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. 276 Annals of the Missouri Botanical Garden 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 Volume 92, Number 2 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- 278 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 Volume 92, Number 2 2005 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- 280 Annals of the Missouri Botanical Garden 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- Volume 92, Number 2 2005 Nie et al. 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 source of the floristic development of the region, with the existence of migration routes for plants from the north to the south and vice versa (H. Sun, 2002a, 2002b; H. Sun & Li, 2003). Brochmann, C. & S. W. Steen. 1999. Sex and genes in the flora of Svalbard—Implications for conservation biology and climate change. Det Norske Videnskaps-Akademi I Matematisk-Naturvidenskapelig Klasse, Skrifter, Ny Serie 38: 33–72. , A. K. Brysting, I. G. Also, L. Borgen, H. H. Grundt, A.-C. Scheen & R. Elven. 2004. Polyploidy in arctic plants. Biol. J. Linn. Soc. 82: 521–536. Cai, J., H. Wang, Z. J. Gu, R. R. Mill & D. Z. Li. 2004. Karyotype morphology of thirteen species of Pedicularis (Orobanchaceae) from the Hengduan Mountains Region, NW Yunnan, China. 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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.