221,A);Tamur
Valley, Yangma Khola, NE of Walunchung Gola,
1b) Var. oresbia (J. Anthony) S. Akiyama
14000 ft. (Stainton 1081, July 23, 1956, A, BM);
& H.Ohba, stat. nov. [Fig. 1] Barbung Khola, E of Charhkka, ca. 4500 m (Grey-
Saxifraga oresbia J. Anthony in Notes Wilson & Phillips 459, July 31, 1973, K); Khung
Roy. Bot. Gard. Edinburgh 18: 28(1933) – Kholaheadwater,ca.5000m(Grey-Wilson&Phillips
623,Aug.16,1973,K).
Pan, Fl. Reipubl. Pop. Sin 34(2): 199
288 植物研究雑誌 第82巻 第5号 平成19年10月
Fig.2. A. Saxifraga nigroglandulifera (S Tibet: Nyalam–Zhangmu, 3690 m, Aug. 22, 2001). B.
S. excellens (E Nepal: Sagarmatha Zone. Solukhumbu distr., Thasing Dingma–Chhatarwa,
3800m,Aug.28,1995).PhotobyF.Miyamoto.C.S.bergenioides(STibet:Maizhokunggar
Xian,Mt.MilhaShan,4920m,Aug.6,2001).
October2007 JournalofJapaneseBotanyVol.82No.5 289
Sikkim:Giagong,16000ft.(Prains.n.,Sept.1903, TI, KUN); Xiaojin, Ganhaizi–southwestern slope of
K);NSikkim(L.R.Wagers.n.,July23–27,1923,K); Mt.Siguniang–Ganhaizi,4100m(Ikeda&al.100417,
Thanga, 14000 ft. (Ribu 2920, in 1909, K); Tangu, Sept.1,1998,TI,KUN).
14000ft.(Smith&Cave2279,Aug.8,1900,K). The flowers of Saxifraga nigroglanduli-
Tibet: STibet, Le La, Chayul-Charme Rd., 15000 fera eventually nod and the petals are
ft. (Ludlow & Sherriff 2450, July 27, 1936, BM);
oblanceolate or narrowly oblong. We have
Salween-Tsangpo divide, NE of Shinden Gompa
seen no specimens transitional with other
(Kingdon-Ward 10727a, in 1933, BM); S of Gyalam,
14000ft.(Kingdon-Ward12277,Aug.31,1935,BM); species.
SE, Nye La, Ozogang District Kham, 14900 ft. (J.
Hambury-Tracys.n.,Sept.10,1936,BM);Vicinityof 3) Saxifraga pseudohirculus Engl. in
Lhasa, 13000 ft. (H. E. Richardson 262, July 1939,
Bot. Jahrb. Syst. 48: 590 (1912) – Pan, Fl.
BM); E of Lhasa (H. J. Walton s. n. in 1904, K); Kyi
Reipubl. Pop. Sin 34(2): 204 (1992) – Pan &
ChuValley,15milesEofLhasa(H.J.Waltons.n.in
al., Fl. China 8: 305 (2001).
Sept.,1904,K);Chala,nearLhasa,15500ft.(Ludlow
&Sherriff8995,Aug.5,1942,BM);Nangtse,20miles Type: S Tibet, Chumbi (Dungboo [Dr.
W ofLhasa, 14500 ft. (Ludlow & Sherriff 982b, July King’s Collector], CAL, not seen).
28, 1943, BM); Hills, north of Lhasa, 15000 ft. Saxifraga hirculoides Engl. ex Maxim. in
(Ludlow & Sherriff 9082, Sept. 19, 1942, BM); SE
Bull. Acad. Sci. St.-Pétersb., Sér. 3, 29: 112
Tibet, Gorpo La, N of Dongkar, 15500 ft. (Ludlow &
(1883), not Decne. (1844). Type: China
Sherriff822,Aug.15,1938,BM);Reting,60milesN
of Lhasa, 14500 ft. (Ludlow & Sherriff 11134, Aug. occidentalis, prov. Kansu in regione alpina
19,1944,BM);nearKampa,14500ft.(C.S.Catting& declivitatis borealis jugi ad austrum fluminis
A. S. Vernay 106B, Sept. 11, 1935, K); above Sigma Tetung, 10–12000’s [3–4000 m] (N. M.
Rhangchung, 11500 ft. (F. Spencer Chapman 181,
Przewalski 648, July 20/1 Aug., 1880, LE–
Aug. 21, 1936, K); Zogang–Dongdala Shan, 4650 m
holo).
(Akiyama & al. 105852, 18 July 2000, TI, KUN);
S. pseudohirculus Engl. var. shensiensis
Nyalam–Zhangmu, 3690 m (Akiyama & al. 106553,
22Aug.2001,TI,KUN). Engl. & Irmsch. in Bot. Jahrb. Syst. 48: 590
Western China: Yung Ning (McLaren’s collector (1913). Types: N Shaanxi (Nord-Shensi),
155inSept.1933,BM,K);loc.sinedescr.,13–14000 T’ai-par-shan (G. Giraldi 5441–5447, B–
ft.(Wilson3595,July19,1903,BM,K).
syn, not seen); Distrikt Pao-Ki-seen, auf
Yunnan: Eastern flank of Likiang Range (Forrest
dem Miao-Wang-san (G. Giraldi 5448, B–
6599, 1910, E, BM, MO); loc. cit., 11–12000 ft.
(Forrest 2621, July 1906, BM, K); Mountains NE of syn, not seen).
Atuntze, 12–13000 ft. (Forrest 20150, Sept. 1921, S. pseudohirculus Engl. var. tenuiflora
BM); in reg. Chung-tien, 3400 m (C. Schneider 3663, Harry Sm. in Acta Hort. Gothob. 1: 12
July 1914, K); Chung-tien. Mountains west of Hsiao
(1924). Types: Sichuan (N Szechuan),
Chung-tien, 10500 ft. (J. F. Rock 24634, BM, MO);
Dongrergo, 4300 m (H. Smith 3515, Aug. 8,
loc. cit. (J. F. Rock 25264, K); Zhongdian, Haba
1922, UPS–syn; K–isosyn), loc. cit., 4100 m
Xueshan,HabaHai,4020m(Wu&al.103476,9Aug.
1999, TI, KUN); Mount Habashan, north of Ndaku, (H. Smith 3520, Aug. 9, 1922, UPS–syn;
north of Likiang Snow Range, Yangtze Drainage BM–isosyn), loc. cit. (H. Smith 3911, UPS–
Basin, 14000 ft. (J. F. Rock 9639, July 1933, BM); syn, not seen).
Mont. niveor. prope Lichiang, 3800 m (C. Schneider
S. selgenensis K. S. Hao in Bot. Jahrb.
1812,June16,1914,K).
Syst. 68: 604 (1938). Type: Xinjiang,
Sichuan (including Sikang): Kangting (Tachienlu)
distr., Yülingkong, mont. orient., Gomba La, ca. 3900 Kokonor, auf dem Selgen, um 4800 m (K. S.
m, in prato alpino (H. Smith 10729, July 22, 1934, Hao 938, Aug. 21, 1930, PE).
BM); Taofu (Dawo) distr., Montes orient. Lhamo S. balongshanensis T. C. Ku in Bull. Bot.
Mondek La, ca. 4600 m (H. Smith 12329, Sept. 22,
Res., Harbin 9(4): 7 (1989). Type: Sichuan,
1934,BM);Daocheng,BowaShan,4460m(Wu&al.
Wenchuan, Mt. Balongshan, 4–4350 m (K.
688, Aug. 16, 1996, TI, KUN); Daocheng, around
Y. Lang, L. Q. Li & Y. Fei 1125, Aug. 18,
Gongga Shan, 4430 m (Wu & al. 847, Aug. 5, 1997,
290 植物研究雑誌 第82巻 第5号 平成19年10月
1982, PE–holo). 11256, Aug. 20, 1934, A, BM); loc. cit., 4200 m, in
S. longipetala T. C. Ku in Bull. Bot. Res., prato alpino (H. Smith 11209, Aug. 3, 1934, A, BM);
Harbin 9(4): 10 (1989). Type: Sichuan, Kangtingdistr.,ChetoLa,42–4400m,inpratoalpino
(H. Smith 11040, Aug. 3, 1934, A, BM); Kangting
Kangding, Mt. Zheduoshan, 4000 m (K. C.
distr., Tapaoshan, west range, ca. 4300 m, in prato
Kuan & al. 1066, July 28, 1963, PE–holo).
alpino (H. Smith 11303, Aug. 20, 1934, BM);
S. spathulifolia T. C. Ku in Bull. Bot. Kangtingdistr.,Tapaoshan,westrange,ca.4700m,in
Res., Harbin 9(4): 11 (1989). Type: Sichuan, rupibus calcareis (H. Smith 11294, Aug. 20, 1934,
Xiaojin, Shepiliangzi, 4400 m (X. S. Zhang BM); Taofu distr., mont. orient., Lhamo Mondeh La,
ca. 4000 m, in prato herboso-praticoso (H. Smith
&Y.X.Ren6555,Aug.13,1958,PE–holo).
12274,Sept.21,1934,BM);Tachienlu,9-13500ft.(A.
Distr.:Tibet,Qinghai,Gansu,Shaanxiand
E.Pratt616&613,BM);Reg.bor.-occid.,interMerge
Sichuan.
etZanskar,ca.4300m,inrupibuscalcareis(H.Smith
Specimens examined. Tibet: Cha La, near Lhasa, 4335,Sept.4,1922,BM);Reg.bor.-occid.,interKerge
15000 ft., among rocks on grassy hillside (Ludlow & et Lanker, ca. 4300 m, in rupibus calcareis (H. Smith
Sherriff 8988, Aug. 4, 1942, BM); Hills S. of Lhasa, 4335, Sept. 4, 1922, K); Sungpan Hsien, at grassy
15500 ft. (Ludlow & Sherriff 2885, Aug. 19, 1943, slope (W. P. Fang 4023, Aug. 8, 1928, K). W
BM); Hills north of Lhasa, 15500 ft., grassy uplands Szechuan and Tibet Frontier, chiefly near Tachienlu,
(Ludlow & Sherriff 9025, Aug. 21, 1942, BM); 9000–13500ft.(A.E.Pratl613,K).
Kongbo Province, Phu Chu, near Paka, 14000 ft., Saxifraga pseudohirculus with its numer-
29º15′, 94º25′, onopen grassy hillside (Ludlow, ous, erect flowers is unique in the series
Sherriff & Taylor 5962, July 27, 1938, BM); Salween
Lychnitideae. The ensiform or oblanceolate
–Tsangpo divide, N.E. of Shinden Gompa, 15–16000
to narrowly oblong petals, and the pedicels
ft., dry ground (Kingdon-Ward 10729, Aug. 11, 1933,
BM);ETibet,GrasslandbetweenLabrangandYellow and calyx with dense dark brown glandular
River, near camp, Wanrgen Valley, 12200 ft. (J. F. hairs support its placement in the series
Rock 14527, July 29, 1926, A, K); Above Singma Lychnitideae. Saxifraga cacuminum Harry
Khangchung, 11500 ft. (Spencer Chapman 580, Aug.
Sm. approaches S. pseudohirculus, but dif-
31,1936,A).
fers in having aristate basal leaves with the
SE Tibet-Yunnan border: Tsarung border, 13000
margins and lower surface hispidulous, and
ft., along water courses (J. F. Rock 23462, Oct.–Nov.
1932,A). solitary flowers. The characters of the basal
Qinghai: Maqin (Maqên) Xien, Xihalong Guo, leaves of S. cacuminum indicate its affinity
between Jungong (Gyumgo) and Maqin on S side of to S. aristulata Hook. f. & Thomson and
the Huang He, 35–3600 m, on wet slopes (Ho,
closely related species. To decide its taxo-
Bartholomew&Gilbert288,A,BM);DawuXiang,SE
nomic status, further information is neces-
of Maqin, 3920 m, alpine meadow with Hedysarum
and N facing slope with very dense turf and frequent sary.
dwarf shrubs, mostly Potentilla (Ho, Bartholomew &
Gilbert 518, A, BM); Heit Shan, Dawu xiang, at pass 4) Saxifraga viscidula Hook. f. &
between Jungong (Gyumgo) and Maqin, 4220 m,
ThomsoninJ.Linn.Soc.Bot.2:68(1857)–
alpine meadow, disturbed by grazing (Ho,
C. B. Clarke in Hook. f., Fl. Birt. India 2:
Bartholomew&Gilbert654,BM).
391 (1878).
Gansu(Kansu):SWKansu,UpperTebbCountry(J.
F. Rock 13089, A, K); Kansu occident, Lou Kio wa Type: Sikkim, regio alp., 13–15,000 ped.
ssen (Abbé E. Licent 4782, Aug. 28, 1918, K); La (J. D. Hooker s. n., K–holo; A–iso).
Chang K’ou, near Sining, 30–3300 m, shaded slopes Saxifraga yarlungzangboensis J. T. Pan in
(R. C. Ching 622, July 24–25, 1923, A); China
ActaPhytotax.Sin.16(2):26(1978)–Pan&
occidentalis, Regio Tangut (Prov. Kansu) (N. M.
al., Fl. China 8: 305 (2002), syn. nov. Type:
Przewalskiin1880,K).
Shaanxi (Shensi): Taipeishan (W. Purdom in 1910, Xizang, (Xizang Medicinal Expd. 4440,
K). HNWP–holo, not seen).
Sichuan [Sikang]: Kangting distr., Tapaoshan, in Distr.: E Himalaya (Sikkim) and SE and E
jugooccid.,ca,4800m,inrupibuscalcareis(H.Smith
October2007 JournalofJapaneseBotanyVol.82No.5 291
Tibet. Hirculoideae of section Ciliatae along with
Specimensexamined.EasternHimalaya:Pa-Sum- S. hirculoides Decne., S. subamplexicaulis
KyeLa,14–15,000ft.(Kingdon-Ward6127,Aug.25, Engl. & Irmsch., S. linearifolia Engl. &
1924,BM). Irmsch., S. erectisepala J. T. Pan, and S.
Tibet: Tse La, 16000 ft., on alpine turf slopes
lepidostolonosa Harry Sm. because of its
(Kingdon-Ward 12261, Aug. 22, 1935, BM); S. E.
curled brownish hairs, sepals with three
Tibet, Kongbo Province, Kyabden, Nyang Chu, 13–
14000 ft., on grassy cliff ledges and open steep grass inconfluent nerves and petals without cal-
slopes (Ludlow, Sherriff & Taylor 6184, Aug. 23, luses. However, these two are different from
1938,BM);S.E.Tibet,TrakaLa,Mago,14500ft.,in all others in the series in having the reddish
stonymoraineslopes(Ludlow&Sherriff816,Aug.5,
or reddish purple petals.
1934, BM); S. E. Tibet, Cha La, Dongkar, 14000 ft.,
Saxifraga pardanthina from SW China
open stony hillside (Ludlow & Sherriff 840, Aug. 18,
(YunnanandSichuan)resemblesS.excellens
1934,BM).
In Saxifraga yarlungzangboensis the glan- in having dense, short, black tipped glandu-
dularhairsonthepetals,theonlyfeaturedis- lar hairs on pedicels and the outer surface of
tinguishing it from S. viscidula, vary from sepals and hypanthium, but differs greatly
nearly absent to moderate. in the reflexed sepals, cordate based ovate
petals, and in the middle to upper cauline
(2) Saxifraga excellens Harry Sm. leaves not amplexicaul. We agree with Pan’s
and resembling species treatment on S. atropurpurea J. Anthony as
Of the Sino-Himalayan Saxifraga, only a synonym of S. pardanthina. Saxifraga
three species with reddish or reddish purple pardanthina has rather dense blackish purple
petals are known in section Ciliatae. They spots throughout petals, but S. excellens and
are S. excellens Harry Sm., S. bergenioides S. bergenioides have petals without any
C. Marquand and S. pardanthina Hand.- spots and calluses.
Mazz. The reddening of petals is considered to
Saxifraga excellens differs from S. occurindependentlyinseveralgroupsinsec-
bergenioides in having dense, short, black tion Ciliatae, and to be inappropriate for
tipped glandular hairs on the pedicels. establishing any infrageneric taxa based on
Saxifraga bergenioides has only brownish this. We refrain from further systematic
curledhairsonthesamestructures.Thepres- arrangement of these taxa because of a lack
ence of glandular hairs and curled hairs is of information.
stable and constant within a species. Pan
et al. (2001) used these differences in hairs 1) Saxifraga bergenioides C. Marquand
as an important character to distinguish the in J. Linn. Soc. Bot. 48: 176 (1929) – Pan &
main groups classified under Key 3 and Key al., Fl. China 8: 314 (2001). [Figs. 2C, 3]
6 insection Ciliatae. Moreover, these two Type: S. E. Tibet, Sang La, 4500–4800 m
species differ in the posture of the flowers, [15–16,000 ft.], amongst boulders and on
which is erect or nodding. Smith (1960) es- cliffs on the exposed side of the mountains
tablished a monotypic grex Cinctae in sec- (Kingdon Ward 5912, July 8, 1924, K–holo,
tion Ciliatae based on Saxifraga excellens iso).
characterized by the basally connate fila- Distr.Tibet,endemictoSETibet(Kongbo
ments. But basal connate filaments are also and Takpo Provinces).
found in several dissimilar species. Specimens examined. SE Tibet: Kongbo Province,
Pan (1992) placed Saxifraga bergenioides Valley above Tripe, Tsangpo Valley, 13500 ft., on
grassy slopes and screes (Ludlow, Sherriff & Taylor
and S. pardanthina Hand.-Mazz. in series
5420,July26,1938,K);PasumLa,DruklaChu,16000
292 植物研究雑誌 第82巻 第5号 平成19年10月
Fig.3. Saxifragabergenioides(Akiyama&al.104533,Aug.17,2000,TI).
a.Petal.b.Sepal.c.Flower.Barindicates3mmforpetalandsepal,and
6mmforflower.
ft., in cleft of rocks (Ludlow, Sherriff & Taylor 6896, al.104533,Aug.17,2000,TI,KUN).
Aug. 23, 1938, K); Nambu La, 15000 ft., on dry The distribution is extremely narrow and
ground (Ludlow, Sherriff & Elliot 15417, July 13, limited to Kongbo and Takpo provinces in
1947,K);Buchu,LechenLa,13000ft.,ondryground
SE Tibet.
(Ludlow, Sherriff & Elliot 15553, Aug. 4, 1947, K);
Nyima La, Rong Chu, 14500 ft., on grassy rock shelf
(Ludlow, Sherriff & Taylor 5122, July 4, 1938, K); 2) Saxifraga excellens Harry Sm. in Bull.
Tsari, Tsari chu, Bimbi La, 13500 ft., on mossy rock Brit. Mus. (Nat. Hist.), Bot. 2(9): 230 (1960)
shelves (Ludlow, Sherriff & Taylor 6321, Oct. 14, – H.Hara, Enum. Flow. Pl. Nepal 2: 152
1938, K); Tsari, Sexx La, 14500 ft., rocky situations
(1979). [Figs. 2B, 4]
(Ludlow&Sherriff1952,July27,1936,K);MiraLa,
Type: Nepal, Annapurna Himal, Mardi
Puchu, 14–15000 ft., on steep grassy open hillside
Khola, 12500 ft. (Stainton, Sykes &
(Ludlow, Sherriff & Taylor 6153, Aug. 15, 1938, K);
NamLa,14500ft.,amongstrocks(Ludlow,Sherriff& Williams 8460, 18 Sept. 1954, BM–holo;
Elliot 14479, Sept. 1, 1947, K); Kulu Phuchu, near iso).
Paka, 14–15000 ft., common on open (Ludlow, Distr. Nepal, endemic to C and E Nepal.
Sherriff & Taylor 5950, July 27, 1935, K); Mira La,
Specimens examined. E Nepal: Sagarmatha Zone.
Nyang Chu, 13500–14000 ft., among rhododendrons
Solukhumbudistr.,ThasingDingma–Chhatarwa,3800
on steep hillside (Ludlow, Sherriff & Taylor 6057,
m, on mossy rocks in Abies forest (Miyamoto & al.
Aug. 13, 1938, K); Le La, Chagul-Charme, 15000 ft.,
9580471, Aug. 28, 1995, TI); loc. cit., 3750 m
on open rocky hillside (Ludlow & Sherriff 2285, July
(Miyamoto&al.9596499,Aug.28,1995,TI).
4,1936,K).TakpoProvince,TseLa,Langong,14000
These collections agree well with the type
ft., on moss covered rocks beside stream (Ludlow,
Sherriff & Taylor 5617, June 21, 1938, K); Tse La, collected in C Nepal, except for the shape of
16000 ft., in shade, under bushes, alpine region the apex of the sepals and petals.
(Kingdon-Ward 12248, Aug. 22, 1935, K);
Maizhokunggar Xian, Mt. Milha Shan, 4920 m
3) Saxifraga pardanthina Hand.-Mazz.
(Akiyama & al. 106407, Aug. 6, 2001, TI, KUN,
in Symb. Sin. 7: 418 (1931) – Pan, Fl.
TNS); Nyingchi, Sezhaila Shan, 4500 m (Akiyama &
October2007 JournalofJapaneseBotanyVol.82No.5 293
Fig.4. Saxifraga excellens (Miyamoto & al. 9580471, Aug. 28, 1995, TI). Bar indicates 3
cmforplant,and3mmforflower,sepalandpetal.