POLYPLOIDY IN THE FLORA
OF THE HENGDUAN
MOUNTAINS HOTSPOT,
SOUTHWESTERN CHINA 1
Ze-Long Nie, 2 Jun Wen, 3 Zhi-Jian Gu, 2
David E. Boufford, 4 and Hang Sun 2
ABSTRACT
The Hengduan Mountain region is located at the eastern end of the Himalayan range and in the southeastern corner
of the Qinghai-Tibet Plateau in eastern Asia. It is probably the richest temperate region in the world biologically, with
about 8000 species recorded from the 500,000 km2 region. A high rate of polyploidy was expected in the Hengduan
Mountains because of the unusually high species diversity and endemism, the widespread alpine environment in this
region, and the importance of polyploidy in plant evolution. Chromosome numbers for 552 taxa of native Hengduan
angiosperms were obtained from published literature. If the criterion of infrageneric polyploidy is used (i.e., the possession of three or more basic chromosome sets in a nucleus), the frequency of polyploidy is only 22%. It is 37% and
58% if taxa with haploid chromosome number n 5 14 or more, or n 5 11 or more are considered to be polyploids,
respectively. Several highly diversified groups such as Aconitum and Delphinium also show a low proportion of polyploidy (18.6% and 0%, respectively) in the Hengduan Mountains. Examples from other large plant groups (e.g., Saxifraga, Ligularia, and Cremanthodium) and endemic genera (e.g., Solms-laubachia, Tibetia, and Nomocharis) again
indicated that polyploidy, especially infrageneric polyploidy, may have played a minor role in the evolutionary diversification of these plants in the Hengduan Mountains.
Key words: chromosome number, diversification, Hengduan Mountains, Hotspot, polyploidy.
The Hengduan Mountains are located at the eastern end of the Himalayan range and in the southeastern corner of the Qinghai-Tibet Plateau, the
highest and youngest plateau in the world. The region comprises more than 500,000 km2 of temperate to alpine mountains in southwestern China and
is bounded in the northwest by the dry QinghaiTibet Plateau, in the north by the Tao River of
southern Gansu, and in the east by the Sichuan
Basin and the plateau of eastern Yunnan. It also
includes small portions of extreme northern Myanmar and extreme eastern India. About 8000 species
of flowering plants have been recorded from this
region (X. W. Li & Li, 1993; W. T. Wang et al.,
1993, 1994), making it one of the two most speciose
floristic subregions of the Sino-Himalayan Floristic
Region, the other being the East Himalayan subregion (Wu, 1979, 1988; X. W. Li, 1994). The region is also renowned as the home of the giant panda (Wu, 1988). Based on a series of expeditions to
the region during 1981–1983 organized by the Chinese Academy of Sciences (Wu, 1988), W. T. Wang
et al. (1993, 1994) and X. W. Li and Li (1993)
published a preliminary appraisal of the flora that
documented its high levels of species diversity and
endemism. Although the Hengduan Mountains are
still poorly known to the western scientific community, they are biologically among the world’s
richest and most diverse regions (Boufford & van
Dyck, 1999) and have recently been recognized as
the biologically richest temperate region in the
world, characterized by the occurrence of some of
1
This research was supported by grants from the Chinese Natural Science Foundation (CNSF, grant numbers
30300023 to Z.-L. Nie, 40332021 and 30270121 to H. Sun), the Innovation Project of the Chinese Academy of Sciences
(CAS, KSCX2-1-09 to H. Sun), in part by a grant from the Biotic Surveys and Inventory, National Science Foundation,
U.S.A. (DEB-9705795, M. J. Donoghue, Principal Investigator) for fieldwork in the Hengduan Mountains, and grants
from the Excellent Overseas Young Scientists Program of CAS and the National Science Foundation (DEB 0108536)
to J. Wen. We thank Eve Emshwiller, Pete Lowry, and editor Victoria Hollowell for their most constructive suggestions
to improve our study.
2
Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan 650204, People’s Republic of China.
hsun@mail.kib.ac.cn (H. Sun).
3
Department of Botany, Field Museum of Natural History, 1400 S. Lake Shore Dr., Chicago, Illinois 60605-2496,
U.S.A., and Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences,
Nanxincun 20, Xiangshan, Beijing 100093, People’s Republic of China. wen@fieldmuseum.org; current address: Department of Botany, National Museum of Natural History, MRC-166, Smithsonian Institution, P.O. Box 37012, Washington, D.C. 20013-7012, U.S.A.
4
Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138-2020, U.S.A. boufford@oeb.harvard.edu.
ANN. MISSOURI BOT. GARD. 92: 275–306. 2005.
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the world’s most distinctive species, such as
Acanthochlamys bracteata P. C. Kao (Velloziaceae),
Sinadoxa corydalifolia C. Y. Wu et al. (Adoxaceae),
Salweenia wardii E. G. Baker (Fabaceae), and Takakia ceratophylla (Mitt.) Grolle (Bryophyta). Other
floristic characteristics of the range include great
regional radiations within a large number of plant
groups, and the occurrence of a large number of
relict plant lineages (Ying & Zhang, 1984; Wu,
1988; H. S. Wang, 1989; Wilson, 1992; X. W. Li
& Li, 1993; Ying et al., 1993; H. S. Wang & Zhang,
1994; Boufford & van Dyck, 1999; H. Sun, 2002a,
b; H. Sun & Li, 2003). The area was previously
included in the Eastern Himalayan ‘‘hot spot’’ (Myers, 1988; Wilson, 1992), but more recently has
been referred to as the South-Central China Hotspot (Myers et al., 2000) or the Hengduan Mountains Hotspot (Boufford & van Dyck, 1999). This
Hotspot is a focus for studies of relationships between florogenesis and plate tectonics, on the origins of the north temperate flora, including the floras of eastern Asia, North America, and Europe,
and on the evolution of seed plants in general (Wu,
1988).
Polyploidy, the possession of three or more basic
chromosome sets in nuclei (Bennett, 2004), has
long been recognized as a prominent force in the
evolutionary diversification of plants (Winge, 1917;
Darlington, 1937; Stebbins, 1940, 1950, 1971;
Grant, 1981; Levin, 2002), especially ferns (Wagner, 1980; Werth et al., 1985) and flowering plants
(Lewis, 1980; Soltis & Soltis, 2000; Wendel, 2000).
This phenomenon occurs in up to 80% of all angiosperm species and is considered to be one of the
most important cytogenetic mechanisms in plant
evolution (Masterson, 1994; Levin, 2002), especially in facilitating rapid speciation (Hiremath &
Salimath, 1991).
At the beginning of this research, our working
hypothesis was that a high proportion of polyploids
occurs in the angiosperm flora of the Hengduan
Mountains. Below we outline the reasoning behind
this hypothesis and provide a general introduction
on the physical and biological characteristics of the
Hengduan Mountains Hotspot.
The Hengduan Mountains are geologically
young, yet possess an extremely rich flora both in
terms of overall species diversity and endemics.
The Hengduan Mountains rose relatively rapidly
from sea level to the present high mountains (up to
7600 m) with rugged and dissected topography following the collision of the Eurasian and Indian
plates about 50 million years ago (Patriat &
Achache, 1984; C. S. Wang & Ding, 1998; Tao,
1992, 2000). The most common mechanism for rap-
id speciation in plants is through polyploidy (Grant,
1981). Recent studies in several plant groups have
revealed that the genesis of allopolyploidy under
both synthetic and natural conditions is often accompanied by rapid genetic and sometimes evolutionarily conserved epigenetic changes (B. Liu et
al., 1998; Ozkan et al., 2001; B. Liu & Wendel,
2003; Osborn et al., 2003; Soltis et al., 2004). The
capacity of angiosperms to form new combinations
via polyploidy may help explain rapid diversification and high endemism in a given region with high
biodiversity. Polyploidization might thus account
for the high species diversity and endemism in the
Hengduan region.
In regions such as the Hengduan Mountains with
highly diversified floras, a high frequency of polyploidy would also be expected because of the crucial role of polyploidy in plant speciation (Grant,
1981). Polyploids can have more than two different
alleles at any given locus. The intrinsic advantages
of heterozygosity may be a key factor in the growth,
performance, and adaptability of a polyploid
(Grant, 1981; Levin, 1983; Ramsey & Schemske,
1998, 2002; Soltis et al., 2004). Polyploid populations often demonstrate extensive genomic rearrangement including the origin of novel regions of
DNA (Song et al., 1995; Arnold, 1997).
Alpine and arctic floras have been reported to
have high frequencies of polyploidy (Hanelt, 1966;
Löve & Löve, 1967, 1975; Ohba, 1988; Abbott &
Brochmann, 2003; Brochmann et al., 2004). Likewise, polyploids are more common in cold climates
associated with harsh environmental conditions
(Löve & Löve, 1949, 1967). Grant (1981) also suggested that disturbed habitats and stressful environments, which are often associated with cold climates, usually favor polyploidy. The great
ecological amplitude that polyploid species can exhibit gives them a high degree of buffering against
environmental changes over long periods of time
due to glaciation, mountain building and degradation, and overall fluctuations in climate (Stebbins,
1971; Grant, 1981). The Hengduan region has extensive alpine areas. Environmental extremes such
as cold and severe climate, and nutrient stress in
high latitude or altitude areas have been suggested
to increase the frequency of 2n gamete formation
(Thompson & Lumaret, 1992; Levin, 2002), which
leads to polyploidy.
Polyploids have been suggested to be more adaptive in disturbed or novel niches than their diploid
progenitors (Ehrendorfer, 1980; Grant, 1981; Stebbins, 1985; Morton, 1993; Otto & Whitton, 2000;
Levin, 2004). The Tertiary Himalayan orogeny has
led to changes in the climate and topography of the
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2005
Nie et al.
Polyploidy in Regional Flora
Hengduan Mountains (J. J. Li et al., 1995; Shi et
al., 1998), whose uplifting and the advance and
retreat of ice sheets and glaciers may have provided
opportunities for the establishment of polyploids
through hybridization and other mechanisms in the
contact zones between existing species. The advantages and fitness polyploidy can bestow on organisms will be most effective when new habitats are
being created and become available for colonization (Morton, 1993). Ehrendorfer (1980) also surveyed various plant communities to determine polyploid frequency in the flora of lower Austria, and
found that the highest proportion of polyploid species was found in disturbed areas.
The objectives of this paper are to: (1) present a
comprehensive review of chromosome numbers for
the flora of the Hengduan Mountains, (2) calculate
the frequency of polyploidy in this region and in
exemplar taxa, and (3) test the hypothesis that polyploidy is important in the evolutionary diversification of its rich angiosperm flora.
number regardless of the ploidy level, while x is
the most probable base number. The large number
of species and the paucity of taxonomic treatments
for the study region made it difficult to assign previously published records to currently recognized
taxa. We thus report chromosome numbers under
the names that were originally published.
METHODS
AND
TERMINOLOGY
For each taxon (including species and subspecies), chromosome numbers were recorded from
published reports. Several chromosome atlases
were also consulted (Darlington & Wylie, 1955;
Bolkhovskikh et al., 1969; Moore, 1973, 1974,
1977; Goldblatt, 1981, 1984, 1985, 1988; Goldblatt & Johnson, 1990, 1991, 1994, 1996, 1998,
2000, 2003).
We employed three methods to evaluate polyploids. First, taxa with three or more basic chromosome sets in nuclei are regarded as polyploids
(Bennett, 2004), i.e., as resulting from infrageneric
polyploidy (Stebbins, 1950). The second method
recognizes polyploidy based on the criterion of haploid number n 5 14 or more for statistical purposes
(Grant, 1963, also see 1981: 296). Goldblatt (1980)
suggested a third method, arguing that Grant’s
(1963) estimate was too conservative, and proposed
that taxa with haploid number n 5 11 or more
should be considered as polyploids. In our study,
we have employed all three methods to estimate the
incidence of polyploidy in the Hengduan region.
We emphasize infrageneric polyploidy in our discussion, because we consider this measure to be
the most accurate of the three. We also inferred the
ploidal level for each taxon based on Stebbins’s
(1950) criterion (results shown in Appendix 1). A
few taxa with both polyploid and diploid counts are
treated as polyploids in our calculation.
In this paper, 2n refers to a taxon’s somatic chromosome number and n to the gametic chromosome
FREQUENCY
MOUNTAINS
OF
POLYPLOIDY
IN THE
277
HENGDUAN
Chromosome counts for 552 species and infraspecific taxa in 152 genera and 44 families from
the Hengduan Mountains are shown in Appendix
1. Most taxa are representatives of genera that are
taxonomically diversified and ecologically common
in the region, such as Rhododendron, Silene, Arenaria, Rubus, Ligularia, Delphinium, Poa, Astragalus, Saussurea, Aconitum, Primula, Gentiana, and
Pedicularis, all of which are characterized by regional radiation. Others, such as Acanthochlamys,
Beesia, Circaeaster, Dichotomanthes, Kingdonia,
Nomocharis, Pomatosace, Przewalskia, Salweenia,
Solms-laubachia, and Souliea, are largely endemics
to the region. At present no chromosome counts are
available for certain large families, such as Lamiaceae, Cyperaceae, and Crassulaceae.
When polyploidy is calculated based on three or
more basic chromosome sets in nuclei recorded in
a genus (infrageneric polyploidy), only 22% (124/
552) of the angiosperm taxa sampled in the Hengduan Mountains are polyploids. Using this same
method, 30%–35% are estimated to be polyploids
for all flowering plants whose chromosome numbers
have been counted (Stebbins, 1950) and 36% for
monocots (Goldblatt, 1980: 225).
Based on the criterion that haploid number n 5
14 or more denotes polyploidy sensu Grant (1963,
1981), 37% (204/552) of the angiosperm taxa from
the Hengduan Mountains are polyploids (Table 1).
These values are relatively low compared with previous estimates. Grant (1963) reported that 47% of
the 17,138 species of flowering plants he sampled
were polyploids. According to the method of Goldblatt (1980) with n 5 11 or above as polyploids,
58% (322/552) of the angiosperm taxa in the Hengduan region are polyploids. Using the same criterion, Goldblatt (1980) reported that 68% of the
10,580 species of monocots he surveyed were polyploids.
COMPARISONS
WITH
OTHER REGIONS
Because of the limited sample size of our chromosome data (ca. 6.9% of the estimated 8000 species present in the Hengduan Mountains), we com-
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Annals of the
Missouri Botanical Garden
Table 1. Polyploidy statistics of angiosperms in the Hengduan Mountains based on three methods: (a) infrageneric
polyploids—taxa with three or more base chromosome number sets (Stebbins, 1950; Bennett, 2004); (b) haploid number
n 5 14 or more (Grant, 1963); (c) n 5 11 or more (Goldblatt, 1980).
Total species (%)
All species sampled
Endemic species
Infrageneric polyploidy Polyploidy (n $ 14)
Species number (%) Species number (%)
552
183 (33)
124 (22)
29 (16)
pared our findings with chromosome data for the
plants of Pakistan. Pakistan is located on the western end of the Himalayan range. The chromosome
data from Pakistan are based on a larger sample
size, i.e., 1777 species and infraspecific taxa sampled (34% of its total angiosperm flora). As shown
in Table 2, the frequency of polyploidy in Pakistan
is similar to what we found for the Hengduan
Mountains, with 29% infrageneric polyploidy, and
44% (n 5 14 or more) or 63% (n $ 11) based on
the two methods for estimating polyploidy, as calculated using data taken from Khatoon and Ali
(1993). The similar incidence of polyploidy in these
two regions may reflect a broader characteristic of
polyploid distribution in the Himalayan range. Although the sample size in our study includes only
a small proportion of the angiosperm flora of the
Hengduan Mountains, it nevertheless provides good
taxonomic and ecological representation. The available data suggest that the flora of the Hengduan
Mountains is characterized by a relatively low frequency of polyploidy.
As indicated above, the frequency of polyploidy
is higher in arctic floras than those at lower latitudes (Hanelt, 1966; Löve & Löve, 1975; Abbott &
Brochmann, 2003). The frequency of polyploid species in the vascular flora of the arctic region as a
Table 2.
Polyploidy (n $ 11)
Species number (%)
204 (37)
49 (27)
322 (58)
89 (49)
whole is nearly 60% in the low arctic zone, but
70% in the high arctic and as high as 80% in higharctic endemics (Löve & Löve, 1975). Brochmann
and Steen’s (1999) research on the isolated arctic
archipelago of Svalbard, which was almost completely ice-covered during the last glaciation,
showed that nearly 80% of the 161 native species
are polyploids. A recent review on arctic plants
based on 1719 taxa showed that the majority
(60.7%) are polyploids, especially for arctic specialist taxa (73.7%) (Brochmann et al., 2004).
Löve and Löve (1967) pointed out that the frequency of polyploidy is significantly high in the
alpine zone, as on Mt. Washington in the White
Mountains of New Hampshire, where 63.6% of the
alpine taxa of vascular plants are polyploids. Another example is from the flora of the Cameroon
Mountains (Morton, 1993), which shows a frequency of 52.9%, higher than generally reported (sensu
Grant, 1981). Hanelt (1966) gave percentages ranging from 45% to 85% polyploids in various high
mountain floras in Eurasia, the Americas, and New
Guinea.
Our finding of only 22% polyploidy in the angiosperm flora of the Hengduan Mountains based
on the same method of calculation (Stebbins, 1950)
stands in sharp contrast with these earlier results.
Comparison of frequency of polyploidy among selected regions or plant groups.
Reference
Hengduan Mountains
Pakistan
Monocots
Angiosperms
High mountain floras
Cameroon Mountains
Mt. Washington
Arctic flora
Svalbard
Juan Fernandez Islands
Hawaii
Khatoon & Ali (1993)
Goldblatt (1980)
Stebbins (1950)
Grant (1963, 1981)
Hanelt (1966)
Morton (1993)
Löve & Löve (1967)
Löve & Löve (1975)
Brochmann et al. (2004)
Brochmann & Steen (1999)
Sanders et al. (1983)
Carr (1998)
Infrageneric
polyploidy, %
22
29
36
30–35
Polyploidy
(n $ 14), %
Polyploidy
(n $ 11), %
37
44
58
63
68
47
45–85
52.9
63.6
60–80
60.7
ca. 80
75
80
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Nie et al.
Polyploidy in Regional Flora
A similar survey on Chihuahuan Desert flora
showed that a relatively low frequency of 17.8%
polyploidy occurs among the gypsophilic species
compared to 32.7% polyploidy for the non-gypsum
vegetation (Powell & Sloan, 1975). Further chromosome reports on some species from the Hohxil
region of Qinghai, a harsh, cold, uninhabited, alpine desert at an average elevation of 5000 m near
the center of the Qinghai-Xizang Plateau, also
showed that most plants there are diploids (Y. P.
Yang & Wu, 1993).
Several studies using the more inclusive criterion
that taxa with gametic numbers of n 5 14 and
above are regarded as polyploids (Grant, 1963)
have found higher levels of polyploidy than the
58% we estimated for the Hengduan region. For
example, Sanders et al. (1983) reported 75% polyploidy for the flora of the Juan Fernandez Islands,
and Carr (1998) reported 80% for Hawaiian plants.
era in this region has resulted mainly from speciation within diploid lineages.
Ligularia and Cremanthodium (Asteraceae) are
two related genera that are also highly diversified
in the Hengduan Mountains (Liu, 2004). Ligularia
has 129 species in the region, with about 100 native and 60 endemic. This genus is morphologically
diverse and has members in a variety of habitats,
from forests to high alpine meadows, ranging from
1000 to 4000 m in altitude, although most species
are found in the mountains (S. W. Liu et al., 1994).
Cremanthodium comprises 67 species growing in
alpine meadows, largely in the Hengduan Mountains and on the Qinghai-Tibet Plateau. Although
these genera do not qualify for Stebbins’s (1950)
definition of infrageneric polyploidy, their high haploid chromosome numbers of n 5 30 and n 5 29
suggest ancient polyploidizations. Variation of karyotype structure at the diploid level seems to be
the predominant feature in these groups, and sympatric speciation via hybridization and polyploidization has played a minor role in their species diversity (J. Q. Liu, 2004).
A similar phenomenon can be seen in several
other groups occurring around or within the Himalaya and Qinghai-Tibet Plateau. Saxifraga (Saxifragaceae) is widely distributed in the temperate
zone of the Northern Hemisphere and is one of the
most important components of the alpine flora of
the Himalayan region (Ohba, 1986). In particular,
some species are quite diverse in the high alpine
region of the Himalaya as well as in the Hengduan
Mountains. In the Hengduan Mountains, four of the
nine species reported are polyploids (44.4%). However, cytological studies of Saxifraga in the Himalayas by Wakabayashi and Ohba (1988) showed
that diploids prevail and polyploids comprise only
17% of the species. Polyploidy thus may not have
played an important role in the diversification of
Saxifraga in the Himalayan range (Wakabayashi &
Ohba, 1988).
POLYPLOIDY
IN
REPRESENTATIVE TAXA
We expected to find a high level of polyploidy
before we began this study, but it seems that polyploidy is not common in the Hengduan Mountains
based on regional cytological statistical comparisons. This result is even more pronounced when
one considers the frequency of polyploidy in several representative taxa from the Hengduan Mountains Hotspot.
HIGHLY DIVERSIFIED GROUPS
Aconitum and Delphinium are two closely related
genera of Ranunculaceae in the tribe Delphineae
and are widely distributed in north temperate areas.
About half of the species of Aconitum and more
than 63% of the species of Delphinium are found
in the Hengduan Mountains (Q. E. Yang et al.,
1989). The species in these two genera are highly
diversified in the region, with both primitive and
advanced representatives, including many local endemics (X. W. Li & Li, 1993; Q. E. Yang, 1996).
The cytological data reveal that of 43 species in
Aconitum in the Hengduan region (41% of the total
104 species), 30 are endemic to the area, and only
eight are polyploids (18.6%). There are no polyploids among the 23 species (11 endemics) of Delphinium reported (32% of the total 71 species in
the region). However, in Simon et al.’s (1999) comprehensive chromosome survey of Delphineae,
which included about 40% of species in the tribe,
48% and 31% of Aconitum and Delphinium species
were found to be polyploids, respectively. These figures are much higher than our estimates from the
Hengduan region. The high diversity of these gen-
279
ENDEMIC TAXA
High levels of endemism are a major feature of
the flora of the Hengduan Mountains. About 37.7%
of the angiosperms are endemic or mainly restricted
to this area (X. W. Li & Li, 1993). Of the 552 taxa
sampled, 33% are endemic to the Hengduan region
(Table 1), a proportion similar to that of the total
angiosperms in the region. But infrageneric polyploidy is particularly low among endemics (just
16%). Below we discuss several endemic genera to
illustrate this phenomenon.
The nine species of Solms-laubachia (Brassica-
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ceae) are nearly restricted to scree slopes and rocky
crevices at altitudes of 3400–5700 m in the Hengduan Mountains (Al-Shehbaz & Yang, 2001; Yue et
al., 2004). All species are 2n 5 14, although one
species (S. pulcherrima Muschl.) has a tetraploid
cytotype (Yue et al., 2003, 2004). Speciation within
Solms-laubachia may have been the result of primarily intrachromosomal variation at the diploid
level, and the genus probably evolved within the
Himalayan-Hengduan Mountains instead of migrating from other regions (Yue et al., 2004).
Another example is the small, localized, nearly
endemic genus Tibetia (Fabaceae). This genus apparently differentiated from the widely distributed
north temperate Gueldenstaedtia and became
adapted to the alpine environment (Tsui, 1979; Nie
et al., 2002) as the Hengduan Mountains uplifted.
The cytological studies of Nie (2002), Nie et al.
(2002), and D. K. Yang (2002) show that the species of Tibetia have all evolved at the diploid level
of 2n 5 16, but with alterations of their karyotypes.
These authors suggest the importance of allopatric
speciation at the diploid level in Tibetia in the
Hengduan Mountains.
Nomocharis (Liliaceae), comprising ca. 10 species with the Sino-Himalayan distribution pattern,
is restricted to the Hengduan Mountains and the
southeastern edge of the Tibetan Plateau. All its
members are not infrageneric polyploids, with 2n
5 24, and appear to have diversified at the diploid
level after ancient polyploidization (Xie et al.,
1992). Other taxa, such as Salweenia (Q. X. Zhou
et al., 2000), Beesia (Q. E. Yang et al., 1995), and
Kingdonia (Z. Y. Zhang, 1982), are likewise all diploids that have developed and become localized in
the Hengduan Mountains area.
nus of Adoxaceae from the Hengduan region, is
closely related to Adoxa (J. Q. Liu et al., 2000b),
but is morphologically very distinct, despite low
ITS sequence divergence (3.4%), which suggests a
recent origin of Sinadoxa. Both genera have the
same chromosome number of 2n 5 36 and a similar
karyotype, and the initial differentiation between
them might have resulted from allopatric isolation
when the first large-scale uplifting of the QinghaiTibet Plateau occurred about 3.4–1.7 million years
ago (J. Q. Liu et al., 1999, 2000b).
Geographical and ecological heterogeneity may
have played an important role in the diversification
of plants in the Hengduan Mountains. Most parts
of the region are characterized by a series of parallel mountain ranges dissected by deep river valleys that run from north to south, which present
physical barriers to gene flow. The high speciation
rate in the Hengduan flora may also be due to highly varied local climates that result from geographic
differences in the mountains. The wide diversity of
climates combined with local differences in geomorphological, edaphic, and ecological features
may have generated distinct niches for different
species. Rhododendron (Ericaceae), for example, is
the largest genus in the region with about 220 species extending throughout the Hengduan area, but
most species have highly localized distributions.
Both diploids and polyploids are common among
Rhododendron in the Hengduan Mountains, with a
high base chromosome number of x 5 13 (Darlington & Wylie, 1955; Gao, 2002). The radiation of
Rhododendron in this region is perhaps related to
geographic heterogeneity in conjunction with polyploidy.
Animals such as insects may have played a vital
role in the dispersal as well as the evolution of
reproductive isolation of some groups in the region.
Pedicularis is a circumpolar genus of Scrophulariaceae, consisting of approximately 500 species,
with about 200 occurring in the Hengduan Mountains, making it the second largest genus of angiosperms in the region (X. W. Li & Li, 1993). Pedicularis shows extreme variation in floral
morphology that is correlated with specific mechanisms of pollination involving a diverse group of
insects. Chromosome numbers of Pedicularis recorded in the Hengduan Mountains are 2n 5 14
and 16 (J. Cai et al., 2004), suggesting that the
evolution has taken place at the diploid level and
that diversification is perhaps facilitated by the behavior of pollinators.
Climatic changes during the Pleistocene glacial
and interglacial periods may have resulted in the
formation of many refugia in the region; these re-
EVOLUTION OF HIGH SPECIES DIVERSITY
HENGDUAN MOUNTAINS HOTSPOT
IN THE
If polyploidy did not play a major role in the
evolution of Hengduan plants, what other mechanisms might have shaped the substantially high
species diversity and endemisms in the region? Below we propose a few hypotheses based on physical
and historical factors.
Alpine plants survive under the environmental
conditions they encounter because of adaptive features such as cushion or rosette habit, pubescence,
and coriaceous leaves (Chaverri-Polini, 1998; Körner, 2003). Morphological differentiation can be accumulated in a relatively short period under certain
alpine environments through natural selection,
without drastic genetic or chromosomal divergence.
For example, Sinadoxa, an endemic monotypic ge-
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Polyploidy in Regional Flora
fugia may have served as shelters for a rich diversity of species, as well as sources for dispersion to
neighboring areas during the warm interval. The
arctic and Tethyan Tertiary floras also served as a
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CONCLUSIONS
Polyploidy (especially infrageneric polyploidy) is
not common in the alpine flora of the Hengduan
Mountains Hotspot based on our cytological statistical analysis. This finding is further strengthened
by case studies of representative taxa in the region.
High levels of speciation activities in the Hotspot
are probably attributable at least in large part to
the geographic, climatic, and geological heterogeneity that resulted from the Tertiary Himalayan
orogeny and Cenozoic ice-age cycles. Further investigations, particularly of taxa characteristic of
the area, are needed to (a) further test our findings
of low frequency of polyploids in the Hengduan region in a phylogenetic framework, (b) examine other prominent evolutionary forces involved in speciation in the Hotspot, and (c) explore the possible
reasons for the apparently low rate of formation
and/or survival of polyploid species. We hope this
cytological study will stimulate further chromosomal and evolutionary work on the Hengduan Mountains Hotspot, and on other conservation hotspots
as well.
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Appendix 1. Chromosome reports and ploidal levels in angiosperms of the Hengduan Mountains. 2n 5 somatic
chromosome number; n 5 gametic or haploid chromosome number; x 5 chromosome base number; GS 5 Gansu; QH
5 Qinghai; SC 5 Sichuan; XZ 5 Xizang; YN 5 Yunnan Province, China.
Taxon
ADOXACEAE
Sinadoxa corydalifolia Z. Y. Wu, Z.
L. Wu & R. F. Huang
Tetradoxa omeiensis (Hara) C. Y. Wu
APIACEAE
Angelica chinghaiensis Shan ex K. T.
Fu
A. dielsii H. Boissieu
A. laxifoliata Diels
A. maowenensis C. Q. Yuan & R. H.
Shan
A. omeiensis C. Q. Yuan & R. H.
Shan
A. setchuenensis Diels
A. sinensis (Oliv.) Diels
Bupleurum marginatum Wall. ex DC.
var. stenophyllum (Wolff) Shan &
Y. Li
Heracleum candicans Wall.
H. franchetii M. Hiroe
H. hemsleyanum Diels
H. henryi H. Wolff
H. millefolium Diels
H. millefolium Diels var. longilobum
C. Norman
H. obtusifolium Wall.
H. stenopterum Diels
H. vicinum H. Boissieu
H. wolongense F. T. Pu & X. J. He
H. yungningense Hand.-Mazz.
Ligusticum acuminatum Franch.
L. brachylobum Franch.
L. pteridophyllum Franch. ex Gliver
Physospermopsis rubrinervis (Franch.)
C. Norman
Pimpinella flaccida C. B. Clarke
Pleurospermum angelicoides Benth.
ex C. B. Clarke
P. rivulorum (Diels) K. T. Fu & Y. C.
Ho
Tongoloa elata Il. Wolff
Trachyspermum scaberulum (Franch.)
H. Wolff var. ambrosiifolium
(Franch.) R. H. Shan
ARACEAE
Acorus calamus L.
Amorphophallus bannanensis H. Li
Arisaema biauriculatum W. W. Sm.
A. dulongense H. Li
Locality
2n
Ploidy
QH: Yushu
36
2x
X. F. Lu et al. (2002)
SC: Mt. Emei, 2300 m
36
2x
H. X. Liang (1986)
SC: Songpan, 3500 m
44
4x
Z. H. Pan et al. (1991)
SC: Songpan, 3000 m
SC: Hanyuan, 1900 m
SC: Songpan, 2800 m
22
22
22
2x
2x
2x
Z. H. Pan et al. (1991)
Z. H. Pan et al. (1991)
Z. H. Pan et al. (1991)
SC: Mt. Emei, 2100 m
22
2x
Z. H. Pan et al. (1991)
SC: Songpan, 2800 m
SC: Songpan, 2900 m
YN: Heqin
22
22
2x
2x
2x
Z. H. Pan et al. (1991)
Z. H. Pan et al. (1991)
Chin et al. (1989)
22
22
22
2x
2x
2x
4x
2x
2x
X.
X.
X.
X.
X.
X.
2x
2x
2x
2x
2x
2x
2x
2x
2x
X. J. He et al. (1994)
X. J. He et al. (1994)
X. J. He et al. (1994)
X. J. He et al. (1994)
X. J. He et al. (1994)
Z. H. Pan et al. (1985)
Chin et al. (1989)
Chin et al. (1989)
Chin et al. (1989)
22
2x
2x
Chin et al. (1989)
Chin et al. (1989)
22
2x
Chin et al. (1989)
2x
2x
Chin et al. (1989)
Chin et al. (1989)
66
26
6x
2x
H. Wang et al. (2001)
Gu et al. (1992)
28
2x
Gu et al. (1992)
28
26
2x
2x
Gu & Sun (1998)
Gu et al. (1992)
SC: Songpan
SC: Xiaojin
SC: Lixian
YN: Lushui
SC: Songpan
SC: Songpan
SC: Wenchuan
SC: Songpan
SC: Wenchuan
SC: Wenchuan
SC: Xiaojin
SC: Xiaojin
YN: Heqin
YN: Dali
YN: Lijiang
YN: Binchuan
YN: Lijiang
n
7
22
22
22, 24
22
22
22
22
22
22
11
11
11
10
YN: Lijiang
YN: Dali
YN: Binchuan
YN: Zhongdian
YN: Gongshan,
1280 m
YN: Gongshan,
2600 m
XZ: Motuo, 1100 m
YN: Gongshan,
2600 m
11
9
Reference
J.
J.
J.
J.
J.
J.
He
He
He
He
He
He
et
et
et
et
et
et
al.
al.
al.
al.
al.
al.
(1994)
(1994)
(1994)
(1994)
(1994)
(1994)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
289
Continued.
Taxon
2n
Ploidy
56
4x
Gu et al. (1992)
28
2x
SC: Mt. Emei, 1700 m
YN: Tengchong,
3700 m
YN: Gongshan,
2800 m
YN: Dali
YN: Dali
YN: Dali
YN: Baoshan, 2290 m
56
28
4x
2x
28
2x
Murata & Iijima
(1983)
Hong & Zhang (1990)
Z. Y. Yang et al.
(2003)
Gu et al. (1992)
28
42
42
26
2x
3x
3x
2x
Long et al. (1989)
H. Li & Hay (1992)
Long et al. (1989)
Bian et al. (2001)
SC: Wenchuan,
1400 m
SC: Lixian, 3800 m
SC: Kanging, 3100 m
SC: Kangding, 4150 m
QH: Yushu, 3850 m
18, 36
2x, 4x
Kondo et al. (1995)
36
18
36
18
4x
2x
4x
2x
Kondo et
Kondo et
Kondo et
J. Q. Liu
QH: Yushu, 3950 m
QH: Yushu, 3850 m
18
18
2x
2x
J. Q. Liu (1999)
J. Q. Liu (1999)
SC: Hongyuan, 3900 m
58
2x
J. Q. Liu et al. (2001)
QH: Maduo, 4900 m
QH: Maduo, 4800 m
QH: Yushu, 4500 m
QH: Maduo, 4800 m
QH: Maqing, 4300 m
QH: Maduo, 4500 m
QH: Yushu, 4700 m
QH: Maduo, 5000 m
SC: Zhaojue, 1950 m
58
58
58
116
60
58
58
58
54
2x
2x
2x
4x
2x
2x
2x
2x
6x
J. Q. Liu et al. (2001)
Huang et al. (1996a)
J. Q. Liu et al. (2001)
Huang et al. (1996a)
J. Q. Liu et al. (2001)
J. Q. Liu et al. (2001)
J. Q. Liu et al. (2001)
J. Q. Liu et al. (2001)
Kondo et al. (1998)
SC: Mt. Emei, 600 m
36
4x
58
2x
2x
2x
L. dentata (A. Gray) Hara
SC: Mt. Emei, 1500 m
L. dictyoneura (Franch.) Hand.-Mazz. YN: Zhongdian
YN: Lijiang, 3300 m
L. hookeri (C. B. Clarke) Hand.YN: Dali
Mazz.
L. kanaitzensis (Franch.) Hand.-Mazz. YN: Lijiang
L. liatroides (C. Winkler) Hand.QH: Yushu, 3500 m
Mazz.
L. latihastata Hand.-Mazz.
YN: Lijiang
L. pleurocaulis Hand.-Mazz.
SC: Daocheng, 4500 m
58
58
58
58
2x
2x
2x
2x
J. W. Wang et al.
(1993)
Hong & Zhang (1990)
Hong & Zhang (1990)
Y. Z. Pan et al.
(2004a)
J. Q. Liu (2004)
Gong et al. (2001)
J. Q. Liu (2004)
Gong et al. (2001)
58
58
2x
2x
Gong et al. (2001)
J. Q. Liu (2004)
58
58
2x
2x
L. phyllocolea Hand.-Mazz.
58
2x
Gong et al. (2001)
Y. Z. Pan et al.
(2004a)
Y. Z. Pan et al.
(2004a)
A. erubescens Schott
A. heterophyllum Blume
A. lobatum Engl.
Colocasia gaoligongensis H. Li & C.
L. Long
Remusatia hookeriana Schott
R. vivipara Schott
Sauromatum gaoligongense Z. L.
Wang & H. Li
ASTERACEAE
Ajania myriantha (Franch.) Ling ex
C. Shih
A. przewalskii Poljakov
A. ramosa (Chang) C. Shih
A. tenuifolia (Jacq.) Tzvelev
Aster diplostephioides Benth. & Hook.
f.
A. souliei Franch.
A. yunnanensis Franch. var. labrangensis (Hand.-Mazz.) Y. Ling
Cremanthodium brunneopilosum S.
W. Liu
C. discoideum Maxim.
C. ellisii (Hook. f.) S. Kitamura
C. humile Maxim.
C. lineare Maxim.
C. microglossum S. W. Liu
C. stenoglossum Ling & S. W. Liu
Dendranthema glabriusculum (W. W.
Sm.) C. Shih
D. lavandulifolium (Fisch. ex Trautv.)
Y. Ling & C. Shih
Erigeron annuus Pers.
Gnaphalium affine D. Don
Ligularia cyathiceps Hand.-Mazz.
Locality
n
YN: Gongshan,
2300 m
SC: Mt. Emei
SC: Baoxing, 1500 m
SC: Mt. Emei, 800 m
YN: Lijiang, 3600 m
YN: Ninglang, 3700 m
9
7
Reference
al. (1992)
al. (1992)
al. (1992)
(1999)
290
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
L.
L.
L.
L.
purdomii (Turrill) Chittenden
stenocephala Matsum. & Koidzumi
tangutorum Pojark.
tongolensis (Franch.) Hand.-Mazz.
L. tsangchanensis (Franch.) Hand.Mazz.
L. vellerea Hand.-Mazz.
L. virgaurea Mattf. ex Rehder & Kobuski
L. yunnanensis (Franch.) Chang
Nannoglottis gynura (C. Winkler)
Ling & Y. L. Chen
Nouelia insignis Franch.
Locality
2n
Ploidy
SC: Hongyuan
YN: Dali
QH: Maqin, 3200 m
YN: Zhongdian,
3200 m
YN: Dali, 3000 m
58
58
58
58
2x
2x
2x
2x
58
2x
J. Q. Liu (2004)
Gong et al. (2001)
J. Q. Liu (2004)
Y. Z. Pan et al.
(2004a)
J. Q. Liu (2004)
YN: Lijiang
SC: Ruoergai
58
58
2x
2x
Gong et al. (2001)
J. Q. Liu (2004)
QH: Maqing, 3700 m
YN: Dali, 3600 m
87
58
3x
2x
QH: Yushu, 3600 m
18
2x
J. Q. Liu (2004)
Y. Z. Pan et al.
(2004a)
J. Q. Liu et al. (2000a)
54
6x
Peng et al. (2002)
2x
Hong & Zhang (1990)
30
2x
Ho et al. (2002c)
30
48
2x
Ho et al. (2002c)
Huang et al. (1996a)
28
2x
Huang et al. (1996a)
28
2x
Huang et al. (1996a)
9
8
2x
2x
Hong & Zhang (1990)
Hong & Zhang (1990)
6
2x
Hong & Zhang (1990)
YN: Zhongdian,
1988 m
Picris hieracioides L. subsp. japonica SC: Baoxing, 1800 m
Krylv.
Saussurea ceterach Hand.-Mazz.
QH: Nangqian,
3940 m
S. eopygmaea Hand.-Mazz.
QH: Yushu, 4060 m
S. gnaphalodes (Royle) Sch. Bip.
QH: BayanHarashan,
5200 m
S. tangutica Maxim.
QH: BayanHarashan,
5200 m
S. wellbyi Hemsl.
QH: Maduo, 4700–
4800 m
Sonchus arvensis L.
SC: Mt. Emei, 800 m
Youngia erythrocarpa (Vaniot) Babc. SC: Baoxing, 1500 m
& Stebbins
Y. heterophylla (Hemsl.) Babc. &
SC: Mt. Emei, 1200 m
Stebbins
BEGONIACEAE
Begonia cavaleriei H. Lév.
n
5
Reference
YN: Eryuan
30
3x
Tian et al. (2002)
YN: Zhongdian,
3200 m
SC: Mt. Emei, 805 m
12
2x
Ma & Hu (1996)
12
2x
Ma & Hu (1996)
YN: Zhongdian,
3200 m
12
2x
Ma & Hu (1996)
BIGNONIACEAE
Incarvillea arguta Royle
YN: Zhongdian
22
2x
I. berezovskii Batalin
SC: Maerkang
22
2x
I. compacta Maxim.
SC: Shiqu
22
2x
I. delavayi Bureau & Franch.
YN: Lijiang
22
2x
I. dissectifolia Q. S. Zhao
SC: Yanyuan
22
2x
I. forrestii Fletcher
YN: Zhongdian
22
2x
S. T. Chen
(2004)
S. T. Chen
(2004)
S. T. Chen
(2004)
S. T. Chen
(2004)
S. T. Chen
(2004)
H. Xiao et
BERBERIDACEAE
Diphylleia sinensis Li
Dysosma versipellis (Hance) M.
Cheng
Sinopodophyllum hexandrum (Royle)
Ying
et al.
et al.
et al.
et al.
et al.
al. (2002)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
291
Continued.
Taxon
Locality
n
2n
Ploidy
Reference
I. lutea Bureau & Franch.
SC: Daocheng
22
2x
I. mairei (H. Lév.) Grierson
YN: Lijiang
YN: Zhongdian
YN: Dali
22
22
22
2x
2x
2x
I. mairei (H. Lév.) Grierson var.
grandiflora (Wehrhahn) Grierson
I. sinensis Lam.
YN: Lijiang
22
2x
SC: Maerkang
22
2x
I. younghusbandii Sprague
XZ: Nielamu
22
2x
I. zhongdianensis Grey-Wilson
YN: Lijiang
YN: Zhongdian
22
22
2x
2x
YN: Deqin, 4600 m
14
2x
Yue et al. (2003)
XZ: Basho, Yelashan
XZ: Lhasa, Resela
YN: Deqin, 4310 m
YN: Zhongdian,
4500 m
XZ: Nyenchengtang La
14
14
14
14
2x
2x
2x
2x
Yue
Yue
Yue
Yue
14
14
28
2x
2x
4x
Yue et al. (2003)
Yue et al. (2004)
Yue et al. (2003)
S. xerophyta (W. W. Sm.) Comber
Thlaspi arvense L.
YN: Lijiang, 4500 m
YN: Lijiang
SC: Xiangcheng,
4000 m
XZ: Zuogong
SC: Daocheng
SC: Baoxing, 2600 m
14
14
2x
2x
2x
Yue et al. (2004)
Yue et al. (2004)
Hong & Zhang (1990)
CAMPANULACEAE
Adenophora lobophylla D. Y. Hong
SC: Jingchuan
3412B
2x
A. potanini Korsh.
SC: Baoxin
34
2x
A. stricta Miq.
SC: Jingchuan
68
4x
K. Q. Wang & Ge
(1998)
K. Q. Wang & Ge
(1998)
K. Q. Wang & Ge
(1998)
CAPRIFOLIACEAE
Lonicera schneideriana Rehder
SC: Baoxing, 2600 m
BRASSICACEAE
Solms-laubachia eurycarpa (Maxim.)
Botsch.
S. lanata Botsch.
S. linearifolia O. E. Schulz
S. minor Hand.-Mazz.
S. platycarpa (Hook. f. & Thomson)
Botsch.
S. pulcherrima Muschl.
S. retropilosa Botsch.
CARYOPHYLLACEAE
Psammosilene tunicoides W. C. Wu & YN: Lijiang
C. Y. Wu
Stellaria neglecta Weihe
SC: Mt. Emei, 2100 m
CHLORANTHACEAE
Chloranthus nervosus Collett &
Hemsl.
CUCURBITACEAE
Gynostemma longipes C. Y. Wu
Thladiantha davidii Franch.
T. dentata Cogn.
S. T. Chen
(2004)
H. Xiao et
H. Xiao et
S. T. Chen
(2004)
S. T. Chen
(2004)
S. T. Chen
(2004)
S. T. Chen
(2004)
H. Xiao et
S. T. Chen
(2004)
et
et
et
et
al.
al.
al.
al.
et al.
al. (2002)
al. (2002)
et al.
et al.
et al.
et al.
al. (2002)
et al.
(2004)
(2004)
(2003)
(2003)
Yue et al. (2004)
7
9
28
22
2x
Hong & Zhang (1990)
2x
4x
Y. Z. Pan et al.
(2004b)
Hong & Zhang (1990)
YN: Yangbi, Meixi
30
2x
Kong (2000)
YN: Lijiang
SC: Pengxian, 1230 m
SC: Nanchuan, 1650 m
44
18
18
4x
2x
2x
X. F. Gao et al. (1995)
J. Q. Li et al. (1993)
J. Q. Li et al. (1993)
292
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
T. lijiangensis A. M. Lu & Zhi Y.
Zhang
T. nudiflora Hemsl. ex Forb. &
Hemsl.
Locality
2n
Ploidy
YN: Lijiang, Longpan
18
2x
J. Q. Li et al. (1993)
SC: Mt. Emei, 1650 m
18
2x
J. Q. Li et al. (1993)
Jiangsu Institute of
Botany (1976)
Jiangsu Institute of
Botany (1976)
Jiangsu Institute of
Botany (1976)
Chin et al. (1985)
Jiangsu Institute of
Botany (1976)
Pei et al. (1979)
Jiangsu Institute of
Botany (1976)
n
Reference
DIOSCOREACEAE
Dioscorea althaeoides Knuth
YN: Weixi
10
20
2x
D. collettii Hook. f.
SC: Emei
10
20
2x
D. deltoidea Wall.
YN: Deqin
10
20
2x
D. opposita Thunb.
D. panthaica Prain & Burkill
SC: Mt. Emei
YN: Lijiang
20
138–142
40
14x
4x
D. parviflora C. T. Ting
D. zingiberensis C. H. Wright
YN: Yongsheng
YN: Liuku
10
10
20
20
2x
2x
SC: Daocheng
24
2x
Y. L. Cao & Lu (1989)
SC: Nanping
24
2x
Y. L. Cao & Lu (1989)
YN: Zhongdian
24
2x
Y. L. Cao & Lu (1989)
SC: Hongyuan
24
2x
Y. L. Cao & Lu (1989)
ERICACEAE
Rhododendron mackenzianum Forrest YN: Gongshan
R. leptothrium Balf. f. & Forrest
YN: Weixi
26
26
2x
2x
L. M. Gao (2002)
L. M. Gao (2002)
FABACEAE
Astragalus acaulis Baker
SC
16
2x
A. ernestii Comber
SC: Kangding
16
2x
A. kialensis N. D. Simpson
YN
16
2x
A. pastorius Tsai & Yu
SC
32
4x
A. przewalskii Bunge ex Maxim.
SC
16
2x
A. strictus Graham
XZ: Bomi, 3300 m
32
4x
A. tongolensis Ulbrich
SC
16
2x
Caragana bicolor Kom.
SC
16
2x
Crotalaria medicaginea Lam.
C. sessiliflora L.
Desmodium yunnanense Franch.
Gueldenstaedtia delavayi Franch.
Mastersia assamica Benth.
Medicago lupulina L.
Salweenia wardii Baker f.
YN: Heqin
XZ: Motuo, 900 m
YN: Heqing
YN: Lijiang
XZ: Motuo, 900 m
SC: Baoxing
SC: Daocheng
16
16
22
16
22
16
2x
2x
2x
2x
2x
2x
2x
Tibetia coelestis (Diels) H. P. Tsui
YN: Zhongdian
16
2x
S. M. Mu & Shue
(1985)
S. M. Mu & Shue
(1985)
S. M. Mu & Shue
(1985)
S. M. Mu & Shue
(1985)
S. M. Mu & Shue
(1985)
W. T. Wang et al.
(1994)
S. M. Mu & Shue
(1985)
S. M. Mu & Shue
(1985)
J. Q. Li (1988)
Gu & Sun (1998)
Cui (1987)
Nie (2002)
Gu & Sun (1998)
Hong (1984)
Q. X. Zhou et al.
(2000b)
Nie (2002)
ELAEAGNACEAE
Hippophae neurocarpa S. W. Liu & T.
N. He
H. rhamnoides L. subsp. sinensis
Rousi
H. rhamnoides L. subsp. yunnanensis
Rousi
H. tibetana Schlecht.
8
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
Continued.
Taxon
T. himalaica (Baker) H. P. Tsui
Locality
2n
Ploidy
16
16
16
16
12
2x
2x
2x
2x
2x
2x
2x
Nie et al. (2002)
Nie (2002)
Nie et al. (2002)
Nie et al. (2002)
Hong (1984)
Hong (1984)
Hong (1984)
24
2x
M. Cao & Zhou (2000)
2x
Hong & Zhang (1990)
18
18
20
18
18
2x
2x
2x
2x
2x
J.
J.
J.
J.
J.
20
24
48
24
2x
2x
4x
2x
Küpfer & Yuan (1996)
J. Q. Liu et al. (2002a)
Chen et al. (1997)
J. Q. Liu et al. (2002a)
14
14
20
16
26
24
24
26
2x
2x
2x
2x
2x
2x
2x
2x
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
Küpfer & Yuan (1996)
Ho et al. (2002b)
Yuan & Küpfer (1997)
J. Q. Liu et al. (2002a)
Ho et al. (2002a)
Yuan et al. (1998)
20
40
18
2x
4x
2x
Yuan et al. (1998)
Yuan et al. (1998)
Yuan & Küpfer (1997)
YN: Lijiang, 2900 m
YN: Lijiang, 2400 m
YN: Zhongdian,
3400 m
XZ: Dingqing, 4250 m
SC: Litang, 3800 m
26
20
20
2x
2x
2x
Yuan & Küpfer (1997)
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
20
18
2x
2x
GS: Maqu, 3650 m
GS: Maqu, 3650 m
GS: Maqu, 3900 m
YN: Dali, 2100 m
QH: Maduo, 4300 m
YN: Lijiang, 3200 m
YN: Lijiang, 2700 m
SC: Ruoergai, 4200 m
XZ: Dingqing, 4200 m
YN: Lijiang, 3200 m
12
12
36
20
12
20
20
36
26
20
2x
2x
6x
2x
2x
2x
2x
4x
2x
2x
Yuan et al. (1998)
S. L. Chen et al.
(1997)
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
Yuan et al. (1998)
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
Yuan et al. (1998)
Yuan et al. (1998)
Yuan et al. (1998)
T. tongolensis (Ulbr.) H. P. Tsui
T. yunnanensis (Franch.) H. P. Tsui
Vicia hirsuta (L.) Gray
V. sativa L.
V. villosa Roth
SC: Daocheng
SC: Litang
SC: Daocheng
YN: Zhongdian
SC: Baoxing
SC: Baoxing
SC: Baoxing
FAGACEAE
Quercus guyavaefolia H. Lév.
YN: Lijiang
FUMARIACEAE
Corydalis omeiana H. Tsaung
SC: Mt. Emei, 1700 m
GENTIANACEAE
Comastoma arrectum (Franch.) Holub
C. beesianum (W. W. Sm.) Holub
C. chiuchiense T. N. Ho & J. Q. Liu
C. pulmonarium (Turcz.) Toyokuni
C. traillianum (Forrest) Holub
Gentiana alsinoides Franch.
G. altorum Harry Sm.
G. ampla Harry Sm.
G. arethusae Burkill subsp. delicatula (C. Marquand) Halda
G. aristata Maxim.
G.
G.
G.
G.
G.
G.
293
asterocalyx Diels
caeruleogrisea T. N. Ho
callistantha Diels & Gilg
cephalantha Franch.
chinensis Kusnezow
crassicaulis Duthie ex Burkill
G. crassula Harry Sm.
G. crassuloides Bureau & Franch.
G. crenulatotruncata (C. Marquand)
T. N. Ho
G. delavayi Franch.
G. exigua Harry Sm.
G. haynaldii Kanitz
G. heleonastes Harry Sm.
G. heterostemon Harry Sm.
G. hyalina T. N. Ho
G. intricata C. Marquand
G. leucomelaena Maxim.
G. lhassica Burkill
G. mairei H. Lév.
n
7
6
8
YN: Deqin, 3900 m
YN: Lijiang, 2800 m
QH: Maqin, 4450 m
QH: Maqin, 4000 m
YN: Zhongdian,
3382 m
YN: Lijiang, 2950 m
SC: Kangding, 3600 m
YN: Lijiang, 3700 m
YN: Deqin, 3800 m
GS: Luqu, 3450 m
GS: Maqu, 3500 m
YN: Lijiang, 2900 m
QH: Maqin, 3500 m
GS: Luqu, 3500 m
YN: Lijiang, 3000 m
SC: Mt. Emei, 3000 m
YN: Zhongdian,
3300 m
YN: Lijiang, 4300 m
YN: Deqin, 4100 m
GS: Maqu, 4200 m
7
Reference
Q.
Q.
Q.
Q.
Q.
Liu
Liu
Liu
Liu
Liu
&
&
&
&
&
Ho
Ho
Ho
Ho
Ho
(2002)
(2002)
(2002)
(2002)
(2002)
294
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
2n
Ploidy
Reference
YN: Lijiang, 3700 m
24
2x
G.
G.
G.
G.
G.
G.
YN: Lijiang, 2900 m
QH: Maqin, 4900 m
YN: Dali, 3200 m
YN: Dali, 2200 m
SC: Sungpan, 3700 m
GS: Luqu, 3500 m
GS: Luqu, 3300 m
GS: Maqu, 3800 m
XZ: Dingqing, 4250 m
SC: Sungpan, 3400 m
GS: Maqu, 3700 m
YN: Lijiang, 2400 m
YN: Lijiang, 2400 m
GS: Luqu, 3700 m
SC: Litang, 3800 m
24
24
20
20
20
18
20
20
20
20
20
34
46
38
26
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
52
20
20
4x
2x
2x
G. tibetica King ex Hook. f.
G. veitchiorum Hemsl.
GS: Maqu, 3500 m
YN: Lijiang, 2900 m
YN: Zhongdian,
3200 m
XZ: Changdu, 3650 m
SC: Litang, 3800 m
S. L. Chen et al.
(1997)
Ho et al. (2002a)
Ho et al. (2002b)
Yuan & Küpfer (1997)
Küpfer & Yuan (1996)
Yuan et al. (1998)
Küpfer & Yuan (1996)
Küpfer & Yuan (1996)
Yuan & Küpfer (1997)
Yuan et al. (1998)
Yuan et al. (1998)
Yuan & Küpfer (1997)
Ho et al. (2002b)
Ho et al. (2002b)
Yuan et al. (1998)
S. L. Chen et al.
(1997)
Yuan & Küpfer (1997)
Küpfer & Yuan (1996)
Yuan et al. (1998)
52
24
4x
2x
XZ: Quxu, 3700 m
SC: Baoxing, 3400 m
YN: Dali, 3700 m
YN: Lijiang, 2900 m
26
24
24
22
2x
2x
2x
2x
QH: Dari, 4200 m
YN: Lijiang
YN: Lijiang
YN: Lijiang
QH: Dari
22
26
52
26
12
2x
2x
4x
2x
2x
YN: Deqin
28
4x
XZ: Changdu, 3800 m
QH: Maqin, 3800 m
QH: Maduo, 3700 m
20
14
28
2x
2x
4x
J. Q. Liu et al. (2002c)
Yuan & Küpfer (1993)
Yuan & Küpfer (1993)
Yuan & Küpfer (1993)
J. Q. Liu et al.
(2002b)
J. Q. Liu et al.
(2002b)
T. N. He et al. (1999)
T. N. He et al. (1999)
T. N. He et al. (1999)
2x
Hong & Zhang (1990)
34
2x
40
4x
Y. Z. Wang & Gu
(1999)
Y. X. Lu et al. (2002)
34
2x
P. Zhou (2003)
32
2x
Y. L. Mu et al. (1990)
G. microdonta Franch.
G.
G.
G.
G.
nubigena Edgew.
panthaica Burkill
praticola Franch.
prattii Kusnezow
G. pseudoaquatica Kusnezow
pseudosquarrosa Harry Sm.
pudica Maxim.
serra Franch.
souliei Franch.
squarrosa Ledeb.
stipitata Edgew. subsp. tizuensis
(Franch.) T. N. Ho
G. straminea Maxim.
G. subintricata T. N. Ho
G. waltonii Burkill
G. wasenensis C. Marquand
G. yunnanensis Franch.
Gentianella gentianoides (Franch.)
Harry Sm.
G. azurea (Bunge) Holub
Gentianopsis barbata (Froel.) Ma
G. grandis (Harry Sm.) Ma
G. lutea Ma
Lomatogoniopsis alpina T. N. Ho &
S. W. Liu
Megacodon stylophorus (C. B. Clarke)
Harry Sm.
Swertia franchetiana Harry Sm.
S. tetraptera Maxim.
S. wolfgangiana Gruning
GERANIACEAE
Geranium eriostemon Fisch. ex DC.
Locality
SC: Baoxing, 2500 m
GESNERIACEAE
Ancylostemon aureus (Franch.) B. L. YN: Binchuan
Burtt
Corallodiscus flabellatus (Craib) B. L. YN: Zhongdian
Burtt
Loxostigma griffithii (Wight) C. B.
YN: Rujiang River
Clarke
JUGLANDACEAE
Juglans sigillata Dode
LAURACEAE
YN: Yangbi
n
14
Yuan et al. (1998)
S. L. Chen et al.
(1997)
Ho et al. (2002a)
Ho et al. (2002b)
Yuan et al. (1998)
J. Q. Liu et al. (2002c)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
295
Continued.
Taxon
Locality
n
2n
Ploidy
Reference
Cinnamomum longipaniculatum
(Gamble) N. Chao ex H. W. Li
Lindera communis Hemsl.
SC: Dujiangyan
24
2x
SC: Dujiangyan
24
2x
L. megaphylla Hemsl.
SC: Dujiangyan
24
2x
L. pulcherrima (Nees) Benth. ex
Hook. f.
SC: Dujiangyan
24
2x
QH: Maduo, 4450 m
YN: Jianchuan,
2800 m
YN: Zhongdian,
3250 m
XZ: Zhayab, 3780 m
SC: Xiangcheng,
2950 m
SC: Xiangcheng,
2800 m
YN: Zhongdian,
3200 m
YN: Dali, 2050 m
YN: Dali, 2050 m
YN: Zhongdian,
3200 m
YN: Zhongdian,
3250 m
YN: Zhongdian,
3250 m
YN: Zhongdian,
3250 m
SC: Wenchuan,
1200 m
SC: Daocheng, 4500 m
SC: Emei, 1400 m
SC: Wenchuan,
2200 m
YN: Zhongdian,
3600 m
SC: Lixian, 3600 m
SC: Litang, 4100 m
QH: Maqin, 3400 m
QH: Nangqian,
3550 m
XZ: Zhayab, 3780 m
YN: Zhongdian,
3300 m
YN: Baoshan, 1500 m
SC: Lixian, 1100 m
YN: Binchuan, 2900 m
YN: Binchuan 3200 m
YN: Zhongdian,
3300 m
SC: Mt. Emei
YN: Gongshan,
2700 m
16
22
2x
2x
16
2x
16
20
2x
2x
L. Yang et al. (1998)
Huang et al. (1995)
20
2x
Xu et al. (1998)
22
2x
Huang et al. (1995)
33
44
22
3x
4x
2x
Huang et al. (1995)
Huang et al. (1995)
Huang et al. (1996b)
28
4x
Huang et al. (1996b)
14
2x
Huang et al. (1995)
28
4x
Huang et al. (1995)
32
4x
16
22
24
2x
2x
3x
S. M. Zhu & Xu
(1999)
Jing et al. (1999)
Huang et al. (1995)
Jing et al. (1999)
16
2x
Jing et al. (1999)
32
16
32
64
4x
2x
4x
6x
Jing
Jing
Xue
Xue
32
16
4x
2x
L. Yang et al. (1998)
Huang et al. (1995)
24
32
14
14
14
3x
4x
2x
2x
2x
Huang et al. (1985)
L. Yang et al. (1998)
Huang et al. (1995)
Huang et al. (1996c)
Huang et al. (1995)
24
24
2x
2x
Tang et al. (1984)
Xie et al. (1992)
LILIACEAE
Allium carolinianum Delar.
A. chienchuanense J. M. Xu
A. cyathophorum E. Bureau &
Franch.
A. fasciculatum Rendle
A. hookeri Thwaites
A. hookeri Thwaites var. muliense
Airy Shaw
A. macranthum Baker
A. macrostemon Bunge
A. nanodes Airy Shaw
A. omeiense Z. Y. Zhu
A. ovalifolium Hand.-Mazz.
A. prattii C. H. Wright
A. przewalskianum Regel
A. trifurcatum (Wang & Tang) J. M.
Xu
A. tuberosum Rottl. ex Spreng.
A. wallichii Kunth
A. wallichii Kunth var. platyphyllum
(Diels) J. M. Xu
Cardiocrinum giganteum Makino
C. B. Chen
(1998)
C. B. Chen
(1998)
C. B. Chen
(1998)
C. B. Chen
(1998)
et al.
et al.
et al.
et al.
Huang et al. (1996a)
Huang et al. (1995)
et
et
et
et
al.
al.
al.
al.
(1999)
(1999)
(2000)
(2000)
296
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
Clintonia udensis Trautv. & Mey.
Disporopsis aspersa (Hua) Engl.
D. fusco-picta Hance
Disporum bodinieri (H. Lév. & Vaniot) F. T. Wang & T. Tang
D. cantoniense Merr.
Fritillaria cirrhosa D. Don
Hemerocallis forrestii Diels
H. multiflora Stout
H. plicata Stapf
Lilium bakerianum Coll. & Hemsl.
var. rubrum Stearn
L. brownii F. E. Brown ex Spae
L.davidii Duch.
Locality
2n
Ploidy
SC: Mt. Emei, 2750 m
14
2x
YN: Lijiang, 3700 m
28
4x
YN: Yunlong, 3300 m
SC: Mt. Emei
YN: Tengchong
XZ: Motuo, 1500 m
14
40
40
16
2x
2x
2x
2x
S. F. Li & Chang
(1996)
S. F. Li & Chang
(1996)
L. Wang et al. (1993)
Gu et al. (1990)
Gu et al. (1990)
Gu & Sun (1998)
14
24
2x
2x
2x
Gu et al. (1993a)
Tang et al. (1984)
Xie et al. (1992)
22
22
33
22
22
24
2x
2x
3x
2x
2x
2x
Xiong & Chen (1998)
Xiong et al. (1997)
Xiong & Chen (1998)
Xiong & Chen (1998)
Xiong et al. (1997)
L. Wang et al. (1993)
24
36
24
24
2x
3x
2x
2x
Xie et al. (1992)
Xie & Wu (1993)
Yu et al. (1996a)
Huang & Li (1996)
24
24
24
2x
2x
2x
L. Wang et al. (1993)
Xie et al. (1992)
Xie et al. (1992)
24
2x
Xie et al. (1992)
24
2x
Xie et al. (1992)
24
2x
Xie et al. (1992)
24
24
24
24
2x
2x
2x
2x
L. Wang et al. (1993)
Xie et al. (1992)
Yu & Huang (1994)
Yu et al. (1996d)
24
24
2x
2x
Yu et al. (1996b)
Xie et al. (1992)
24
24
2x
2x
L. Wang et al. (1993)
Xie et al. (1992)
24
2x
Xie et al. (1992)
24
24
2x
2x
L. Wang et al. (1993)
Yu et al. (1996c)
24
24
72
2x
2x
4x
L. Wang et al. (1993)
Yu et al. (1996c)
D. M. Zhang (1998)
108
6x
D. M. Zhang (1998)
YN: Lijiang, 2600 m
SC: Mt. Emei
YN: Gongshan,
4100 m
YN: Lijiang, 3200 m
YN: Lijiang
YN: Dali, 2500 m
YN: Dali, 2300 m
YN: Dali, 2300 m
YN: Dali, 3200 m
YN: Lushui, 2000 m
YN: Gongshan
YN: Dali, 2400 m
L. duchartrei Franch.
YN: Zhongdian,
3200 m
L. henrici Franch.
YN: Lijiang, 2800 m
L. nepalense D. Don
YN: Lushui, 2700 m
L. souliei (Franch.) Sealy
YN: Gongshan,
3500 m
L. taliense Franch.
YN: Gongshan,
2000 m
Nomocharis aperta W. W. Sm. & W. YN: Gongshan,
E. Evans
3300 m
N. basilissa Farrer ex W. E. Evans
YN: Gongshan,
3400 m
N. biluoensis S. Y. Liang
YN: Lijiang, 3000 m
N. farreri Cox
YN: Lushui, 3100 m
N. forrestii Balf. f.
YN: Zhongdian
YN: Zhongdian,
3800 m
N. mairei H. Lév.
YN: Dali, 3500 m
N. meleagrina Franch.
YN: Gongshan,
3300 m
N. pardanthina Franch.
YN: Dali, 3200 m
YN: Gongshan,
3000 m
N. pardanthina Franch. f. punctulata YN: Gongshan,
Sealy
3300 m
N. saluenensis Balf. f.
YN: Yunlong, 3100 m
Notholirion bulbiliferum (Lingelsh.)
YN: Zhongdian,
Stearn
3400 m
N. campanulatum Cotton & Stearn
YN: Dali, 3200 m
YN: Ninglang, 3500 m
Ophiopogon angustifoliatus (Wang & YN: Weixi, 3100 m
Tang) S. C. Chen
O. bodinieri H. Lév.
YN: Zhongdian,
3600 m
n
8
Reference
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
297
Continued.
Taxon
O. clarkei Hook. f.
O. intermedius D. Don
O. japonicus Ker Gawl.
Paris bashanensis F. T. Wang & T.
Tang
P. dulongensis H. Li & S. Kurita
P. fargesii Franch.
P. forrestii (Takht.) H. Li
P. mairei H. Lév.
P. marmorata Stearn
P. polyphylla Sm.
Locality
2n
Ploidy
36
72
108
10
2x
4x
6x
2x
D. M. Zhang (1998)
D. M. Zhang (1998)
D. M. Zhang (1998)
Gu & Na (1986)
10
2x
Gu et al. (1992)
10
10
2x
2x
2x
Tang et al. (1984)
Miyamoto et al. (1992)
Gu et al. (1992)
10
10
10
2x
2x
2x
Gu & Na (1986)
Miyamoto et al. (1992)
Gu et al. (1992)
10
10
10
20
2x
2x
2x
4x
10
2x
Gu & Na (1986)
Gu & Na (1986)
Hong & Zhu (1987)
S. F. Wang & Xu
(1989)
Gu et al. (1992)
10
10
10
2x
2x
2x
Miyamoto et al. (1992)
Gu & Na (1986)
Gu et al. (1992)
32
2x
S. F. Chen (1989)
32
2x
L. Wang et al. (1993)
52
56
30
78
36
22
30
28
24
38
16
4x
4x
2x
6x
4x
2x
2x
2x
2x
2x
2x
YN: Yunlong, 2800 m
36
2x
Gu et al. (1993b)
S. F. Chen (1989)
L. Wang et al. (1993)
L. Wang et al. (1993)
Gu & Sun (1998)
Fang (1989)
Gu & Sun (1998)
J. Yang et al. (1992)
S. F. Chen (1989)
Tang et al. (1984)
K. Y. Ding et al.
(1998)
L. Wang et al. (1993)
YN: Zhongdian,
3800 m
YN: Lushui, 3100 m
XZ: Motuo, 1600 m
YN: Yunlong, 2800 m
YN: Gongshan,
1300 m
YN: Dali, 3200 m
YN: Zhongdian,
3700 m
YN: Gongshan,
3300 m
YN: Dali, 3200 m
72
4x
Meng et al. (2005)
36
54
36
72
2x
3x
2x
4x
Meng et al. (2005)
Gu & Sun (1998)
L. Wang et al. (1993)
Gu et al. (1992)
36
36
2x
2x
L. Wang et al. (1993)
Meng et al. (2005)
36
2x
Meng et al. (2005)
36
2x
L. Wang et al. (1993)
YN:
YN:
YN:
YN:
Lijiang, 2750 m
Lijiang, 2750 m
Weixi, 2500 m
Dali
YN: Gongshan,
1320 m
SC: Mt. Emei
YN: Tengchong
YN: Gongshan,
2350 m
YN: Dali
YN: Dali
YN: Gongshan,
1700 m
YN: Dali
YN: Dali
SC: Wenchuan
SC: Mt. Emei
P. rugosa H. Li & S. Kurita
YN: Gongshan,
1500 m
P. thibetica Franch.
YN: Dali
YN: Dali
P. thibetica Franch. var. apetala
YN: Gongshan,
Hand.-Mazz.
2300 m
Polygonatum alternicirrhosum Hand.- SC: Luding, 2800 m
Mazz.
P. cathcartii Baker
YN: Gongshan,
2600 m
P. cirrhifolium Royle
XZ: Bomi, 3000 m
SC: Kangding, 3300 m
YN: Yunlong, 2900 m
P. curvistylum Hua
YN: Dali, 3000 m
P. griffithii Baker
XZ: Motuo, 1900 m
P. odoratum Druce
SC: Mt. Emei, 800 m
P. oppositifolium Royle
XZ: Motuo, 2100 m
P. prattii Baker
YN: Dali
P. sibiricum Redouté
SC: Nanping, 3200 m
Reineckea carnea (Andr.) Kunth
SC: Mt. Emei
Scilla sinensis (Lour.) Merr.
YN: Eryuan, Lijiang
Smilacina atropurpurea (Franch.)
Wang & Tang
S. forrestii (W. W. Sm.) Hand.-Mazz.
S. fusca Wall.
S. henryi (Baker) Hara
S. lichiangense (W. W. Sm.) W. W.
Sm.
S. purpurea Wall.
S. tatsienensis (Franch.) Wehrh.
n
5
Reference
298
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
2n
Ploidy
XZ: Motuo, 1700 m
SC: Mt. Emei
SC: Mt. Emei
32
10
20
4x
2x
4x
Gu & Sun (1998)
S. F. Wang (1989)
S. F. Wang & Xu
(1989)
ONAGRACEAE
Circaea alpina L. subsp. angustifolia YN: Hengduan Mts.
(Hand.-Mazz.) D. E. Boufford
22
2x
Seavey & Boufford
(1983)
Streptopus simplex D. Don
Trillium tschonoskii Maxim.
Locality
n
Reference
ORCHIDACEAE
Cymbidium hookerianum Rchb. f.
C. eburneum Lindl.
YN: Tengchong
YN: Baoshan
40
40
PAEONIACEAE
Paeonia delavayi Franch.
YN: Lijiang, 2400 m
10
2x
YN: Dali, 2910 m
10
2x
D. Q. Yang & Zhu
(1989)
Gong et al. (1999)
YN: Zhongdian,
3200 m
YN: Dali, 3000 m
10
2x
Gong et al. (1991)
10
2x
SC: Wolong Nat. Res.
20
4x
D. Q. Yang & Zhu
(1989)
Hong et al. (2001)
YN: Zhangdian,
4150 m
YN: Weixi, 4100 m
YN: Zhongdian,
4100 m
YN: Zhongdian,
4400 m
76
4x
Meng et al. (in prep.)
56
56
4x
4x
Meng et al. (in prep.)
Meng et al. (in prep.)
56
4x
Meng et al. (in prep.)
4x
Hong & Zhang (1990)
P. delavayi Franch. f. lutea (Delavay
ex Franch.) S. G. Haw
P. obovata Maxim.
PAPAVERACEAE
Meconopsis integrifolia Franch.
M. horridula Hook. f. & Thomson
M. racemosa Maxim.
M. pseudovenusta G. Taylor
PLANTAGINACEAE
Plantago erosa Wall.
POACEAE
Elymus anthosachnoides (Keng) Á.
Löve
E. atratus (Nevski) Hand.-Mazz.
E. cylindricus Honda
E. dahuricus Turcz.
E. nutans Griseb.
E. sibiricus L.
E. submuticus Keng ex Keng f.
SC: Baoxing, 1850 m
12
Y. G. Li et al. (2003)
Y. G. Li et al. (2003)
SC: Batang
28
4x
W. J. Li et al. (1996)
SC:
SC:
SC:
SC:
SC:
SC:
SC:
SC:
SC:
SC
42
28
42
42
42
42
28
28
42
42
6x
4x
6x
6x
6x
6x
4x
4x
6x
6x
42
42
42
6x
6x
6x
42
ca. 21
6x
3x
Y. H. Liu (1985)
B. R. Lu et al. (1990)
Y. H. Liu (1985)
B. R. Lu et al. (1990)
Y. H. Liu (1985)
Y. H. Liu (1985)
Y. H. Liu (1985)
B. R. Lu et al. (1990)
B. R. Lu et al. (1990)
L. B. Cai & Feng
(1997)
Y. H. Liu (1985)
B. R. Lu et al. (1990)
X. Q. Zhang et al.
(1998)
Y. H. Zhou (1994)
Huang et al. (1996a)
28
4x
Hongyuan
Rourgai
Ganzi
Hongyuan
Kangding
Ruoergai
Kangding
Rourgai
Nanping
E. tangutorum (Nevski) Hand.-Mazz.
SC: Shiqu
SC: Maowen
Kengyilia laxiflora (Keng) S. L. Chen SC: Ganzi
K. melanthera (Keng) S. L. Chen
SC: Ruoergai
Poa attenuata Trin. ex Bunge var. vi- QH: BayanHarashan,
vipara Rendl
5200 m
Roegneria altissima Keng ex Y. L.
SC: Batang
Keng & S. L. Chen
Y. H. Zhou et al.
(1993)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
299
Continued.
Taxon
2n
Ploidy
Reference
SC: Ruoergai
28
4x
B. R. Lu et al. (1990)
SC: Wenchuan
SC: Yaan
28
28
4x
4x
SC: Nanping
SC: Wenchuan
14
28
2x
4x
XZ: Chuangdu
28
4x
R. gmelini (Griseb.) Kitag.
R. kamoji (Ohwi) Ohwi
SC: Ruoergai
SC: Lixian
SC: Yaan
28
42
42
4x
6x
6x
R. nutans (Keng) Keng ex Y. L.
Keng & S. L. Chen
SC: Ruoergai
42
6x
B. R. Lu et al. (1990)
G. L. Sun et al.
(1993b)
G. L. Sun et al. (1992)
Y. H. Zhou et al.
(1993)
C. B. Ding et al.
(2004)
B. R. Lu et al. (1990)
B. R. Lu et al. (1990)
X. Q. Zhang et al.
(1998)
B. R. Lu et al. (1990)
SC: Hongyuan
28
4x
SC: Hongyuan
SC: Maerkang
42
28
6x
4x
G. L. Sun et al.
(1993a)
B. R. Lu et al. (1990)
B. R. Lu et al. (1990)
SC:
SC:
SC:
SC:
Lixian
Lixian
Nanping
Aba
42
28
28
28
6x
4x
4x
4x
B.
B.
B.
B.
SC: Maowen
28
4x
B. R. Lu et al. (1990)
YN: Zhongdian,
3400 m
YN: Lijiang, 3200 m
QH: Maduo, 4600 m
QH: Maqin
YN: Jianchuan,
3100 m
YN: Lijiang, 2800 m
YN: Lijiang, 2500 m
YN: Lijiang, 2800 m
YN: Lijiang, 2700 m
YN: Zhongdian,
3200 m
SC: Wenchuan,
3700 m
YN: Zhongdian,
3300 m
YN: Zhongdian,
3300 m
YN: Lijiang, 2800 m
SC: Kangdian, 2500 m
YN: Deqin, 4300 m
YN: Zhongdian,
3100 m
YN: Deqin, 4300 m
YN: Luquan, 2500 m
YN: Zhongdian,
3100 m
20
2x
Nakata et al. (1997)
20
40
20
22
2x
4x
2x
2x
Nakata et al. (1997)
Huang (2001)
Kong & Liu (1999)
H. F. Zhu et al. (2001)
22
22
22
24
18
2x
2x
2x
2x
2x
H. F. Zhu et al. (2001)
Nakata et al. (1997)
H. F. Zhu et al. (2001)
Nakata et al. (1997)
Nakata et al. (1997)
16
2x
H. F. Zhu et al. (2001)
22
2x
H. F. Zhu et al. (2001)
22
2x
Nakata et al. (1997)
16
22
22
22
2x
2x
2x
2x
Nakata et al. (1997)
H. F. Zhu et al. (2001)
H. F. Zhu et al. (2001)
Nakata et al. (1997)
20
24
22
2x
2x
2x
H. F. Zhu et al. (2001)
Nakata et al. (1997)
Nakata et al. (1997)
R. breviglumis Keng ex Y. L. Keng
& S. L. Chen
R. ciliaris (Trin.) Nevski
R. dolichathera Keng ex Y. L. Keng
& S. L. Chen
R. elytrigioides C. Yen & J. L. Yang
R. parvigluma Keng ex Y. L. Keng
& S. L. Chen
R. pendulina Nevski
R. sinica Keng ex Y. L. Chen & S.
L. Chen
R. stricta Keng ex Y. L. Keng & S.
L. Chen
PRIMULACEAE
Androsace bulleyana Forrest
A. spinulifera Knuth
A. yargongensis Petitm.
Pomatosace filicula Maxim.
Primula aurantiaca W. W. Sm.
P. beesiana Forrest
P. bulleyana Forrest
P. forrestii Balf. f.
P. malacoides Franch.
P. monticola (Hand.-Mazz.) F. H.
Chen & C. M. Hu
P. poissonii Franch.
P. pulchella Franch.
P. pulverulenta Duthie
P. secundiflora Franch.
P. sikkimensis Hook.
P. sinolisteri Balf. f.
Locality
n
R.
R.
R.
R.
Lu
Lu
Lu
Lu
et
et
et
et
al.
al.
al.
al.
(1990)
(1990)
(1990)
(1990)
300
Annals of the
Missouri Botanical Garden
Appendix 1.
Continued.
Taxon
2n
Ploidy
Reference
YN: Zhongdian,
4050 m
22
2x
H. F. Zhu et al. (2001)
RANUNCULACEAE
Aconitum brachypodum Diels
YN: Lijiang, 3700 m
16
2x
A. brachypodium Diels var. laxiflorum H. R. Fletcher & Lauener
A. brevicalcaratum Diels
YN: Zhongdian,
3200 m
YN: Heqing, 3700 m
16
2x
32
4x
YN: Lijiang
32
4x
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1994)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1994)
Q. E. Yang (1996)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1994)
Q. E. Yang (2001a)
Q. E. Yang et al.
(1993a)
Q. E. Yang (2001a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1993a)
Q. E. Yang (2001a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
J. Q. Liu (2002)
Q. E. Yang et al.
(1993a)
Q. E. Yang (2001a)
Shang & Li (1984)
P. vialii Delavay ex Franch.
Locality
n
A. brevicalcaratum (Finet & Gagnep.) YN: Lijiang, 4000 m
Diels var. parviflorum Chen & Liu
A. brunneum Hand.-Mazz.
SC: Songpan
A. bulleyanum Diels
YN: Heqing
32
4x
16
16
2x
2x
A. carmichaelii Debeaux
YN: Dali
64
8x
A. contortum Finet & Gagnep.
YN: Yunlong, 3400 m
16
2x
YN: Dali
16
2x
32
4x
A. delavayi Franch.
YN: Zhongdian,
3700 m
SC: Yajiang
YN: Lijiang, 2900 m
32
16
4x
2x
A. episcopale H. Lév.
YN: Dali
YN: Dali
16
16
2x
2x
32
4x
16
2x
A. forrestii Stapf
YN: Zhongdian,
3700 m
YN: Zhongdian,
3970 m
YN: Weixi
YN: Lijiang
16
16
2x
2x
A. georgei Comber
YN: Lijiang, 3200 m
16
2x
A. gymnandrum Maxim.
A. hemsleyanum E. Pritz. ex Diels
QH: Dari, 4050 m
YN: Deqin, 3200 m
16
16
2x
2x
YN: Weixi
SC: Mt. Emei, 2100 m
16
16
2x
2x
SC: Shimian
SC: Yajiang
YN: Yunlong
16
16
16
2x
2x
2x
YN: Weixi, 3600 m
16
2x
YN: Tengchong
16
2x
YN: Deqin, 3600–
4000 m
YN: Deqin, 4200 m
16
2x
16
2x
A. crassiflorum Hand.-Mazz.
A. fengii W. T. Wang
A. hemsleyanum E. Pritz. var. circinatum W. T. Wang
A. legendrei Hand.-Mazz.
A. liljestrandii Hand.-Mazz.
A. nagarum Stapf
A. nagarum Stapf var. heterotrichum
H. R. Fletcher & Lauener
A. nagarum Stapf var. dielsianum
(Airy Shaw) W. T. Wang
A. ouvrardianum Hand.-Mazz.
Q. E. Yang (2001a)
Q. E. Yang (2001a)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1993a)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
301
Continued.
Taxon
A. pendulicarpum Chang ex W. T.
Wang
A. pendulum Busch
Locality
YN: Deqin
n
2n
Ploidy
16
2x
16
2x
32
4x
16
2x
YN: Zhongdian,
3200 m
A. piepunense Hand.-Mazz.
YN: Zhongdian,
3200 m
YN: Zhongdian,
3400 m
YN: Zhongdian
A. piepunense Hand.-Mazz. var. pilos- YN: Zhongdian,
um H. F. Comber
3700 m
A. pulchellum Hand.-Mazz.
YN: Deqin, 4200 m
16
16
2x
2x
16
2x
A. rockii Fletcher & Lauener
16
2x
YN: Zhongdian,
3750 m
YN: Zhongdian,
3970 m
YN: Dali
SC: Ruoergai
32
4x
16
16
2x
2x
SC: Nanping
YN: Deqin, 4300 m
16
16
2x
2x
A. spathulatum W. T. Wang
YN: Heqing,
16
2x
A. stapfianum Hand.-Mazz.
YN: Lijiang
16
2x
A. stramineiflorum Chang ex W. T.
Wang
A. tanguticum (Maxim.) Stapf
A. tongolense Ulbr.
A. transsectum Diels
YN: Weixi
16
2x
QH: Yushu, 3500 m
SC: Lixian
YN: Lijiang, 3600 m
16
16
16
2x
2x
2x
YN: Lijiang, Yulong
16
2x
YN: Lushui, 3200–
3800 m
YN: Zhongdian,
2800 m
YN: Zhongdian
16
2x
16
2x
16
2x
A. wangii Q. E. Yang
YN: Lijiang, 3200 m
16
2x
A. wardii Fletcher & Lauener
YN: Deqin
32
4x
Actaea asiatica Hara
YN:
YN:
YN:
YN:
16
16
16
16
2x
2x
2x
2x
YN: Ninglang
16
2x
YN: Weixi
SC: Nanchuan
YN: Zhongdian
YN: Weixi
32
16
14
14
4x
2x
2x
2x
A. scaposum Franch.
A. sessiliflorum (Finet & Gagnep.)
Hand.-Mazz.
A. sinomontanum Nakai
A. souliei Finet & Gagnep.
A. tsaii W. T. Wang
A. tuguancunense Q. E. Yang
Adonis brevistyla Franch.
Anemoclema glaucifolium (Franch.)
W. T. Wang
Anemone davidii Franch.
A. begoniifolia H. Lév. & Vaniot
A. demissa Hook. f. & Thomson
A. flaccida Fr. Schmidt
Dali
Zhongdian
Zhongdian
Lijiang
Reference
Q. E. Yang
(1989)
Q. E. Yang
(1993a)
Q. E. Yang
(1993a)
Q. E. Yang
(1993a)
Q. E. Yang
Q. E. Yang
(1993a)
Q. E. Yang
(1993a)
Q. E. Yang
(1993a)
Q. E. Yang
(1993a)
Q. E. Yang
Q. E. Yang
et al.
et al.
et al.
et al.
(2001a)
et al.
et al.
et al.
et al.
(2001a)
(1996)
Q. E. Yang (1996)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
J. Q. Liu (2002)
Q. E. Yang (2001a)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1989)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1993a)
Q. E. Yang & Gong
(1995)
Q. E. Yang et al.
(1993a)
Q. E. Yang et al.
(1994)
Q. E. Yang (1998)
Q. E. Yang (2002a)
Q. E. Yang (2001b)
Q. E. Yang (2002b)
G. L. Zhang & Gong
(2002)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
302
Appendix 1.
Annals of the
Missouri Botanical Garden
Continued.
Taxon
2n
Ploidy
YN: Zhongdian
16
2x
Q. E. Yang (2002b)
YN:
YN:
YN:
YN:
Lijiang
Zhongdian
Zhongdian
Dali
16
14
16
14
2x
2x
2x
2x
Q.
Q.
Q.
Q.
YN: Zhongdian
14
2x
Q. E. Yang (2002b)
YN: Gongshan
16
2x
SC: Mt. Emei, 2000 m
YN: Zhongdian
YN: Zhongdian
SC: Mt. Emei
YN: Dali
YN: Weixi
XZ: Motuo
16
24
16
16
32
16
16
2x
3x
2x
2x
4x
2x
2x
YN: Dali, 2800 m
SC: Mt. Emei, 2400 m
YN: Dali
YN: Zhongdian
16
16
16
32
2x
2x
2x
4x
Q. E. Yang et al.
(1993b)
Z. Y. Zhang (1982)
Q. E. Yang (2001b)
Q. E. Yang (2001b)
Shang (1985)
Q. E. Yang (1999a)
Q. E. Yang (2002a)
Q. E. Yang et al.
(1995)
Q. E. Yang (1995)
Z. Y. Zhang (1982)
Q. E. Yang (2002a)
Q. E. Yang (2001b)
YN:
YN:
YN:
YN:
YN:
YN:
Dali
Zhongdian
Deqin
Zhongdian
Dali
Zhongdian
32
48
64
16
32
16
4x
6x
8x
2x
4x
2x
Q.
Q.
Q.
Q.
Q.
Q.
C. foetida L. var. velutina Franch. ex YN: Lijiang
Finet & Gagnep.
C. simplex Wormsk. ex DC.
SC: Muli
C. yunnanensis Hsiao
YN: Lijiang
YN: Deqin
Circaeaster agrestis Maxim.
QH: Datong
Clematis brevicaudata DC.
YN: Deqin
C. chrysocoma Franch.
YN: Lijiang
C. kockiana C. K. Schneid.
YN: Yunlong
C. puberula Hook. f. & Thomson var. YN: Lijiang
ganpiniana (H. Lév. & Vaniot) W.
T. Wang
C. ranunculoides Franch.
YN: Zhongdian
C. rehderiana Craib
YN: Deqin
Coptis teeta Wall.
YN: Gongshan
16
2x
E. Yang
E. Yang
E. Yang
E. Yang
E. Yang
E. Yang
(1993b)
Q. E. Yang
16
16
16
30
16
16
16
16
2x
2x
2x
2x
2x
2x
2x
2x
Q. E. Yang (1999b)
Q. E. Yang (1999b)
Q. E. Yang (2002a)
Kong & Yang (1997)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
Q. E. Yang (2002b)
16
16
18
2x
2x
2x
YN: Zhongdian
SC: Muli
YN: Zhongdian
SC: Ruoergai
QH: Yushu, 4650 m
16
16
16
16
16
2x
2x
2x
2x
2x
Q. E. Yang (2002b)
Q. E. Yang (2002b)
Q. E. Yang et al.
(1993b)
Q. E. Yang (2001a)
Q. E. Yang (2001a)
Q. E. Yang (2001a)
Q. E. Yang (1996)
J. Q. Liu & He (1999)
YN: Weixi
YN: Lijiang
YN: Lijiang
16
16
16
2x
2x
2x
Q. E. Yang (2001a)
Q. E. Yang (2001a)
Q. E. Yang (2001a)
A. hupehensis Hort. ex Boynton f.
alba W. T. Wang
A. rivularis Wall.
A. rupestris Wall.
A. stolonifera Maxim.
A. trullifolia Hook. f. & Thomson
var. colestina (Franch.) Finet &
Gagnep.
A. trullifolia Hook. f. & Thomson
var. holophylla Diels
Asteropyrum peltatum J. R. Drumm.
& Hutchinson
Batrachium bungei (Steud.) L. Liu
Beesia calthifolia Ulbr.
B. deltophylla C. Y. Wu
Calathodes oxycarpa Sprague
Callianthemum pimpinelloides (D.
Don) Hook. f. & Thomson
Caltha palustris L.
Cimicifuga foetida L.
Delphinium beesianum W. W. Sm.
D. bonvalotii Franch.
D. bulleyanum Forrest ex Diels
D. caeruleum Jacquem.
D. candelabrum Ostenf. var. monanthum (Hand.-Mazz.) W. T. Wang
D. ceratophorum Franch.
D. coleopodum Hand.-Mazz.
D. delavayi Franch.
Locality
n
Reference
E.
E.
E.
E.
Yang
Yang
Yang
Yang
(2002b)
(2002b)
(2002b)
(2002b)
(2002a)
(2002a)
(2002a)
(1999b)
(1999b)
et al.
(1999b)
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
303
Continued.
Taxon
2n
Ploidy
YN: Zhongdian
16
2x
Q. E. Yang (2001a)
YN: Zhongdian
SC: Lixian
SC: Nanping
QH: Yushu, 4020 m
16
16
16
16
2x
2x
2x
2x
Q. E. Yang (2001a)
Q. E. Yang (2001a)
Q. E. Yang (1996)
J. Q. Liu & He (1999)
YN: Zhongdian
QH: Yushu, 3540 m
YN: Zhongdian
QH: Yushu, 4000 m
YN: Zhongdian
YN: Dali
QH: Maduo, 4430 m
YN: Zhongdian
QH: Yushu, 3850 m
YN: Weixi
YN: Zhongdian
16
16
16
16
16
16
16
16
16
16
32
2x
2x
2x
2x
2x
2x
2x
2x
2x
2x
4x
Q. E. Yang (2001a)
J. Q. Liu & He (1999)
Q. E. Yang (2001a)
J. Q. Liu & He (1999)
Q. E. Yang (2001a)
Q. E. Yang (2001a)
J. Q. Liu & He (1999)
Q. E. Yang (2001a)
Liu & He (1999)
Q. E. Yang (2001a)
Q. E. Yang (2001b)
SC: Mt. Emei, 2750 m
18
2x
Z. Y. Zhang (1982)
YN: Weixi, 3900 m
YN: Deqin, 3900 m
YN: Zhongdian,
4100 m
YN: Lijiang, 3600 m
YN: Weixi
QH: Maduo, 4840 m
16
16
16
2x
2x
2x
Q. E. Yang (2000a)
Q. E. Yang (2000a)
Q. E. Yang (2000a)
32
32
ca. 42
4x
4x
6x
Q. E. Yang (2000b)
Q. E. Yang (2001b)
Huang et al. (1996a)
R. japonicus Thunb.
SC: Yaan
R. laetus Wall.
YN: Dali
R. nematolobus Hand.-Mazz.
YN: Dali
R. potanini Komarov
YN: Zhongdian
R. sinovaginatus W. T. Wang
YN: Deqin
R. tanguticus (Finet & Gagnep.) Hao SC: Lixian
R. trigonus Hand.-Mazz.
YN: Lijiang
R. wangianus Q. E. Yang
YN: Weixi
R. yunnanensis Franch.
YN: Dali, 3400 m
Souliea vaginata Franch.
YN: Zhongdian
Trollius yunnanensis Ulbr.
YN: Lijiang
14
14
16
32
32
32
16
48
16
16
16
2x
2x
2x
4x
4x
4x
2x
6x
2x
2x
2x
Q.
Q.
Q.
Q.
Q.
Q.
Q.
Q.
Q.
Q.
Q.
ROSACEAE
Dichotomanthes tristaniaecarpa Kurz
YN: Tengchong
34
2x
Malus asiatica Nakai
M. halliana Koehne
SC: Baxian
SC: Zhaojiao
SC: Zhaojiao
68
34
34
4x
2x
2x
M. hupehensis (Pamp.) Rehder
SC: Maerkang
SC: Yanyuan
SC: Xiaojin
51
68
34
3x
4x
2x
L. H. Zhou et al.
(2000)
G. L. Liang (1987)
G. L. Liang (1987)
G. L. Liang & Li
(1993)
G. L. Liang (1987)
G. L. Liang (1987)
G. L. Liang (1987)
SC: Yanyuan
SC: Yanyuan
34
34
2x
2x
D. delavayi Franch. var. pogonanthum (Hand.-Mazz.) W. T. Wang
D. forrestii Diels
D. grandiflorum L.
D. kamaonense Huth
D. kamaonense var. glabrescens (W.
T. Wang) W. T. Wang
D. likiangense Franch.
D. nangchienense W. T. Wang
D. oxycentrum W. T. Wang
D. pylzowi Maxim.
D. smithianum Hand.-Mazz.
D. spirocentrum Hand.-Mazz.
D. tangkulaense W. T. Wang
D. tenii H. Lév.
D. trichophorum Franch.
D. umbrosum Hand.-Mazz.
Halerpestes sarmentosa (Adams) Komarov
Kingdonia uniflora Balf. f. & W. W.
Sm.
Oxygraphis delavayi Franch.
O. glacialis Bunge
O. tenuifolia W. E. Evans
Ranunculus felixii H. Lév.
R. ficariifolia H. Lév. & Vaniot
R. involucratus Maxim.
M. kansuensis (Batalin) C. K.
Schneid.
M. ombrophila Hand.-Mazz.
Locality
n
Reference
E.
E.
E.
E.
E.
E.
E.
E.
E.
E.
E.
Yang
Yang
Yang
Yang
Yang
Yang
Yang
Yang
Yang
Yang
Yang
(2001b)
(2001b)
(2001b)
(2001b)
(2001b)
(2001b)
(2001b)
(2000b)
(2000b)
(2002a)
(2002a)
G. L. Liang (1987)
G. L. Liang & Li
(1993)
304
Annals of the
Missouri Botanical Garden
Appendix 1.
Continued.
Taxon
M. rockii Rehder
M. sikkimensis (Wenzig) Koehne ex
C. K. Schneid.
M. toringoides (Rehd.) Hughes
Locality
2n
Ploidy
Reference
SC: Yanyuan
SC: Zhaojiao
SC: Zhaojiao
34
51
51
2x
3x
3x
SC: Yanyuan
34
2x
SC: Yanyuan
34
2x
SC: Maerkang
SC: Aba
51
34
3x
2x
SC: Maerkang
51
3x
SC: Aba
68
4x
G. L. Liang (1987)
G. L. Liang (1987)
G. L. Liang & Li
(1993)
G. L. Liang & Li
(1993)
G. L. Liang & Li
(1993)
G. L. Liang (1987)
G. L. Liang & Li
(1993)
G. L. Liang & Li
(1993)
G. L. Liang & Li
(1993)
G. L. Liang & Li
(1993)
G. L. Liang (1987)
G. L. Liang & Li
(1993)
Iwatsubo & Naruhashi
(1992)
n
M. xiaojinensis M. H. Cheng & N. G. SC: Xiaojin
Jiang
M. yunnanensis C. K. Schneid.
SC: Yanyuan
SC: Yanyuan
68
4x
34
34
2x
2x
Rubus ichangensis Hemsl. & Kuntze
SC: Mt. Emei
28
4x
RUBIACEAE
Kelloggia chinensis Franch.
YN: Zhongdian
22
2x
Tu et al. (in prep.)
SABIACEAE
Sabia yunnanensis Franch.
SC: Baoxing, 2600 m
2x
Hong & Zhang (1990)
SAXIFRAGACEAE
Chrysosplenium griffithii Hook. f. &
Thomson
12
SC: Kangding, 3560 m
18
2x
Parnassia aff. cacuminum Hand.Mazz. f. yushuensis T. C. Ku
P. brevistyla (Brieg.) Hand.-Mazz.
SC: Songpan, 3850 m
18
2x
SC: Songpan, 3420 m
14
2x
P. delavayi Franch.
SC: Nanping, 3050 m
14
2x
P. viridifolia Batalin
SC: Nanping, 3050 m
36
4x
P. yunnanensis Franch. var. longistip- SC: Kangdin, 3750 m
itata Z. P. Jien
Rodgersia aesculifolia Batalin
SC: Aba, Maoxian
32
4x
30
2x
60
ca. 75
4x
5x
Funamoto et al.
(1997b)
Funamoto et al.
(2000b)
Funamoto et al.
(2000b)
Funamoto et al.
(2000b)
Funamoto et al.
(2000b)
Funamoto et al.
(2000b)
Funamoto et al.
(2001a)
Funamoto et al.
(2001a)
Funamoto et al.
(2001a)
Funamoto et al.
(2001a)
Funamoto et al.
(1997a)
Funamoto et al.
(2000a)
Akiyama et al. (1990)
Akiyama et al. (1990)
SC: Songpan, 3400 m
36
4x
C. lanuginosum Hook. f. & Thomson SC: Dujianyan,
1900 m
C. aff. lixianense Jien ex J. T. Pan
SC: Nanping, 3050 m
16
2x
12
2x
36
4x
52
4x
28
4x
Funamoto et al. (1998)
C. pillosum Maxim. var. valdipilosum SC: Songpan, 3400 m
Ohwi
C. qinlingense Jien ex J. T. Pan
SC: Nanping, 3050 m
R. pinnata Franch.
YN: Lijiang
YN: Lijiang
Saxifraga aculeate Balf. f.
SC: Kangdin, 2400 m
Volume 92, Number 2
2005
Appendix 1.
Nie et al.
Polyploidy in Regional Flora
305
Continued.
Taxon
Locality
n
S. aurantiana Franch.
2n
Ploidy
Reference
16
2x
S. cardiophylla Franch.
S. divaricata Engl. & Irmscher
S. egregia Engl.
SC: Kangdin, 3620 m
SC: Kangdin, 3750 m
SC: Songpan, 3500 m
32
22
16
4x
2x
2x
S. heleonastes Harry Sm.
SC: Songpan, 3350 m
16
2x
S. hirculus L.
SC: Songpan, 3850 m
32
4x
S. aff. melanocentra Franch.
SC: Songpan, 3850 m
22, 23
2x
S. montana Harry Sm.
Tiarella polyphylla D. Don
SC: Kangdin, 3540 m
SC: Mt. Emei
32
14
4x
2x
Funamoto et al.
(2001b)
Funamoto et al. (1998)
Funamoto et al. (1998)
Funamoto et al.
(2001b)
Funamoto et al.
(2001b)
Funamoto et al.
(2001b)
Funamoto et al.
(2001b)
Funamoto et al. (1998)
Soltis & Bohm (1984)
YN: Zhongdian
16
2x
J. Cai et al. (2004)
YN: Zhongdian
YN: Zhongdian
YN: Zhongdian
16
16
14
2x
2x
2x
J. Cai et al. (2004)
J. Cai et al. (2004)
J. Cai et al. (2004)
YN: Zhongdian
16
2x
J. Cai et al. (2004)
YN: Zhongdian
YN: Zhongdian
YN: Zhongdian
16
16
14
2x
2x
2x
J. Cai et al. (2004)
J. Cai et al. (2004)
J. Cai et al. (2004)
YN: Zhongdian
YN: Deqin
YN: Zhongdian
YN: Zhongdian
YN: Zhongdian
SC: Mt. Emei, 1700 m
SC: Mt. Emei, 1000 m
SC: Baoxing, 1600 m
SC: Baoxing, 2500 m
16
16
16
16
16
2x
2x
2x
2x
2x
4x
2x
2x
2x
J. Cai et al. (2004)
J. Cai et al. (2004)
J. Cai et al. (2004)
J. Cai et al. (2004)
J. Cai et al. (2004)
Hong & Zhang (1990)
Hong & Zhang (1990)
Hong & Zhang (1990)
Hong & Zhang (1990)
SCROPHULARIACEAE
Pedicularis densispica Franch. ex
Maxim.
P. dichotoma Bonati
P. dolichocymba Hand.-Mazz.
P. integrifolia Hook. f. subsp. integerrima (Pennell & Li) Tsoong
P. longiflora Rudolph var. tubiformis
(Klotzsch) Tsoong
P. oxycarpa Franch. ex Maxim.
P. rex C. B. Clarke ex Maxim.
P. siphonantha D. Don var. delavayi
(Franch. ex Maxim.) Tsoong
P. strobilacea Franch. ex Hemsl.
P. trichoglossa Hook. f.
P. tricolor Hand.-Mazz.
P. tsekouensis Bonati
P. umbelliformis Li
Veronica henryi T. Yamaz.
V. javanica Blume
V. laxa Benth.
V. serpyllifolia L.
SMILACACEAE
Heterosmilax yunnanensis Gagnep.
16
16
16
7
SC: Mt. Emei, 1700 m
32
2x
S. menispermoidea A. DC.
SC: Mt. Emei
SC: Mt. Emei
SC: Mt. Emei
32
32
32
2x
2x
2x
S. tsingchengshanensis Wang
SC: Mt. Emei, 1500 m
32
2x
Fu et al. (1995a,
1995b)
Fu et al. (1993)
Fu et al. (1992)
Fu et al. (1995a,
1995b)
Fu et al. (1993)
48
4x
Tu et al. (2005)
48
48
4x
4x
Tu et al. (2005)
Tu et al. (2005)
48
48
48
34
4x
4x
4x
2x
Tu
Tu
Tu
Tu
Smilax chapaensis Gagnep.
SOLANACEAE
Anisodus acutangulus C. Y. Wu & C. YN: Lijiang, 3100 m
Chen
SC: Daocheng, 3750 m
A. carniolicoides (C. Y. Wu & C.
YN: Zhongdian,
Chen) D’Arcy & Zhi Y. Zhang
3370 m
A. luridus Link & Otto
YN: Heqing, 2390 m
A. sinensis Pascher
SC: Shizhu, 2900 m
A. tanguticus Pascher
SC: Daocheng, 3750 m
Hyoscyamus niger L.
SC: Ganzi, 3400 m
et
et
et
et
al.
al.
al.
al.
(2005)
(2005)
(2005)
(2005)
306
Annals of the
Missouri Botanical Garden
Appendix 1.
Continued.
Taxon
Mandragora caulescens C. B. Clarke
Przewalskia tangutica Maxim.
Solanum spirale Roxb.
THEACEAE
Camellia pitardii Cohen Stuart
C. reticulata Lindl.
C. saluenensis Stapf ex Bean
Locality
YN: Zhongdian,
3700 m
QH: Maduo, 4000 m
SC: Shiqu, 4320 m
XZ: Motuo, 900 m
SC: Huili, 2150 m
SC: Yanbian, 1650 m
SC: Miyi, 1850 m
YN: Tengchong
YN: Tengchong
SC: Yanbian
YN: Tengchong
SC: Panzhihua, 1700
m
SC: Dukou, 2000 m
SC: Huili, 2650–
2900 m
SC: Huili, 2650–
2900 m
SC: Yanbian, 2600 m
YN: Huaping, 1900 m
YN: Baoshan
SC: Huili, 1650–
1900 m
VALERIANACEAE
Valeriana hardwickii Wall. subsp. la- SC: Mt. Emei, 1600 m
tifolia (Rehd. & Wils.) Y. F. Wu
VELLOZIACEAE
Acanthochlamys bracteata P. C. Kao
n
15
30
45
45
15
45
45
15
45
15
15
2n
Ploidy
48
4x
Tu et al. (2005)
44
44
48
4x
4x
4x
Huang et al. (1996a)
Tu et al. (2005)
Gu & Sun (1998)
30
60
90
90
2x
4x
6x
6x
6x
2x
6x
6x
Gu (1997)
Gu (1997)
Gu (1997)
Gu et al. (1988)
T. J. Xiao et al. (1993)
Xia et al. (1994)
Gu et al. (1988)
Gu (1997)
90
30
6x
2x
T. J. Xiao et al. (1996)
T. J. Xiao et al. (1996)
60
4x
T. J. Xiao et al. (1996)
30
90
2x
6x
2x
2x
Gu (1997)
Gu (1997)
T. J. Xiao et al. (1993)
Gu (1997)
2x
Hong & Zhang (1990)
30
30
8
Reference
SC: Daofu, 2700–
3400 m
38
2x
Kao et al. (1993)
ZINGIBERACEAE
Caulokaempferia yunnanensis (Gagnep.) R. M. Sm.
Globba emeiensis Z. Y. Zhu
SC: Dukou
42
2x
SC: Mt. Emei
24
2x
Roscoea tibetica Batalin
YN: Lijiang
24
2x
Z. Y. Chen et al.
(1988)
Z. Y. Chen et al.
(1988)
Z. Y. Chen et al.
(1988)
Volume 92, Number 2, pp. 139–306 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN
was published on July 26, 2005.