全 文 ：青藏高原一新特有属———异型株属及其传粉模式*
何廷农 1, 刘尚武1, 刘建全1,2**
(1 中国科学院西北高原生物研究所高原生物进化与适应重点实验室, 青海 西宁摇 810001;
2 兰州大学生命科学院草地农业生态系统国家重点实验室, 甘肃 兰州摇 730000)
摘要: 胚胎学和分子证据表明广义獐牙菜属的异形花组应该独立为一个新属。 在重新检查该属唯一种在各
居群的形态变化时, 发现该物种具有异形植株和异形花; 前一性状在以前的研究中被忽略了, 同时还发现
该物种主要是采用闭花受精结实, 这一繁殖特点可能是对青藏高原极端环境生存适应的结果。 这些特征明
显不同于獐牙菜属的其余种和相关属。 综合其它证据讨论了该种的系统位置, 认为其与花锚属有最近的亲
缘关系, 这一物种独特的形态性状和系统位置支持其独立为属, 并正式描述和发表这一新属———异型株
属。 但是必需指出的是, 广义獐牙菜属排除这一物种后, 分子证据表明其还是一个复系属。 分类上处理这
种由于快速物种多样化和性状趋同进化导致的复系属仍面临巨大的挑战, 还有待今后更详细的研究。
关键词: 异型株属; 异型植株; 异型花; 传粉式样
中图分类号: Q 949摇 摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2013)03-393-08
A New Qinghai鄄Tibet Plateau Endemic Genus
Sinoswertia and Its Pollination Mode*
HE Ting鄄Nong
1
, LIU Shang鄄Wu1, LIU Jian鄄Quan1,2**
(1 Key Laboratory of Evolution and Adaptation of Plateau Biology, Northwest Plateau Institute of Biology, Chinese Academy
of Sciences, Xining 810001, China; 2 State Key Laboratory of Grassland Agro鄄Ecosystem, College of
Life Science, Lanzhou University, Lanzhou 730000, China)
Abstract: Available data from both embryological and molecular studies of the monotypic section Heteranthos within
the genus Swertia s. l. suggested that this taxon should be erected as a separate genus. We examined morphological
variations of the only species across multiple populations and found that it has two types of individuals and one type
contains both large and small flowers while the other only small flowers. However, all small and most large flowers
set seeds through a cleistogamous pollination mode, which may have evolved in response to the reproductive adapta鄄
tion to the arid habitats of the Qinghai鄄Tibet Plateau. However, this pollination mode is different from the outcrossing
breeding system observed for the other Swertia species. Dimorphic plants and flowers distinguish this species from
the other Swertia species and related genera. We further discussed the systematic position of this species based on
the comprehensive data and the current data support its close relationship with Halenia. The morphological distinct鄄
ness and systematic position of this species from all comparisons justified its generic status. We here place this spe鄄
cies in a new genus Sinoswertia, and describe and illustrate it. However, after excluding this species, Swertia s. l.
is still polyphyletic based on the molecular evidence. The taxonomic treatment of this polyphyletic genus possibly due
to the recent radiative diversification and convergent evolution poses an extreme challenge at the present and pends
further detailed studies.
植 物 分 类 与 资 源 学 报摇 2013, 35 (3): 393 ~ 400
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 7677 / ynzwyj201313042
*
**
Foundation item: Key Project of the National Basic Research Program of China (2007FY110100)
Author for correspondence; E鄄mail: liujq@ lzu. edu. cn
Received date: 2013-03-19, Accepted date: 2013-04-09
作者简介: 何廷农 (1938-2011) 女, 研究员, 已故中国龙胆科植物分类学专家。
Kew words: Sinoswertia; Dimorphic individuals; Dimorphic flowers; Pollination mode
摇 With more than 160 species accepted, Swertia
is one of the large genera in the tribe Gentianinae of
the family Gentianaceae (Struwe and Albert, 2002).
It is widely distributed in Asia, Africa, North Amer鄄
ica and Europe. This genus was defined based on
the rotate corolla and diverse nectaries not surroun鄄
ded by fringes or appendages (Ho, 1988; Ho and
Pringle, 1995). However, all phylogenetic analyses
based on the molecular data suggested that Swertia
in its present circumscription is polyphyletic and that
the rotate corolla as well as fringed nectaries may
have derived from homoplastic evolutions due to the
radiative diversifications of the common ancestor to鄄
gether with the related genera or taxa ( Yuan and
K俟pfer, 1995; Chassot et al., 2001; Liu et al.,
2001; Hagen and Kadereit, 2002). It is necessary
to use distinct morphological traits to define these i鄄
dentified monophyletic lineages inferred from molec鄄
ular data.
Within this in mind, we aimed to examine mor鄄
phological traits and taxonomic rank of a special tax鄄
on, sect. Heteranthos within the genus Swertia,
which is endemic to the Qinghai鄄Tibet Plateau.
The only species of this monotypic section, S. tet鄄
raptera, was found to cluster with Halenia with
spurred flowers (Yuan and K俟pfer, 1995; Chassot
et al., 2001; Liu et al., 2001; Hagen and Kadere鄄
it, 2002). The embryological comparisons also sug鄄
gested that unique characters united these two taxa
together and the exclusion of this taxon from Swertia
s. l. was further inferred (Xue et al., 1999a, b).
In the present study, we reported dimorphic plants
and flowers found across all populations of this spe鄄
cies and we also found that most flowers of this spe鄄
cies set seeds through a cleistogamous pollination
mode. All these traits clearly distinguish this species
from the other Swertia species and related genera.
Therefore, this species should be elevated to a sepa鄄
rate genus and the new genus is described here for
the first time.
1摇 Materials and methods
1. 1摇 Morphology
The previously collected specimens in HNWP
were initially examined and two types of individuals
were found. We explored this morphological varia鄄
tion in the field populations between 2002 and
2010. In total, we visited 14 populations at altitudes
between 2 420 m and 4 310 m. We randomly collect鄄
ed 100 individuals and examined the distribution of
two types of individuals in each population.
1. 2摇 Pollination mode
In 2008, we subjected three types of flowers
found in S. tetraptera, i. e., small flowers of the
homogeneous individuals, large and small flowers of
the heterogeneous individuals for the following five
treatments: (1) control; (2) bagged isolation; (3)
isolation without emasculation; (4) emasculation for
natural pollination; and (5) emasculation and hand鄄
pollination with pollen grains from the other individ鄄
uals. We marked flowers for treatments (1), (2)
and (3) at the bud phase while emasculations in
treatment for (4) and (5) were completed before
stigma lobe opened. In order to provide enough
space for flower opening and closing, we isolated all
flowers with paper bags with at least two sizes of
flowers. All these experiments were finished at one
population in Datong, Qinghai (2 700 m) and we al鄄
so monitored how many flowers had opened in this
population through the total growing season. We
used one鄄way ANOVA and Post hoc鄄LSD to examine
seed sets associated with different treatments.
1. 3 摇 Systematic position inferred from molecu鄄
lar evidence
We examined the sequence variations between
different populations and two different individuals
from three populations. The extracting of the total
DNA and amplifying and sequencing of three DNA
fragments ( trnL鄄F, trnS鄄Ycf9, and ITS) followed
the previous standard procedures and primers (Yuan
and K俟pfer, 1995; Chassot et al., 2001; Liu et al.,
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2001; Hagen and Kadereit, 2002). We also down鄄
loaded the corresponding sequences for the other
genera or species (Table 1). Two Gentiana species
were chosen as outgroups. Phylogenetic analyses of
cpDNA and ITS sequences were performed by Mr鄄
Bayes 3. 0 (Ronquist and Huelsenbeck, 2003). All
gaps ( indels) were excluded. For Bayesian analy鄄
ses, four chains were used per run ( three heated
and one cold), and each analysis was repeated three
times, twice for two million generations, with the fi鄄
nal analyses running for 10 million generations. We
used Bayesian posterior probabilities to assess
branch support and the supports were calculated
from 1000 replicates using a heuristic search with
simple addition with TBR and MULPARS options on
in PAUP*(Swofford, 2002).
2摇 Results
2. 1摇 Morphology
Swertia tetraptera was described by Maximowicz
in 1881 and he described only one type of individu鄄
als. We firstly examined the specimens kept in the
herbaria and found that most sheets comprised only
one heterogeneous type of individuals with large and
small flowers. However, in a few sheets, two types
of individuals, one heterogeneous type and the other
homogeneous individuals with only small flowers. In
all examined populations in field, plants conspicu鄄
ously have formed two types: heterogeneous and
homogeneous plants. Heterogeneous plants are tall
and stout, with a distinct erect main stem and few to
numerous, procumbent, weak branches at stem
base. Inflorescence is a panicle of cymes, few鄄 to
many鄄flowered, spreading. However, flowers are di鄄
morphic. On main stem or rarely also on terminal
flowers of basal weak branches, flowers are large鄄
sized; nectaries are linear鄄oblong, canaliculate,
with a very narrow scale and few irregular divisions;
capsules are large (10-14 mm), narrowly ellipsoid
with seeds 13-18 (-27) in number. On weak bran鄄
ches, flowers are small鄄sized, 1 / 4-1 / 3 as large as
those of main stem; nectaries have broadly oblong,
entire, elevated scales and indistinct flat gland pat鄄
ches; capsules are small鄄sized (4-5 mm), broadly
ovoid to subglobose with seeds (2-) 4-6 (-7) in
number. Homogeneous plants are dwarf; stems are
flaccid, slender, few鄄branched without basal weak
branches at stem base. Cymes are lax, few鄄flow鄄
ered. All flowers are uniform and small鄄sized, their
calyx, corolla, nectaries, stamens, pistil, capsules
and seeds ( in shape, size, color, structure and
number) are the same as those of the small鄄sized
flowers on heterogeneous plants. The ratio of the two
plant types changes with the altitudes and a highly
significant negative correlation was found between
frequencies of heterogeneous versus homogeneous
Table 1摇 Accession numbers of the sequences used for constructing phylogenetic relationships of Sinoswertia tetraptera
Species trnL鄄trnF trnS鄄Ycf9 ITS
Halenia brevicornis GI: 15149891 GI: 15209270 GI: 15149919 GI: 21261681 GI: 21261650
Halenia weddelliana GI: 15149893 GI: 15209272 GI: 15149921 GI: 15149957 GI: 15149986
Halenia elliptica GI: 15149892 GI: 15209271 GI: 15149920 GI: 15384520
Sinoswertia tetraptera GI: 15150018 GI: 15209307 GI: 15150036 GI: 1486469摇 GI: 1486470摇
Swertia bimaculata GI: 15149903 GI: 15209282 GI: 15149931 GI: 15149967 GI: 15149996
Veratrilla baillonii GI: 15150001 GI: 15209266 GI: 15150019 GI: 13507073
Swertia punicea GI: 15150014 GI: 15209299 GI: 15150032 GI: 15150050 GI: 15150069
Swertia pubescens GI: 15150013 GI: 15209298 GI: 15150031 GI: 15150049 GI: 15150068
Swertia crassiuscula GI: 15150004 GI: 15209288 GI: 15150022 GI: 15150040 GI: 15150059
Comastoma pulmonarium GI: 15149908 GI: 15209303 GI: 15149936 GI: 15384516
Swertia cordata GI: 15150003 GI: 15209287 GI: 15150021 GI: 18152465 GI: 18152482
Swertia volkensii GI: 15150016 GI: 15209301 GI: 15150034 GI: 15150052 GI: 15150071
Gentiana frigida GI: 4092155摇 GI: 15209309 GI: 15149941 GI: 13235130 GI: 13235177
Gentiana phyllocalyx GI: 15149888 GI: 15209267 GI: 15149916 GI: 15149952 GI: 15149981
5933 期摇 摇 HE Ting鄄Nong et al. : A New Qinghai鄄Tibet Plateau Endemic Genus Sinoswertia and Its Pollination Mode摇 摇 摇
individuals in the 14 examined populations (correla鄄
tion index r= -0. 582; P<0. 05). With the altitudi鄄
nal increases, homogeneous plants with only small
flowers occurred more frequently at high altitude.
2. 2摇 Pollination mode
The seed sets of five pollination treatments were
listed in the Table 2. The direct isolations with the
bags indicated that both large flowers and small flow鄄
ers of the heterogeneous individuals and all small
flowers of the homogeneous plants always set numer鄄
ous seeds with high rates (>98% ). The seed sets of
these isolation treatments showed no distinct differ鄄
ence from the control treatments (P>0. 1). Howev鄄
er, the emasculation and hand鄄pollination treatments
still produced high rates of seeds, indicating that
outcrossing in this species is still fertile. The emas鄄
culation and bagged isolation produced no seeds,
which suggested that the agamospermy does not exist
in this species ( Table 2 ). Our pollination treat鄄
ments together suggested that all flowers from both
types of individuals have developed a perfect self鄄
pollination mechanism. In the field, we found all
flowers have nectaries, which can produce sweet se鄄
cretions. However, most flowers did not open, and
only 1-3% of the large flowers of the heterogeneous
plants were open in one monitored population when
they became mature with their pistil and stamens
equal in height. We found that only ants and thrips
visited these open flowers with very low frequency
( four times within the monitored 12 hours for two
open flowers). Because most flowers of this species
are self鄄pollinated without being open, it seems to be
a cleistogamous pollination mode (Lord, 1981).
2. 3 摇 Systematic position inferred from molecu鄄
lar evidence
We examined the sequence variations among 12
heterogeneous or homogeneous individuals at the
trnL鄄F, trnS鄄Ycf9, and ITS. We found no variation
in these sequenced fragments between different indi鄄
viduals of S. tetraptera, and all sequences were the
same as those obtained before (Table 1). We con鄄
structed the phylogeny trees based on two chloroplast
DNA fragments ( trnL鄄F and trnS鄄Ycf9) and nuclear
ITS respectively. Both phylogenetic trees ( Fig. 1)
suggested that S. tetraptera is closely related to Hale鄄
nia with high supports while the other Swertia repre鄄
sentative species clustered with the other genera as
indicated before (Yuan and K俟pfer, 1995; Chassot
et al., 2001; Liu et al., 2001; Hagen and Kadere鄄
it, 2002).
3摇 Discussions
In this study, we found two different types of
individuals in S. tetraptera, one heterogeneous type
with both large and small flowers and the other hom鄄
ogeneous individuals with only small flowers. This
dimorphic species was firstly recorded in the Swertia
and also rarely found in the family Gentianaceae.
Table 2摇 Seed sets under various treatments on both large and small flowers of the heterogeneous individuals
and small flowers of the homogeneous ones found in Sinoswertia tetraptera
Flower types Naturalpollination
Bagged
isolation
Emasculation for
natural pollination
Emasculation and
bagged isolation
Emasculation and
hand鄄pollination
Seed numbers per flower (Mean 依 SE) 13. 2 依 0. 5a 13. 1 依 0. 3a 2. 2 依 0. 2b 0 6. 8 依 1. 2c
Lf Seed set ratios (Mean 依 SE% ) >98% a >98% a <20% b 0 51. 8 依 6. 5% a
Numbers of flowers examined 159 151 15 12 18
Seed numbers per flower (Mean 依 SE) 5. 2 依 0. 2a 5. 3 依 0. 9a 0 0 1. 8 依 0. 2b
Sf Seed set ratios (Mean 依 SE% ) >98% a >98% a 0 0 <10% b
Numbers of flowers examined 224 10 12 10 14
Seed numbers per flower (Mean 依 SE) 5. 6 依 0. 2a 5. 8 依 0. 8a 0 0 1. 0 依 0. 2b
Shf Seed set ratios (Mean 依 SE% ) 100% a 100% a 0 0 <10% b
Numbers of flowers examined 108 25 8 10 8
Lf = large flowers on a main stem of a heterogeneous individual; Sf = small flowers on basal weak branches of a heterogeneous individual;
Shf = small flowers of the homogeneous individuals.
Values with the same superscript letter in the same column, or the same superscript number in the same row, do not differ significantly at the 0. 05 level
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Fig. 1摇 Two phylogenetic trees constructed for Sinoswertia tetraptera and related genera based on chloroplast DNA fragments and ITS sequences
We further found that most flowers were self鄄pollina鄄
ted through a cleistogamous mode with flowers
closed. This pollination mode and the increase of the
homogeneous individuals with altitude may have the
survival advantages for this species to occur in the
arid habitats (He et al., 2006; Duan et al., 2007,
2010) although such a probability needs further
tests. However, no dimorphic individual was recor鄄
ded for the other Swertia species and all species with
pollination mode studied in this genus were found to
be outcrossing with all flowers open at the flowering
stage ( Duan and Liu, 2007 and references cited
within it) . All molecular phylogenies based on dif鄄
ferent DNA sequences suggested that S. tetraptera is
sister to Halenia, a genus distributed in Asian
mountains but with its great diversification in the
southern America (Yuan and K俟pfer, 1995; Chas鄄
sot et al., 2001; Liu et al., 2001; Hagen and Ka鄄
dereit, 2002). In addition, as pointed out by Xue
et al. (1999a, b), S. tetraptera and Halenia share
two important embryological characters: orthotropous
ovules and well鄄developed hypostase. However, in
the remaining studied Swertia species, ovules are a鄄
natropous or ana鄄campylotropous and both placenta
intrusion and hypostase are not distinctly developed
(Xue et al., 2002a, b). All these lines of evidence
suggest that S. tetraptera should be established as one
separate genus, sister to the well鄄defined Halenia.
However, Halenia differs from this new monotypic
genus with spurred corolla.
After excluding S. tetraptera, the remaining
species or sections in the genus Swertia seem still to
be polyphyletic. The representative species sampled
from this genus for phylogenetic studies, clustered
respectively with the other well鄄circumscribed gene鄄
ra, for example, Comastoma and Veratrilla (Fig. 1
and also see Chassot et al., 2001 and Hagen and
Kadereit, 2002). Although the clade represented by
S. bimaculata is sister to Halenia鄄S. tetraptera, it is
similar to the other Swertia species or sections with
rotate corolla and nectaries surrounded by fringed or
tubular appendages. Swertia is also similar to Loma鄄
gogonium, but the latter differs from it with decur鄄
rent stigma (Ho and Pringle, 1995). In addition,
all these genera differ from one another in both mor鄄
phological and embryological traits ( Xue et al.,
1999a, b, 2002a, b and references in them) (Ta鄄
ble 3). The rotate and deeply lobed corolla and di鄄
verse nectaries shared by S. tetraptera, the other
Swertia species and the other genera may have resul鄄
ted from the convergent evolutions due to common
selection pressure during the rapid diversification of
Swertia in the Qinghai鄄Tibet Plateau and adjacent re鄄
gions. In fact, such a scenario was also found for
other genera. For example, in the Ligularia鄄Crema鄄
nthodium鄄Parasenecio complex (Asteraceae), inter鄄
generic circumscriptions are extremely difficult be鄄
cause these three species鄄abundant genera with dia鄄
7933 期摇 摇 HE Ting鄄Nong et al. : A New Qinghai鄄Tibet Plateau Endemic Genus Sinoswertia and Its Pollination Mode摇 摇 摇
gnostic morphological traits are polyphyletic because
of radiative diversification and common selection
pressure (Liu et al., 2006). The taxonomic treat鄄
ment of the polyphyletic Swertia in the current cir鄄
cumscription is still a challenge at the present and
pends further studies in the future.
4摇 Taxonomic treatments
Sinoswertia T. N. Ho, S. W. Liu & J. Q. Liu
gen. nov.
—Type: Sinoswertia tetraptera (Maximowicz)
T. N. Ho & S. W. Liu
Swertia sect. Heteranthos T. N. He & S. W.
Liu, Acta Phytotax. Sin. 18 (1 ): 85 (1980 ).
Type: S. tetraptera Maxim.
This new genus is closely related to Swertia s.
l. and Halenia. It differs from the latter by spurless
corolla and from both by dimorphic plants and di鄄
morphic flowers (Fig. 2).
A monotypic genus endemic to the northeastern
Qinghai鄄Tibet Plateau (Fig. 3).
Sinoswertia tetraptera ( Maximowicz ) T. N.
Ho, S. W. Liu & J. Q. Liu, comb. nov.
=Swertia tetraptera Maxim., Bull. Acad. Imp.
Sci. St. 鄄P佴tersbourg 27: 503 (1881). Type: Chi鄄
na, Gansu (Kansu), in shrubs, in 1872-1880, Przew鄄
alski s. n. (LE鄄holotype, not seen; PE! 鄄isotype).
= Swertia dimorpha Batalin, Act. Hort. Petrop.
13: 379 (1894). Type: China, N Sichuan (Szechuan),
[Nanpin鄄Songpan鄄Pingwu], Dshangla, Nereku, 25
July 1885, G. N. Potanin s. n. ( LE鄄holotype, not
seen; HNWP! 鄄photo. ) .
= Swertia pusilla Diels, Notizbl. Bot. Gart.
Berlin 11: 215 (1931), non Pursh (1814). Type:
China, W Ningxia (Kansu), Mt. Helan Shan (Ho
Shan), He鄄lan鄄wu鄄kou, 2 000 m, 11 May 1923,
R. C. Ching 70 (US! 鄄holotype).
=Anagallidium dimorpha (Batalin) Ma. Acta
Phytotax. Sin. 14: 65 (1976).
Additionalis description homogeneae plantae:
Plantae homogeneae nanae, (1-) 3-8 (-15) cm.
Caules flaccidi, graciles, fere quadrangulares, pau鄄
co鄄ramosi. Folia 2-4鄄binata, late disposita; petioli
usque 1 cm; laminae 5 -10 伊3 -8 mm, apice obtu鄄
sae, base angustatae, nervis 1 -3, prominentibus.
Cymae pauciflorae. Pedicelli 5-50 mm. Flores uni鄄
formes, parvi. Calyx corollaque lobata fere ad ba鄄
ses; tubus calycis 0. 3-0. 5 mm, lobis 1. 5-4 mm,
late ovatis, apice acutis, costis prominentibus; co鄄
rolla viridis usque flavido鄄viridis, interdum purpura鄄
to鄄tingens, tubo 0. 3-0. 5 mm, lobis 2. 5-5 mm, obo鄄
vato鄄oblongis usque oblongis, apice subroundatis ero鄄
sisque, base contractis; nectaria 2 in unoquoque lobo
corollae, una squama integra late oblonga et maculis
indistinctis instructa; filamenta 1-1. 5 mm, linearia,
complanata, base una squama semiorbiculata ornata;
antherae 0. 5 -1 mm, ellipsoideae, flavidae; pistil鄄
lum 1. 2-1. 7 mm; stylus indistinctus; lobi stigmatis
capitati. Capsulae 4 - 5 mm, late ovoideae usque
subglobosae. Semina numero (2-) 4-6 (-4) vari鄄
antia, (0. 8-) 1. 2 -1. 5 伊0. 9 -1. 1 mm, flavida,
late ellipsoidea usque subglobosa, tenuiter reticulata
usque fere laevia.
Plantae homogeneae a eis heterogeneis caulibus
nanis, pauci鄄ramosis, absque debilibus ramosis,
omnibus floribus parvis et uniformibus recedit.
Table 3摇 Diagnostic features of Sinoswertia, the other Swertia sections, Halenia and Lomatogonium
Characters Sinoswertia Other Swertia, sections Halenia Lomatogonium
Dimorphic individuals Yes No No No
Dimorphic flowers Yes No No No
Corolla spur No No Distinct No
Stigma Normal Normal Normal Decurrent
Placenta intrusion Strong No Strong No
Ovule Orthotropous Anatropous (ana鄄campylotropous) Orthotropous Campylotropous
Hypostase Developed No Developed No
Pollination model Complete cleistogamy Outcrossing Outcrossing or self鄄pollination Outcrossing
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Fig. 2摇 Sinoswertia tetraptera. A. a heterogeneous individual with large and small flowers; B: a large flower of the heterogeneous individual;
C: a corolla lobe; D: a homogeneous plant only with small flowers; E: a small flower of a homogeneous individual;
F: a part of opened corolla lobes of a small flower
Fig. 3摇 Distributional range of Sinoswertia tetraptera
in the Qinghai鄄Tibet Plateau
摇 摇 Distribution and habitat: endemic to Qinghai鄄
Tibet Plateau ( Fig. 3), including northeastern Ti鄄
bet, eastern and southwestern Qinghai, northwestern
Sichuan, and Gansu, It grows in diverse environ鄄
ments, i. e., alpine meadows, alpine shrub鄄mead鄄
ows, shrubs, scattered forests, grasslands, wet
places near to the river, stream or gutter, roadsides,
gravelly lands, and stone pits, mainly at elevations
between 2 000 m and 4 310 m.
Representative specimens. CHINA, Qing鄄
hai: Minhe, Ho T. N., Liu S. W. & Wang Z. Q.
110 (HNWP); Datong, = Ho T. N., Liu S. W. &
Wang Z. Q. 18 (HNWP); Ping爷an, Ho T. N., Liu
S. W. & Wang Z. Q. 34 (HNWP); Xunhua, Ho
T. N., Liu S. W. & Wang Z. Q. 91 ( HNWP);
Huangyuan Ho T. N., Liu S. W. & Wang Z. Q. 117
(HNWP); Gonghe, Ho T. N., Liu S. W. & Wang
Z. Q. 41 (HNWP); Chindu, Ho T. N., Liu S. W.
& Wang Z. Q. 31 (HNWP); Yushu, Ho T. N.,
Liu S. W. & Wang Z. Q. 135 (HNWP). Tibet:
9933 期摇 摇 HE Ting鄄Nong et al. : A New Qinghai鄄Tibet Plateau Endemic Genus Sinoswertia and Its Pollination Mode摇 摇 摇
Jomda, Ho T. N., Liu S. W. & Wang Z. Q. 166
( HNWP ). Gansu: Zuoni, LiuJQ鄄2012鄄GN鄄162
(LZU, KIB); Maqu, LiuJQ鄄GN鄄2011鄄011 ( LZU,
KIB); Hezuo, LiuJQ鄄GN鄄2011鄄022 (LZU, KIB);
Sichuan: Serxu, Ho T. N., Liu S. W. & Wang Z.
Q. 168 (HNWP); Dege, Ho T. N. , Liu S. W. &
Wang Z. Q. 167 (HNWP).
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