青藏高原特有属翅果蓼属的生物地理学研究
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摘要
翅果蓼属(Parapteropyrum)是青藏高原特有的单种属,该属与分布于中亚地区的木蓼属(Atraphaxis)、沙拐枣属(Calligonum)以及双翅蓼属(Pteropyrum)起组成蓼科(Polygonaceae)木蓼族(Atraphaxideae)。尽管翅果蓼属与该族其它三个属的地理分布区相隔很远,但是由于翅果蓼适应生存于干旱河谷地区的木质化灌木生存状态,以及在形态学上与木蓼族植物的若干相似性状,植物分类学家长期以来都一致地把它归入到木蓼族中,并认为与该族中的双翅蓼属有很近的亲缘关系。近期分子系统学的研究表明,翅果蓼与荞麦属有很近的亲缘关系,在分子系统树中与荞麦属聚为一支。然而翅果蓼确切的系统位置仍然模糊不清,其起源进化问题有待进一步深入的研究。本文通过细胞染色体、分子系统学等研究手段对翅果蓼的系统位置、起源进化等问题展开深入的研究和探讨。细胞染色体的研究证据表明,木蓼族各属的染色体总数和基数完全不一致,核型类型也存在明显差异：翅果蓼的染色体总数是2n=48=16m+16sm+16st,核型属于2B型,对称性很低；木蓼属两个种(东北木蓼,沙木蓼)的染色体总数都是2n=22=22m,核型属于1A型,对称性相当高；沙拐枣属的染色体总数为2n=18或者36。在已报道的分布于青藏高原及其周边地区的蓼科植物中,只有荞麦属的染色体基数X=8(2n=16或32),与翅果蓼相同。进一步通过核糖体DNA的内转录间隔区(ITS)和叶绿体片段rbcL、accD以及两套序列联合分别构建的系统发育树表明：木蓼族不是一个单系群；翅果蓼属与该族的其它三个属存在较大的遗传分化,而是深嵌于荞麦属(Fagopyrum)中,并以很高的支持率与荞麦属的一支形成姊妹群。形态学上,翅果蓼与荞麦属植物在几个主要的形态特征上也有非常大的相似性：总状花序,花被片5、深裂,8个雄蕊排成内外两轮(5个在外轮,3个在内轮),头状柱头3,三棱形的瘦果。综合形态学、细胞学以及分子系统学的证据,翅果蓼不能归入木蓼族中,应撤销其属级分类地位,并入到荞麦属中。通过分子进化速率标记的分化时间表明：翅果蓼与荞麦的分化时间大约发生在6.35-14.8百万年之间。这个时间段(尤其在15-13百万年和8-7百万年)正是青藏高原隆升的重要时期。我们推测第三纪中新世以来,由于青藏高原的强烈隆升,同时伴随着气候的剧烈变化,高原生境发生了严重的破碎化。在这种情况下,辐射式物种形成和趋同进化成为青藏高原物种形成和进化的主要方式。为了适应不断变化的环境和气候,提高自身的适合度,防止由于地理隔离而导致的近交衰退,翅果蓼从草本的荞麦属中分化出来,进化出了木质化的茎。因此,岛屿木质化这(insular woodiness)一岛屿特有的物种进化方式在青藏高原这个内陆大岛屿上也存在。另外,翅果蓼这样一种木本荞麦作为一种新资源将会在以后的种质资源开发应用中扮演重要的角色,这在解决日益严重的粮食危机方面可能有重要的应用意义。
Parapteropyrum, a monotypic shrubby genus with the only endangered species endemic to the arid valleys of the central Qinghai-Tibetan Plateau (QTP), was universally placed in the Atraphaxideae because of their shared morphological traits although the other woody species of this tribe occurred mainly in the central Asia. It is suggested that Parapteropyrum is closely related to the small genus Pteropyrum based on morphological traits. However, recent molecular phylogenetic studies suggested that Parapteropyrum is closely related to Fagopyrum. The accurate position of Parapteropyrum is still uncertain and further research works are still needed to clarify the origination and evolution of this monotypic genus. In our studies, the karyological data show that the chromosome number and karyotype of P. tibeticum are formulated as 2n=48=16m+16sm+16st, belonging to 2B type. Both Atraphaxis manshurica and A. bracteata have the same chromosome number of 2n=22 and similar karyotypes (22m, belonging to 1A). The basic chromosome number of another genus (Calligonum) of the Atraphaxideae is x=9 (2n=18 or 36). Across the published karyological data of Polygonaceae that occurred in the QTP or adjacent areas, only Fagopyrum (2n=16 or 32) shares the same basic chromosomes with Parapteropyrum. We studied sequence variations of the nuclear internal transcribed spacer (ITS) and chloroplast DNA (cpDNA; rbcL and accD) for this genus and the other ten related genera of the Polygonaceae occurred in the QTP and adjacent areas. The constructed phylogenies based on ITS, cpDNA or a combination of both datasets suggested that currently recognized tribe Atraphaxideae is polyphyletic. The woody Parapteropyrum nested deeply within the herbal Fagopyrum and it showed no any relationships with the other genera of the Atraphaxideae. Morphologically, many common characters were also shared by Fagopyrum and Parapteropyrum:racemous inflorescence,5 deeply lobed perianth, stamens 8 (5 outer,3 inner), styles 3 with head-like stigmas, triangular achenes. Totally, the karyological, genetic and morphological evidences all suggested that shrubby P. tibeticum has derived from herbal Fagopyrum. Our results suggested that Parapteropyrum should be reduced and its only species should be incorporated under the genus Fagopyrum. Molecular datings revealed that this woody species probably originated between 6.35 and 14.8 million years ago (Mya). Meanwhile, during this stage, geological evidence suggested that the Qinghai-Tibetan Plateau had extensively uplifted, especially between 15-13 Mya and 8-7 Mya. The high species diversity in the QTP was suggested to result mainly from radiative diversification and convergent evolution of plants because of the rapid plateau uplifts and the dramatic climate change. These results together suggested that the large-scale QTP uplifts since the Miocene promoted not only extensive speciation in this region, but also the secondary acquirements of woodiness in a few herbal lineages under the strong counter-selection of inbreeding depression because of the geographical isolation. It is confirmed that insular woodiness has also happened in the QTP as occurred in most islands and its evolutionary significance is also discussed in detail. In addition, the confirmation of this woody species within Fagopyrum further implies its possible usage as an important buckwheat resource because of the rising food crisis.
Parapteropyrum, a monotypic shrubby genus with the only endangered species endemic to the arid valleys of the central Qinghai-Tibetan Plateau (QTP), was universally placed in the Atraphaxideae because of their shared morphological traits although the other woody species of this tribe occurred mainly in the central Asia. It is suggested that Parapteropyrum is closely related to the small genus Pteropyrum based on morphological traits. However, recent molecular phylogenetic studies suggested that Parapteropyrum is closely related to Fagopyrum. The accurate position of Parapteropyrum is still uncertain and further research works are still needed to clarify the origination and evolution of this monotypic genus. In our studies, the karyological data show that the chromosome number and karyotype of P. tibeticum are formulated as 2n=48=16m+16sm+16st, belonging to 2B type. Both Atraphaxis manshurica and A. bracteata have the same chromosome number of 2n=22 and similar karyotypes (22m, belonging to 1A). The basic chromosome number of another genus (Calligonum) of the Atraphaxideae is x=9 (2n=18 or 36). Across the published karyological data of Polygonaceae that occurred in the QTP or adjacent areas, only Fagopyrum (2n=16 or 32) shares the same basic chromosomes with Parapteropyrum. We studied sequence variations of the nuclear internal transcribed spacer (ITS) and chloroplast DNA (cpDNA; rbcL and accD) for this genus and the other ten related genera of the Polygonaceae occurred in the QTP and adjacent areas. The constructed phylogenies based on ITS, cpDNA or a combination of both datasets suggested that currently recognized tribe Atraphaxideae is polyphyletic. The woody Parapteropyrum nested deeply within the herbal Fagopyrum and it showed no any relationships with the other genera of the Atraphaxideae. Morphologically, many common characters were also shared by Fagopyrum and Parapteropyrum:racemous inflorescence,5 deeply lobed perianth, stamens 8 (5 outer,3 inner), styles 3 with head-like stigmas, triangular achenes. Totally, the karyological, genetic and morphological evidences all suggested that shrubby P. tibeticum has derived from herbal Fagopyrum. Our results suggested that Parapteropyrum should be reduced and its only species should be incorporated under the genus Fagopyrum. Molecular datings revealed that this woody species probably originated between 6.35 and 14.8 million years ago (Mya). Meanwhile, during this stage, geological evidence suggested that the Qinghai-Tibetan Plateau had extensively uplifted, especially between 15-13 Mya and 8-7 Mya. The high species diversity in the QTP was suggested to result mainly from radiative diversification and convergent evolution of plants because of the rapid plateau uplifts and the dramatic climate change. These results together suggested that the large-scale QTP uplifts since the Miocene promoted not only extensive speciation in this region, but also the secondary acquirements of woodiness in a few herbal lineages under the strong counter-selection of inbreeding depression because of the geographical isolation. It is confirmed that insular woodiness has also happened in the QTP as occurred in most islands and its evolutionary significance is also discussed in detail. In addition, the confirmation of this woody species within Fagopyrum further implies its possible usage as an important buckwheat resource because of the rising food crisis.
引文
顾志建,王丽,孙航,武素功.1993.青藏高原一些种子植物的核型研究.云南植物研究.15(4)：377-384.
李安仁.1998.中国植物志第25卷第1册,科学出版社
李吉均,施雅风,李炳元.1995.青藏高原隆升与全球变化.甘肃人民出版社,兰州
李吉均,方小敏.1998.青藏高原隆起与环境变化研究。《科学通报》,年,43(15)：1569-1574.
施雅风,李吉均,李炳元.1998.青藏高原晚新生代隆升与环境变化.广东：广东科技出版社
潘保田,李吉钧.1996.青藏高原：全球气候变化的驱动机与放大器,Ⅲ.青藏高原隆起对气候变化的影响.兰州大学学报.32：108-115.
吴征镒.1987.西藏植物区系的起源及其演化.北京,科学出版社.5：874-902.
李文华,周兴民.1998.青藏高原生态系统及优化利用模式.广东：广东科技出版社.
Baltisberger M., Widmer A.2006. Chromosome numbers of plant species from the Canary Islands. Botanica Helvetica 116:9-30.
Bao, B.J., Li, A.R.1993. A study of the genus Atraphaxis in China and the system of Atraphaxideae (Polygonaceae). Acta Phytotaxonomica. Sinica 31 (2); 127-139.
Beerling, D.J., Bailey, J.P., Conolly, A.P.1994. Fallopia japonica (Houtt.) Ronse Decraene. Journal of Ecology 82:959-979.
Ballard, H.E., Sytsma, K.J.2000. Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae):Arctic origins, herbaceous ancestry and bird dispersal. Evolution 54: 1521-1532.
Brandbyge, J.1993. Polygonaceae. In:K. Kubitzki, J.C. Rohwer, and V. Bittrich, eds The families and genera of vascularplants, Springer-Verlag, Berlin.2:531-544.
Bohle, U.R., Hilger, H.H., Martin, W.F.1996. Island colonization and evolution of the insular woody habit in Echium L. (Boraginaceae). Proceedings of the National Academy of Sciences of the United States of America 93:11740-11745.
Carlquist, S.1970. Hawaii, a natural history:geology, climate, native flora and fauna above the shoreline. Pacific Tropical Botanical Garden, Lawai, HI.
Carlquist, S.1974. Island biology. Columbia Univ. Press, New York.
Carlquist, S.2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Journal of Linnean Society 141:25-51.
Chen, Q.F.1999. A study of resources of Fagopyrum (Polygonaceae) native to China. Botanical Journal of Linnean Society 130:53-64.
Chung, S.L., Lo, C.H., Lee, T.Y., Zhang, Y.Q., Xie, Y.W., li, X.H., Wang, P.L.1998. Diachronomous uplift of the Tibetan plateau starting 40 Myr ago. Nature 394:769-773.
Cui, X.K., Ao, C.Q., Zhang, Q., Chen, L.T., Liu, J.Q.2008. Diploid and tetraploid distribution of Allium przewalskianum Regel. (Liliaceae) at the regional scale:implications for cryptic speciation through intraspecific polyploidization. Caryologia 61(2):192-200.
Darwin, C.1859. The Origin of Species by Means of Natural Selection.John Murray, London.
Doyle, J.J., Doyle, J.L.1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19:11-15.
Drummond, A.J., Ho, S.Y., Rawlence, N., Rambaut, A.2007. A rough guide to BEAST 1.4. University of Auckland, Aukland, New Zealand.
Carlquist, S.2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Botanical Journal of Linnean Society 141:25-51.
Farris, J.S., Kallersjo, M., Kluge, A.G., Bult, C.1995. Testing signifcance of incongruence. Cladistics 10:315-319.
Fay, M.F., Swensen, S.M., Chase, M.W.1997. Taxonomic affinities of Medusagyne oppositifolia (Medusagynaceae). Kew Bull.52:111-120. Academic Press, New York.
Felsenstein, J.1985. ConWdence limits on phylogenies:an approach using the bootstrap. Evolution 39:783-791.
Friesen, N., Fritsch, R.M., Pollner, S., Blattner, F.R.2000. Molecular and morphological evidence for an origin of the aberrant genus Milula within Himalayan species of Allium (Alliaceae). Molecular Phylogenetics and Evolution 17:209-218.
Grant, V.1981. Plant speciation,2nd edn. Columbia University Press, New York.
Givnish, T.J.1998. Adaptive radiation of plants on oceanic islands:classical patterns, molecular data, new insights. Pp.281-304 in P. Grant, ed. Evolution on islands. Oxford Univ. Press, Oxford, U.K.
Givnish, T.J., Kendra, M.C., Mast, A.R., Paterson, T.B., Theim, T.J., Hipp, A.L., Henss, J.M., Smith, J.F., Kenneth, R., Wood, K.R., Sytsma, K.J.2009. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales:Campanulaceae). Proceeding of the Royal Society 276:407-416.
Gross, H.1913. Bertrage ur der Polygonaceae. Botanische Jahrbucher fur Systematik 49: 234-248.
Guindon, S., Cascuel, O.2003. A Simple, Fast, and Accurate Algorihm to Estimate Large Phylogenies by Maximum Likelihood. Sysematic Biology 52:696-704.
Hanldson, K.1978. Anatomy and taxonomy in Polygonaceae subfam. Polygonoideae hleisn. emend. Jaretzky. Symbolae Botanicae Upsaliensis 22:1-95.
Jaretzky, R.1925. Beitriige zur Systematik der Polygonaceae unter Berucksichtigung des Oxymethyl-anthrachinon-Vorkommens. Feddes Repertorium Specierum Novarum Regni Vegetabilis 22:49-83.
Hedberg, O.1997. The genus Koenigia L. emend. Hedberg (Polygonaceae). Botanical Journal of Linnean Society 124:295-330.
Hong, D.Y.1990. Plant cytotaxonomy. Science Press, Beijing.
Hong, S.P.1995. Pollen morphology of Parapteropyrum and some putatively related genera (Polygonaceae- Atraphaxideae). Grana 34:153-159.
Hong, S.P.1998. Systematic significance of tepal surfac morphology in tribes Persicarieae and Polygoneae (Polygonaceae). Journal of Linnean Society 127:91-116.
Hu, Y.P., Xie, X.L., Wen, Q., Zhao, X.D., Wang, L., Li, Y.2007. Studies on karyotype of five polulations of Rheum tanguticum. Acta Botanica Yunnanica 29(4):429-433.
Huelsenbeck, J.P., Rannala, B.1997. Phylogenetic Methods Come of Age:Testing Hypotheses in an Evolutionary Context. Science 276:227-232.
Huelsenbeck, J.P, Ronquist, F.2001. MrBayes:Bayesian inference of phygeny. Bioinformatics 17: 754-755.
Hughes, C., Eastwood, R.2006. Island radiation on a continental scale:exceptional rates of plant diversification after uplift of the Andes. Proceedings of the National Academy of Sciences of the United States of America 103:10334-10339.
Kadereit. J.W.1994. Molecules and morphology, phylogenetics and genetics. Botanica Acta 107: 369-373.
Kong, H.Z., Liu, J.Q.1999. Karyomorphology of the genus Pomatosace Maxim. (Primulaceae). Acta Phytotaxonomica Sinica 37:445-450.
Lamb-Frye, A.S., Kron, K.A.2003. rbcL phylogeny and character evolution in Polygonaceae. Systematic Botany.28(2):326-332.
Li, A.R.1981. Parapteropyrum A. J. Li—Unum genus novum Polygonacearum sinicum. Acta Phytotaxonomica Sinica 19:330-331.
Li, J.J., Shi, Y.F.. Li, B.Y.1995. Uplift of the Qinghai-Xizang (Tibet) Plateau and Global Change. Lanzhou:Lanzhou University Press.
Liedo, M.D., Crespo, M.B., Cameron, K.M., Fay, M.F., Chase. M.W.1998. Systematics of Plumbaginaceae Based upon Cladistic Aanlysis of rbcL Sequence Data. Systematic Botany 23:21-29.
Liu, J.Q., Zhou, G.Y., Ho, T.N., Lu, A.M.1999. Karyomorphology of Sinadoxa and its systematic significance. Caryologia 52:159-164.
Liu, J.Q.2000. Karyomorphology of 4 species in Sinacalia and Parasenecio (Astetaceae: Senecioneae). Acta Botanical Yunnanica 22:447-450.
Liu, J.Q.. Chen, Z.D., Lu. A.M.2000. The phylogenetic relationships of the Qinghai-Tibet endemic Sinadoxa, revealed by the ITS data. Acta Phytotaxonomica Sinica 42:656-658.
Liu, J.Q., Ho, T.N., Chen, S.L.2002. The first chromosome data documentations of Megacodon and Lomatogoniopsis and their systematic significance. Acta Biologica Plateau Sinica 15: 41-47.
Liu, J.Q., Gao, T.G., Chen, Z.D., Lu, A.M.2002. Molecular phylogeny and biogeography of the Qinghai-Tibet Plateau endemic Nannoglottis (Asteraceae). Molecular Phylogenetics and Evolution 23:307-325.
Liu, J.Q.2004. Uniformity of karyotypes in Ligularia (Asteraceae:Senecioneae), a highly diversified genus of the eastern Qinghai-Tibet Plateau highlands and adjacent areas. Botanical Journal of Linnean Society 144:329-342.
Liu, J.Q., Wang, Y.J., Wang, A.L., Hideaki, O., Abbott, R.J.2006. Radiation and diversification within the Ligularia-Cremanthodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution 38:31-49.
Liu, J.Q., Tian, B.2007. Origin, evolution, and systematics of Himalaya endemic genera. Newsletters of Himalayan Botany 40:20-27.
Mao, Z.M., Yang, G., Wang, C.G.1983. Studies on chromosome number and anatomy of young branches of Calligonum of Xinjiang in relation to the evolution of some species of the genus. Acta Phytotaxonomica Sinica 21:44-48.
Mao, K.S., Hao, G., Liu, J.Q., Adams, R.P., Milne, R.I.2010. Diversification and biogeography of Juniperus (Cupressaceae):variable diversification rates and multiple intercontinental dispersals. New Phytologist 188:254-272.
Maude, P.F.1939. A list of the chromosome numberals of species of British flowering plants. New Phytologist 38:1-31.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., Fonseca, G.A.B., Kent, J.2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.
Nair, P.K., Rehman, K., Saxena, A.K.1976. Contribution to the pollen morphology of Indian Polygonaceae. Journal of Palynotogy 12:1-18.
Ni, Z.C., Cheng, S.Z.1987. On genera endemic to China and occurring on the Xizang Plateau flora. Acta Phytotaxonomica Sinica 25:416-429.
Nie, Z.L., Gu, Z.J., Sun, H.2002. Cytological study of genus Tibetia (Fabaceae) in the Hengduan Mountains region, China. Journal of Plant Research 115:17-22.
Nie, Z.L., Wen, J., Gu, Z.J., Boufford, D.E., Sun, H.2005. Polyploidy in the flora of the Hengduan mountains hotspot, southwestern China. Annals of the Missouri Botanical Garden 92:275-306.
Nowicke, J.V., Skvarla, J.J.1977. Pollen morphology amd the relationship of the Plumbaginaceae, Polygonaceae and Primulaceae to the order Centrospermae. Smithsonian Contributions to Botany 37:1-64.
Nowicke, J.V., Skvarla, J.J.1979. Pollen morphology:The potential influence in higher order systematics. Annals of the Missouri Botanical Garden 66:633-700.
Ohba, H.,1988. The alpine flora of the Nepal Himalayas:An introductory note. In:Ohba H, Malla SB (Eds.), The Himalayas Plants, vol.1. Tokyo:University of Tokyo Press,19-46.
Ohnishi, O., Matsuoka, Y.1996. Search for the wild ancestor of buckwheat Ⅱ. Taxonomy of Fagopyrum (Polygonaceae) species based on morphology, isozymes and cpDNA variability. Gene & Genetic Systems 71:383-390.
Ohnishi, O.1998. Search for the wild ancestor of buckwheat I. Description of new Fagopyrum (Polygonaceae) species and their distribution in China and the Himalayan hills. Fagopyrum 15:18-28.
Ohsako, T., Ohnishi, O.2000. Intra- and interspecific phytogeny of wild Fagopyrum (Polygonaceae) species based on nucleotide sequences of noncoding regions in chloroplast DNA. American Journal of Botany 87:573-582.
Raven, P.H.1975. The bases of angiospenn phylogeny:cytology. Annals of the Missouri Botanical Garden 62:724-764.
Panero, J.L., Francisco-Ortega, J., Jansen, R.K., Santos-Guerra, A.1999. Molecular evidence for multiple origins of woodiness and a New World biogeographic connection of the Macaronesian Island endemic Pericallis (Asteraceae:Senecioneae). Proceedings of the National Academy of Sciences of the United States of America 96 (24):13886-13891.
Posada, D., Crandall, K.A.1998. Modeltest:testing the model of DNA substitution. Bioinformatics 14:817-818.
Pryer, K.M., Schneider, H., Smith, A.R., Cranfill, R., Wolf, P.G., Hunt, J.S., Sipes, S.D.2001 Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Nature 409:618-622.
Ramsey, J.2011. Polyploidy and ecological adaptation in wild yallow. Proceedings of the National Academy of Sciences of the United States of America 108(17):7096-7101.
Rambaut, A., Drummond, A.J.2010. Tracer v 1.5. Availableat:http://beast.bio.ed.ac.uk/Tracer (ac cessed 10 January 2010).
Roberty, G., Vaurier, S.1964. Les genres de Polygonacees. Boissien 10:7-128.
Richardson, J.E., Pennington, R.T., Pennington, T.D., Hollingsworth, P.M.2001. Rapid diversification of a species-rich genus of neotropical rain forest trees. Science 293:2242-2245.
Sanchez, A., Kron, K.A.2008. Phylogenetics of Polygonaceae with an Emphasis on the Evolution of Eriogonoideae. Systematic Botany 33 (1):87-96.
Sanchez, A., Kron, K.A.2008. Preliminary phylogenetic relationships of Calligonum L. (Polygonaceae) using two chloroplast genes. Botany conference, Abstract ID:98. Website:http://www.2008.botanyconference.org/engine/search/index.php?func=detail&aid= 98.
Sanchez, A., Schuster, T.M., Kron, K.A.2009. A large-scale phylogeny of polygonaceae based on molecular data. International Journal of Plant Sciences 1708:1044-1055.
Saiki, R.K., Celfand, D.H., Stoffel, S., Scharf, S.J., Higuchi, R., Horn, G.T., Mullis, K.B., Erlich, H.H.A.1988. Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase. Science 239:487-491
Shi, Y.F., Li, J.J., Li. B.Y.1998. Uplift and environmental changes of Qinghai-Tibetan Plateau in the late Cenozoic. Guangzhou:Guangdong Science and Technology Press.
Swofford, D.L.2002. paup*:Phylogenetic Analyses Using Parsimony (* and Other Methods), Version 4. Sinauer & Associates, Sunderland, MA.
Soltis, D., Soltis, P., Jennifer, A.T.2003. Advances in the study of polyploidy since Plant speciation. New Phytologist 161:173-191.
Stebbing, G.L.1971. Chromosome evolution in higher plants. Edward Arnold, London.
Stuessy, T.F.1998. Evolution and speciation of island plants.Cambridge University Press.
Simmonds, N.W.1945. Biological flora of the British Isles. Journal of Ecology 33(1):117-120.
Sun, W., Zhou, Z.Z., Liu, M.Z., Wan, H.W., Dong, X.2008. Reappraisal of the generic status of Pteroxygonum (Polygonaceae) on the basis of morphology, anatomy and nrDNA ITS sequence analysis. Journal of Systematics and Evolution 46 (1):73-79.
Tang, H.G., Meng, L.H., Ao, C.Q., Liu, J.Q.2005. Origin of the Qinghai-Tibetan Plateau endemic Milula (Liliaceae):further insights from karyological comparisons with Allium. Caryologia 58:320-331.
Tamura, K., Dudley, J., Nei, M., Kumar, S.2007. MEGA4:Molecular Evolutionary Genetics Analysis (MEGA) v755 software version 4.0. Molecular Phylogenetics and Evolution 24: 1596-1599.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.2008. Morphological cladistic analysis of Calligonum and Pteropyrum (Polygonaceae) in Iran. Iranian Journal of Biotechnology 14: 117-125.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.2010. The phylogeny of Calligonum and Pteropyrum (Polygonaceae) based on nuclear ribosomal DNA ITS and chloroplast trnL-Fsequences. Iranian Journal of Biotechnology 8:7-15.
Tapponnier, P., Xu, Z.Q.. Roger, F., Meyer, B., Arnaud, N., Wittlinger, G., Yang J.S.2001. Oblique stepwise rise and growth of the Tibet Plateau. Science 294:1671-1677.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F., Higgins, D.G.1997. Clustal-X windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic acids research 25:4876-4882.
Tsukaya, H., Tsuge. T.2001. Morphological adaptation of inflorescences in plants that bloom atlow temperature in early spring:the convergent evolution of downy plants. Plant Biology 3:536-543.
Tsumura, H.2002. Optical and anatomical characteristics of bracts from the Chinese "glasshouse" plant, Rheum alexandrae Batalin (Polygonaceae), Yunnan, China. Journal of Plant Research 115:59-63.
Tu, T.Y, Sun, H., Gu, Z.J., Yue, J.P.2005. Cytological studies on the Sino-Himalayan endemic Anisodus and four related genera from the tribe Hyoscyameae (Solanaceae), and their systematic and evolutionary implications. Botanical Journal of Linnean Society 147: 457-468.
Tu, T.Y., Volis, S., Dillon, M.O., Sun, H., Wen, J.2010. Dispersals of Hyoscyameae and Mandragoreae (Solanaceae) from the New World to Eurasia in the early Miocene and their biogeographic diversification within Eurasia. Molecular Phylogenetics and Evolution 57: 1226-1237.
Wallace AR.1878. Tropical Nature and Other Essays. Macmillan, London.
Wang, A.L., Yang, M.H., Liu, J.Q.2005. Molecular phylogeny, recent Radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequence. Annals of Botany 96:489-498.
Wang, Y.J., Susanna, A., Von Raab-Straube, E., Milne, R., Liu, J.Q.2009. Island-like radiation of Saussurea (Asteraceae:Cardueae) triggered by uplift of the Qinghai-Tibet Plateau. Biological Journal of the Linnean Society 97:893-903.
White, T.J., Birns, T., Lee, S., Taylor, J.1990. Amplification and direct sequencing of fungal ribosomal RNA genes fro phylogenetics. Pp.315-322 in M. A. Innis, D. H. Gelfand, J. J.Sninsky, and T. J. White, eds. PCR Protocols:A Guide to Methods and Applications.
Wu, C.Y.1987. Flora of Tibet, Vol.5. Science Press, Beijing.
Wu, C.Y.1988. Hengduan Mountains flora and her sighificane. Journal of Japanese Botany 63: 297-311.
Wu, S.G., Yang, Y.P., Fei, Y.1995. On the flora of the alpine region in the Qinghai-Xizang (Tibet) plateau. Acta Botanical Yunnanica.17:233-250.
Wulf, E.V.1943. An Introduction to Historical Plant Geography. Chronica Botanica, Waltham.
Yasui, Y., Ohnishi, O.1998a. Interspecific relationships in Fagopyrum (Polygonaceae) revealed by the nucleotide sequences of the rbcL and accD genes and their intergenic region. American Journal of Botany 85:1134-1142.
Yasui, Y, Ohnishi, O.1998b. Phylogenetic relationships among Fagopyrum species revealed by the nucleotide sequences of the ITS region of the nuclear rRNA gene. Genes and Genetic Systems 73:201-210.
Yue, J.P., Gu, Z.J., Ai-Shehbaz, LA., Sun, H.2004. Cytological studies on the Sino-Himalayan endemic Solms-laubachia (Brassicaceae) and two related genera. Botanical Journal of Linnean Society 145:77-86.
Zhang, J.W., Sun, H., Nie, Z.L.2007. Karyological studies on the Sino-Himilayan endemic Soroseris and two related genera of tribe Lactrceae (Asteraceae). Botanical Journal of Linnean Society 154:79-87.
Zhang, J.W., Nie, Z.L., Wen, J., Sun, H.2010. Molecular phylogeny and biogeography of three closely related genera, Soroseris, Stebbinsia, and Syncalathium (Asteraceae, Cichorieae), endemic to the Tibetan Plateau, SW China. Taxon 60:15-26.
Zhang, X.S.1983. The Tibetan Plateau in relation to the vegetation of China. Annals of Missouri Botanical Garden 70:564-570.
Zurawski, G., Clegg, M.T.1987. Evolution of higher-plant chloroplast DNA -coded genes: Implications for structure-function, Phylogenetic studies. Annual Review of Plant Physiology 38:391-418.
References
Ballard, H.E., Sytsma, K.J.,2000. Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae):arctic origins, herbaceous ancestry and bird dispersal. Evolution 54, 1521-1532.
Bao, B.J., Li, A.R.,1993. A study of the genus Atraphaxis in China and the system of Atraphaxideae (Polygonaceae). Acta Phytotax. Sin.31 (2),127-139.
Bohle, U.R., Hilger, H.H., Martin, W.F.,1996. Island colonization and evolution of the insular woody habit in Echium L. (Boraginaceae). Proc. Natl. Acad. Sci. USA 93,11740-11745.
Carlquist, S.,1970. Hawaii, a natural history:geology, climate, native flora and fauna above the shoreline. Pacific Tropical Botanical Garden, Lawai, HI.
Carlquist, S.,1974. Island biology. Columbia Univ. Press, New York.
Carlquist, S.,2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Bot. J. Linn. Soc.141,25-51.
Chen, Q.F.,1999. A study of resources of Fagopyrum (Polygonaceae) native to China. Bot. J. Linn. Soc.130,53-64.
Darwin, C.,1859. The Origin of Species by Means of Natural Selection. John Murray, London.
Doyle, J.J., Doyle, J.L.,1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull.19,11-15.
Drummond, A.J., Ho, S.Y., Rawlence, N., Rambaut, A.,2007. A rough guide to BEAST 1.4. University of Auckland, Auckland, New Zealand.
Farris, J.S., Kallersjo, M., Kluge, A.G., Bult, C.,1995. Testing significance of incongruence. Cladistics 10,315-319.
Felsenstein, J.,1985. Confidence limits on phylogenies:an approach using the bootstrap. Evolution 39,783-791.
Friesen. N., Fritsch, R.M., Pollner. S., Blattner. F.R.,2000. Molecular and morphological evidence for an origin of the aberrant genus Milula within Himalayan species of Allium (Alliaceae). Mol. Phylogenet. Evol.17,209-218.
Givnish, T.J.,1998. Adaptive radiation of plants on oceanic islands:classical patterns, molecular data, new insights. Pp.281-304 in P. Grant, ed. Evolution on islands. Oxford Univ. Press, Oxford, U.K.
Givnish, T.J., Kendra, M.C., Mast, A.R., Paterson, T.B., Theim, T.J., Hipp, A.L., Henss, J.M., Smith, J.F., Kenneth, R., Wood, K.R., Sytsma, K.J.,2009. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales:Campanulaceae). Proc. Roy. Soc. B 276, 407-416.
Guindon, S., Cascuel, O.,2003. A simple, fast, and accurate algorihm to estimate large phylogenies by maximum likelihood. Syst. Biol.52,696-704.
Harrison, TM, Copeland, P, Kidd, WSF, Yin, A.1992. Raising Tibet. Science 255,1663-1670.
Hong, S.P.,1995. Pollen morphology of Parapteropyrum and some putatively related genera (Polygonaceae- Atraphaxideae). Grana 34,153-159
Hong, S.P.,1998. Systematic significance of tepal surfac morphology in tribes Persicarieae and Polygoneae (Polygonaceae). Bot. J. Linn. Soc.127,91-116.
Huelsenbeck, J.P., Rannala, B.,1997. Phylogenetic Methods Come of Age:Testing Hypotheses in an Evolutionary Context. Science 276,227-232.
Huelsenbeck, J.P., Ronquist, F.,2001. MrBayes:Bayesian inference of phygeny. Bioinformatics 17,754-755.
Hughes, C., Eastwood, R.,2006. Island radiation on a continental scale:exceptional rates of plant diversification after uplift of the Andes. Proc. Natl. Acad. Sci. USA 103,10 334-10 339.
Kadereit, J.W.,1994. Molecules and morphology, phylogenetics and genetics. Bot. Acta.107, 369-373.
Lamb-Frye, A.S., Kron, K.A.,2003. rbcL phylogeny and character evolution in Polygonaceae. Syst. Bot.28(2),326-332.
Li, A.R.,1981. Parapteropyrum A. J. Li—Unum genus novum Polygonacearum sinicum. Acta Phytotax. Sin.19,330-331.
Li. A.R.,1998. Flora Republicae popularis Sinicae. Beijing:Science Press.
Li, J.J., Shi, Y.F., Li. B.Y.,1995. Uplift of the Qinghai-Xizang (Tibet) Plateau and Global Change. Lanzhou:Lanzhou University Press.
Liedo, M.D., Crespo, MB., Cameron. K.M., Fay, M.F., Chase, M.W.,1998. Systematics of Plumbaginaceae based upon cladistic aanlysis of rbcL sequence data. Syst. Bot.23,21-29.
Liu. J.Q., Chen, Z.D., Lu, A.M.,2000. The phylogenetic relationships of the Qinghai-Tibet endemic Sinadoxa, revealed by the ITS data. Acta Phytotax. Sin.42,656-658.
Liu, J.Q., Wang, Y.J., Wang, A.L., Hideaki, O., Abbott, R.J.,2006. Radiation and diversification within the Ligularia-Cremanlhodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau. Mol. Phylogenet. Evol.38,31-49.
Mao, Z.M., Yang. G., Wang C.G.,1983. Studies on chromosome number and anatomy of young branches of Calligonum of Xinjiang in relation to the evolution of some species of the genus. Acta Phytotaxon. Sin.21,44-48.
Ohnishi, O., Matsuoka, Y.,1996. Search for the wild ancestor of buckwheat Ⅱ. Taxonomy of Fagopyrum (Polygonaceae) species based on morphology, isozymes and cpDNA variability. Gene & Genet. Syst.71,383-390.
Ohnishi, O.,1998. Search for the wild ancestor of buckwheat I. Description of new Fagopyrum (Polygonaceae) species and their distribution in China and the Himalayan hills. Fagopyrum 15, 18-28.
Ohsako, T., Ohnishi, O.,2000. Intra- and interspecific phylogeny of wild Fagopyrum (Polygonaceae) species based on nucleotide sequences of noncoding regions in chloroplast DNA. Am. J. Bot.87,573-582.
Panero, J.L., Francisco-Ortega, J., Jansen, R.K., Santos-Guerra, A.,1999. Molecular evidence for multiple origins of woodiness and a New World biogeographic connection of the Macaronesian Island endemic Pericallis (Asteraceae:Senecioneae). Proc. Natl. Acad. Sci. USA 96 (24),13886-13891.
Posada, D., Crandall, K.A.,1998. Modeltest:testing the model of DNA substitution. Bioinformatics 14,817-81
Richardson, J.E., Pennington, R.T., Pennington, T.D., Hollingsworth, P.M.,2001. Rapid diversification of a species-rich genus of neotropical rain forest trees. Science 293,2242-2245.
Sanchez, A., Schuster, T.M., Kron, K.A.,2009. A large-scale phylogeny of Polygonaceae based on molecular data. Int J. Plant Sci.1708,1044-1055.
Shi. Y.F., Li, J.J., Li, B.Y.,1998. Uplift and environmental changes of Qinghai-Tibetan Plateau in the late Cenozoic. Guangzhou:Guangdong Science and Technology Press.
Spicer, RA, Harris, NBW, Widdowson, M, Herman, AB, Guo, S, Valdes, PJ, Wolfe, JA, Kelley, SP. 2003. Constant elevation of southern Tibet over the past 15 million years. Nature 421, 622-624.
Sun, H.,2002. Tethys retreat and Himalayas-Hengduanshan Mountains uplift and their significance on the origin and development of the Sino-Himalayan elements and alpine flora. Acta Bot. Yunnan.24,273-288.
Swofford, D.L.,2002. paup*:Phylogenetic Analyses Using Parsimony (* and Other Methods), Version 4. Sinauer & Associates, Sunderland, MA.
Tamura, K., Dudley, J., Nei, M., Kumar, S.,2007. MEGA4:Molecular Evolutionary Genetics Analysis (MEGA) v755 software version 4.0. Mol. Biol. Evol.24,1596-1599.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.,2008. Morphological cladistic analysis of Calligonum and Pteropyrum (Polygonaceae) in Iran. Iran. J. Biotechnol.14,117-125.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.,2010. The phylogeny of Calligonum and Pteropyrum (Polygonaceae) based on nuclear ribosomal DNA ITS and chloroplast trnL-F sequences. Iran. J. Biotechnol.8,7-15.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F., Higgins, D.G.,1997. Clustal-X windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucl. Acids Res.25,4876-4882.
Tian, X.M., Liu, R.R., Tian, B., Liu, J.Q.,2009. Karyological studies of Parapteropyrum and Atraphaxis (Polygonaceae). Caryologia 62 (4),261-266.
Tu, T.Y., Volis, S., Dilon, M.O., Sun, H., Wen, J.,2010. Dispersals of Hyoscyameae and Mandragoreae (Solanaceae) from the New World to Eurasia in the early Miocene and their biogeographic diversification within Eurasia. Mol. Phylogenet. Evol.57,1226-1237.
Wallace, A.R.,1878. Tropical Nature and Other Essays. Macmillan, London.
Wang, A.L., Yang, M.H., Liu, J.Q.,2005. Molecular phylogeny, recent Radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequence. Ann. Bot.96,489-498.
Wang, Y.J., Susanna, A., Von, Raab-Straube., Milne, R., Liu, J.Q.2009. Island-like radiation of Saussurea (Asteraceae:Cardueae) triggered by uplift of the Qinghai-Tibet Plateau. Biol. J. Linn. Soc.97,893-903.
Wu, Y.,2004. The floristic characteristics in the region of Bayan Har Mountains. Acta Bot. Yunnan.26,587-603.
Yasui, Y., Ohnishi, O.,1998a. Interspecific relationships in Fagopyrum (Polygonaceae) revealed by the nucleotide sequences of the rbcL and accD genes and their intergenic region. Am. J. Bot.85,1134-1142.
Yasui, Y., Ohnishi, O.,1998b. Phylogenetic relationships among Fagopyrum species revealed by the nucleotide sequences of the ITS region of the nuclear rRNA gene. Genes Genet. Syst.73, 201-210.
Zhang, J.W., Nie,Z.L., Wen, J., Sun, H.,2011. Molecular phylogeny and biogeography of three closely related genera Soroseris, Stebbinsia,and Syncalathium (Asteraceae, Cichorieae) endemic to the Tibetan Plateau, SW China. Taxon 60,1-12.
李安仁.1998.中国植物志第25卷第1册,科学出版社
李吉均,施雅风,李炳元.1995.青藏高原隆升与全球变化.甘肃人民出版社,兰州
李吉均,方小敏.1998.青藏高原隆起与环境变化研究。《科学通报》,年,43(15)：1569-1574.
施雅风,李吉均,李炳元.1998.青藏高原晚新生代隆升与环境变化.广东：广东科技出版社
潘保田,李吉钧.1996.青藏高原：全球气候变化的驱动机与放大器,Ⅲ.青藏高原隆起对气候变化的影响.兰州大学学报.32：108-115.
吴征镒.1987.西藏植物区系的起源及其演化.北京,科学出版社.5：874-902.
李文华,周兴民.1998.青藏高原生态系统及优化利用模式.广东：广东科技出版社.
Baltisberger M., Widmer A.2006. Chromosome numbers of plant species from the Canary Islands. Botanica Helvetica 116:9-30.
Bao, B.J., Li, A.R.1993. A study of the genus Atraphaxis in China and the system of Atraphaxideae (Polygonaceae). Acta Phytotaxonomica. Sinica 31 (2); 127-139.
Beerling, D.J., Bailey, J.P., Conolly, A.P.1994. Fallopia japonica (Houtt.) Ronse Decraene. Journal of Ecology 82:959-979.
Ballard, H.E., Sytsma, K.J.2000. Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae):Arctic origins, herbaceous ancestry and bird dispersal. Evolution 54: 1521-1532.
Brandbyge, J.1993. Polygonaceae. In:K. Kubitzki, J.C. Rohwer, and V. Bittrich, eds The families and genera of vascularplants, Springer-Verlag, Berlin.2:531-544.
Bohle, U.R., Hilger, H.H., Martin, W.F.1996. Island colonization and evolution of the insular woody habit in Echium L. (Boraginaceae). Proceedings of the National Academy of Sciences of the United States of America 93:11740-11745.
Carlquist, S.1970. Hawaii, a natural history:geology, climate, native flora and fauna above the shoreline. Pacific Tropical Botanical Garden, Lawai, HI.
Carlquist, S.1974. Island biology. Columbia Univ. Press, New York.
Carlquist, S.2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Journal of Linnean Society 141:25-51.
Chen, Q.F.1999. A study of resources of Fagopyrum (Polygonaceae) native to China. Botanical Journal of Linnean Society 130:53-64.
Chung, S.L., Lo, C.H., Lee, T.Y., Zhang, Y.Q., Xie, Y.W., li, X.H., Wang, P.L.1998. Diachronomous uplift of the Tibetan plateau starting 40 Myr ago. Nature 394:769-773.
Cui, X.K., Ao, C.Q., Zhang, Q., Chen, L.T., Liu, J.Q.2008. Diploid and tetraploid distribution of Allium przewalskianum Regel. (Liliaceae) at the regional scale:implications for cryptic speciation through intraspecific polyploidization. Caryologia 61(2):192-200.
Darwin, C.1859. The Origin of Species by Means of Natural Selection.John Murray, London.
Doyle, J.J., Doyle, J.L.1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19:11-15.
Drummond, A.J., Ho, S.Y., Rawlence, N., Rambaut, A.2007. A rough guide to BEAST 1.4. University of Auckland, Aukland, New Zealand.
Carlquist, S.2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Botanical Journal of Linnean Society 141:25-51.
Farris, J.S., Kallersjo, M., Kluge, A.G., Bult, C.1995. Testing signifcance of incongruence. Cladistics 10:315-319.
Fay, M.F., Swensen, S.M., Chase, M.W.1997. Taxonomic affinities of Medusagyne oppositifolia (Medusagynaceae). Kew Bull.52:111-120. Academic Press, New York.
Felsenstein, J.1985. ConWdence limits on phylogenies:an approach using the bootstrap. Evolution 39:783-791.
Friesen, N., Fritsch, R.M., Pollner, S., Blattner, F.R.2000. Molecular and morphological evidence for an origin of the aberrant genus Milula within Himalayan species of Allium (Alliaceae). Molecular Phylogenetics and Evolution 17:209-218.
Grant, V.1981. Plant speciation,2nd edn. Columbia University Press, New York.
Givnish, T.J.1998. Adaptive radiation of plants on oceanic islands:classical patterns, molecular data, new insights. Pp.281-304 in P. Grant, ed. Evolution on islands. Oxford Univ. Press, Oxford, U.K.
Givnish, T.J., Kendra, M.C., Mast, A.R., Paterson, T.B., Theim, T.J., Hipp, A.L., Henss, J.M., Smith, J.F., Kenneth, R., Wood, K.R., Sytsma, K.J.2009. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales:Campanulaceae). Proceeding of the Royal Society 276:407-416.
Gross, H.1913. Bertrage ur der Polygonaceae. Botanische Jahrbucher fur Systematik 49: 234-248.
Guindon, S., Cascuel, O.2003. A Simple, Fast, and Accurate Algorihm to Estimate Large Phylogenies by Maximum Likelihood. Sysematic Biology 52:696-704.
Hanldson, K.1978. Anatomy and taxonomy in Polygonaceae subfam. Polygonoideae hleisn. emend. Jaretzky. Symbolae Botanicae Upsaliensis 22:1-95.
Jaretzky, R.1925. Beitriige zur Systematik der Polygonaceae unter Berucksichtigung des Oxymethyl-anthrachinon-Vorkommens. Feddes Repertorium Specierum Novarum Regni Vegetabilis 22:49-83.
Hedberg, O.1997. The genus Koenigia L. emend. Hedberg (Polygonaceae). Botanical Journal of Linnean Society 124:295-330.
Hong, D.Y.1990. Plant cytotaxonomy. Science Press, Beijing.
Hong, S.P.1995. Pollen morphology of Parapteropyrum and some putatively related genera (Polygonaceae- Atraphaxideae). Grana 34:153-159.
Hong, S.P.1998. Systematic significance of tepal surfac morphology in tribes Persicarieae and Polygoneae (Polygonaceae). Journal of Linnean Society 127:91-116.
Hu, Y.P., Xie, X.L., Wen, Q., Zhao, X.D., Wang, L., Li, Y.2007. Studies on karyotype of five polulations of Rheum tanguticum. Acta Botanica Yunnanica 29(4):429-433.
Huelsenbeck, J.P., Rannala, B.1997. Phylogenetic Methods Come of Age:Testing Hypotheses in an Evolutionary Context. Science 276:227-232.
Huelsenbeck, J.P, Ronquist, F.2001. MrBayes:Bayesian inference of phygeny. Bioinformatics 17: 754-755.
Hughes, C., Eastwood, R.2006. Island radiation on a continental scale:exceptional rates of plant diversification after uplift of the Andes. Proceedings of the National Academy of Sciences of the United States of America 103:10334-10339.
Kadereit. J.W.1994. Molecules and morphology, phylogenetics and genetics. Botanica Acta 107: 369-373.
Kong, H.Z., Liu, J.Q.1999. Karyomorphology of the genus Pomatosace Maxim. (Primulaceae). Acta Phytotaxonomica Sinica 37:445-450.
Lamb-Frye, A.S., Kron, K.A.2003. rbcL phylogeny and character evolution in Polygonaceae. Systematic Botany.28(2):326-332.
Li, A.R.1981. Parapteropyrum A. J. Li—Unum genus novum Polygonacearum sinicum. Acta Phytotaxonomica Sinica 19:330-331.
Li, J.J., Shi, Y.F.. Li, B.Y.1995. Uplift of the Qinghai-Xizang (Tibet) Plateau and Global Change. Lanzhou:Lanzhou University Press.
Liedo, M.D., Crespo, M.B., Cameron, K.M., Fay, M.F., Chase. M.W.1998. Systematics of Plumbaginaceae Based upon Cladistic Aanlysis of rbcL Sequence Data. Systematic Botany 23:21-29.
Liu, J.Q., Zhou, G.Y., Ho, T.N., Lu, A.M.1999. Karyomorphology of Sinadoxa and its systematic significance. Caryologia 52:159-164.
Liu, J.Q.2000. Karyomorphology of 4 species in Sinacalia and Parasenecio (Astetaceae: Senecioneae). Acta Botanical Yunnanica 22:447-450.
Liu, J.Q.. Chen, Z.D., Lu. A.M.2000. The phylogenetic relationships of the Qinghai-Tibet endemic Sinadoxa, revealed by the ITS data. Acta Phytotaxonomica Sinica 42:656-658.
Liu, J.Q., Ho, T.N., Chen, S.L.2002. The first chromosome data documentations of Megacodon and Lomatogoniopsis and their systematic significance. Acta Biologica Plateau Sinica 15: 41-47.
Liu, J.Q., Gao, T.G., Chen, Z.D., Lu, A.M.2002. Molecular phylogeny and biogeography of the Qinghai-Tibet Plateau endemic Nannoglottis (Asteraceae). Molecular Phylogenetics and Evolution 23:307-325.
Liu, J.Q.2004. Uniformity of karyotypes in Ligularia (Asteraceae:Senecioneae), a highly diversified genus of the eastern Qinghai-Tibet Plateau highlands and adjacent areas. Botanical Journal of Linnean Society 144:329-342.
Liu, J.Q., Wang, Y.J., Wang, A.L., Hideaki, O., Abbott, R.J.2006. Radiation and diversification within the Ligularia-Cremanthodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution 38:31-49.
Liu, J.Q., Tian, B.2007. Origin, evolution, and systematics of Himalaya endemic genera. Newsletters of Himalayan Botany 40:20-27.
Mao, Z.M., Yang, G., Wang, C.G.1983. Studies on chromosome number and anatomy of young branches of Calligonum of Xinjiang in relation to the evolution of some species of the genus. Acta Phytotaxonomica Sinica 21:44-48.
Mao, K.S., Hao, G., Liu, J.Q., Adams, R.P., Milne, R.I.2010. Diversification and biogeography of Juniperus (Cupressaceae):variable diversification rates and multiple intercontinental dispersals. New Phytologist 188:254-272.
Maude, P.F.1939. A list of the chromosome numberals of species of British flowering plants. New Phytologist 38:1-31.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., Fonseca, G.A.B., Kent, J.2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.
Nair, P.K., Rehman, K., Saxena, A.K.1976. Contribution to the pollen morphology of Indian Polygonaceae. Journal of Palynotogy 12:1-18.
Ni, Z.C., Cheng, S.Z.1987. On genera endemic to China and occurring on the Xizang Plateau flora. Acta Phytotaxonomica Sinica 25:416-429.
Nie, Z.L., Gu, Z.J., Sun, H.2002. Cytological study of genus Tibetia (Fabaceae) in the Hengduan Mountains region, China. Journal of Plant Research 115:17-22.
Nie, Z.L., Wen, J., Gu, Z.J., Boufford, D.E., Sun, H.2005. Polyploidy in the flora of the Hengduan mountains hotspot, southwestern China. Annals of the Missouri Botanical Garden 92:275-306.
Nowicke, J.V., Skvarla, J.J.1977. Pollen morphology amd the relationship of the Plumbaginaceae, Polygonaceae and Primulaceae to the order Centrospermae. Smithsonian Contributions to Botany 37:1-64.
Nowicke, J.V., Skvarla, J.J.1979. Pollen morphology:The potential influence in higher order systematics. Annals of the Missouri Botanical Garden 66:633-700.
Ohba, H.,1988. The alpine flora of the Nepal Himalayas:An introductory note. In:Ohba H, Malla SB (Eds.), The Himalayas Plants, vol.1. Tokyo:University of Tokyo Press,19-46.
Ohnishi, O., Matsuoka, Y.1996. Search for the wild ancestor of buckwheat Ⅱ. Taxonomy of Fagopyrum (Polygonaceae) species based on morphology, isozymes and cpDNA variability. Gene & Genetic Systems 71:383-390.
Ohnishi, O.1998. Search for the wild ancestor of buckwheat I. Description of new Fagopyrum (Polygonaceae) species and their distribution in China and the Himalayan hills. Fagopyrum 15:18-28.
Ohsako, T., Ohnishi, O.2000. Intra- and interspecific phytogeny of wild Fagopyrum (Polygonaceae) species based on nucleotide sequences of noncoding regions in chloroplast DNA. American Journal of Botany 87:573-582.
Raven, P.H.1975. The bases of angiospenn phylogeny:cytology. Annals of the Missouri Botanical Garden 62:724-764.
Panero, J.L., Francisco-Ortega, J., Jansen, R.K., Santos-Guerra, A.1999. Molecular evidence for multiple origins of woodiness and a New World biogeographic connection of the Macaronesian Island endemic Pericallis (Asteraceae:Senecioneae). Proceedings of the National Academy of Sciences of the United States of America 96 (24):13886-13891.
Posada, D., Crandall, K.A.1998. Modeltest:testing the model of DNA substitution. Bioinformatics 14:817-818.
Pryer, K.M., Schneider, H., Smith, A.R., Cranfill, R., Wolf, P.G., Hunt, J.S., Sipes, S.D.2001 Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Nature 409:618-622.
Ramsey, J.2011. Polyploidy and ecological adaptation in wild yallow. Proceedings of the National Academy of Sciences of the United States of America 108(17):7096-7101.
Rambaut, A., Drummond, A.J.2010. Tracer v 1.5. Availableat:http://beast.bio.ed.ac.uk/Tracer (ac cessed 10 January 2010).
Roberty, G., Vaurier, S.1964. Les genres de Polygonacees. Boissien 10:7-128.
Richardson, J.E., Pennington, R.T., Pennington, T.D., Hollingsworth, P.M.2001. Rapid diversification of a species-rich genus of neotropical rain forest trees. Science 293:2242-2245.
Sanchez, A., Kron, K.A.2008. Phylogenetics of Polygonaceae with an Emphasis on the Evolution of Eriogonoideae. Systematic Botany 33 (1):87-96.
Sanchez, A., Kron, K.A.2008. Preliminary phylogenetic relationships of Calligonum L. (Polygonaceae) using two chloroplast genes. Botany conference, Abstract ID:98. Website:http://www.2008.botanyconference.org/engine/search/index.php?func=detail&aid= 98.
Sanchez, A., Schuster, T.M., Kron, K.A.2009. A large-scale phylogeny of polygonaceae based on molecular data. International Journal of Plant Sciences 1708:1044-1055.
Saiki, R.K., Celfand, D.H., Stoffel, S., Scharf, S.J., Higuchi, R., Horn, G.T., Mullis, K.B., Erlich, H.H.A.1988. Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase. Science 239:487-491
Shi, Y.F., Li, J.J., Li. B.Y.1998. Uplift and environmental changes of Qinghai-Tibetan Plateau in the late Cenozoic. Guangzhou:Guangdong Science and Technology Press.
Swofford, D.L.2002. paup*:Phylogenetic Analyses Using Parsimony (* and Other Methods), Version 4. Sinauer & Associates, Sunderland, MA.
Soltis, D., Soltis, P., Jennifer, A.T.2003. Advances in the study of polyploidy since Plant speciation. New Phytologist 161:173-191.
Stebbing, G.L.1971. Chromosome evolution in higher plants. Edward Arnold, London.
Stuessy, T.F.1998. Evolution and speciation of island plants.Cambridge University Press.
Simmonds, N.W.1945. Biological flora of the British Isles. Journal of Ecology 33(1):117-120.
Sun, W., Zhou, Z.Z., Liu, M.Z., Wan, H.W., Dong, X.2008. Reappraisal of the generic status of Pteroxygonum (Polygonaceae) on the basis of morphology, anatomy and nrDNA ITS sequence analysis. Journal of Systematics and Evolution 46 (1):73-79.
Tang, H.G., Meng, L.H., Ao, C.Q., Liu, J.Q.2005. Origin of the Qinghai-Tibetan Plateau endemic Milula (Liliaceae):further insights from karyological comparisons with Allium. Caryologia 58:320-331.
Tamura, K., Dudley, J., Nei, M., Kumar, S.2007. MEGA4:Molecular Evolutionary Genetics Analysis (MEGA) v755 software version 4.0. Molecular Phylogenetics and Evolution 24: 1596-1599.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.2008. Morphological cladistic analysis of Calligonum and Pteropyrum (Polygonaceae) in Iran. Iranian Journal of Biotechnology 14: 117-125.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.2010. The phylogeny of Calligonum and Pteropyrum (Polygonaceae) based on nuclear ribosomal DNA ITS and chloroplast trnL-Fsequences. Iranian Journal of Biotechnology 8:7-15.
Tapponnier, P., Xu, Z.Q.. Roger, F., Meyer, B., Arnaud, N., Wittlinger, G., Yang J.S.2001. Oblique stepwise rise and growth of the Tibet Plateau. Science 294:1671-1677.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F., Higgins, D.G.1997. Clustal-X windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic acids research 25:4876-4882.
Tsukaya, H., Tsuge. T.2001. Morphological adaptation of inflorescences in plants that bloom atlow temperature in early spring:the convergent evolution of downy plants. Plant Biology 3:536-543.
Tsumura, H.2002. Optical and anatomical characteristics of bracts from the Chinese "glasshouse" plant, Rheum alexandrae Batalin (Polygonaceae), Yunnan, China. Journal of Plant Research 115:59-63.
Tu, T.Y, Sun, H., Gu, Z.J., Yue, J.P.2005. Cytological studies on the Sino-Himalayan endemic Anisodus and four related genera from the tribe Hyoscyameae (Solanaceae), and their systematic and evolutionary implications. Botanical Journal of Linnean Society 147: 457-468.
Tu, T.Y., Volis, S., Dillon, M.O., Sun, H., Wen, J.2010. Dispersals of Hyoscyameae and Mandragoreae (Solanaceae) from the New World to Eurasia in the early Miocene and their biogeographic diversification within Eurasia. Molecular Phylogenetics and Evolution 57: 1226-1237.
Wallace AR.1878. Tropical Nature and Other Essays. Macmillan, London.
Wang, A.L., Yang, M.H., Liu, J.Q.2005. Molecular phylogeny, recent Radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequence. Annals of Botany 96:489-498.
Wang, Y.J., Susanna, A., Von Raab-Straube, E., Milne, R., Liu, J.Q.2009. Island-like radiation of Saussurea (Asteraceae:Cardueae) triggered by uplift of the Qinghai-Tibet Plateau. Biological Journal of the Linnean Society 97:893-903.
White, T.J., Birns, T., Lee, S., Taylor, J.1990. Amplification and direct sequencing of fungal ribosomal RNA genes fro phylogenetics. Pp.315-322 in M. A. Innis, D. H. Gelfand, J. J.Sninsky, and T. J. White, eds. PCR Protocols:A Guide to Methods and Applications.
Wu, C.Y.1987. Flora of Tibet, Vol.5. Science Press, Beijing.
Wu, C.Y.1988. Hengduan Mountains flora and her sighificane. Journal of Japanese Botany 63: 297-311.
Wu, S.G., Yang, Y.P., Fei, Y.1995. On the flora of the alpine region in the Qinghai-Xizang (Tibet) plateau. Acta Botanical Yunnanica.17:233-250.
Wulf, E.V.1943. An Introduction to Historical Plant Geography. Chronica Botanica, Waltham.
Yasui, Y., Ohnishi, O.1998a. Interspecific relationships in Fagopyrum (Polygonaceae) revealed by the nucleotide sequences of the rbcL and accD genes and their intergenic region. American Journal of Botany 85:1134-1142.
Yasui, Y, Ohnishi, O.1998b. Phylogenetic relationships among Fagopyrum species revealed by the nucleotide sequences of the ITS region of the nuclear rRNA gene. Genes and Genetic Systems 73:201-210.
Yue, J.P., Gu, Z.J., Ai-Shehbaz, LA., Sun, H.2004. Cytological studies on the Sino-Himalayan endemic Solms-laubachia (Brassicaceae) and two related genera. Botanical Journal of Linnean Society 145:77-86.
Zhang, J.W., Sun, H., Nie, Z.L.2007. Karyological studies on the Sino-Himilayan endemic Soroseris and two related genera of tribe Lactrceae (Asteraceae). Botanical Journal of Linnean Society 154:79-87.
Zhang, J.W., Nie, Z.L., Wen, J., Sun, H.2010. Molecular phylogeny and biogeography of three closely related genera, Soroseris, Stebbinsia, and Syncalathium (Asteraceae, Cichorieae), endemic to the Tibetan Plateau, SW China. Taxon 60:15-26.
Zhang, X.S.1983. The Tibetan Plateau in relation to the vegetation of China. Annals of Missouri Botanical Garden 70:564-570.
Zurawski, G., Clegg, M.T.1987. Evolution of higher-plant chloroplast DNA -coded genes: Implications for structure-function, Phylogenetic studies. Annual Review of Plant Physiology 38:391-418.
References
Ballard, H.E., Sytsma, K.J.,2000. Evolution and biogeography of the woody Hawaiian violets (Viola, Violaceae):arctic origins, herbaceous ancestry and bird dispersal. Evolution 54, 1521-1532.
Bao, B.J., Li, A.R.,1993. A study of the genus Atraphaxis in China and the system of Atraphaxideae (Polygonaceae). Acta Phytotax. Sin.31 (2),127-139.
Bohle, U.R., Hilger, H.H., Martin, W.F.,1996. Island colonization and evolution of the insular woody habit in Echium L. (Boraginaceae). Proc. Natl. Acad. Sci. USA 93,11740-11745.
Carlquist, S.,1970. Hawaii, a natural history:geology, climate, native flora and fauna above the shoreline. Pacific Tropical Botanical Garden, Lawai, HI.
Carlquist, S.,1974. Island biology. Columbia Univ. Press, New York.
Carlquist, S.,2003. Wood anatomy of Polygonaceae:analysis of a family with exceptional wood diversity. Bot. J. Linn. Soc.141,25-51.
Chen, Q.F.,1999. A study of resources of Fagopyrum (Polygonaceae) native to China. Bot. J. Linn. Soc.130,53-64.
Darwin, C.,1859. The Origin of Species by Means of Natural Selection. John Murray, London.
Doyle, J.J., Doyle, J.L.,1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull.19,11-15.
Drummond, A.J., Ho, S.Y., Rawlence, N., Rambaut, A.,2007. A rough guide to BEAST 1.4. University of Auckland, Auckland, New Zealand.
Farris, J.S., Kallersjo, M., Kluge, A.G., Bult, C.,1995. Testing significance of incongruence. Cladistics 10,315-319.
Felsenstein, J.,1985. Confidence limits on phylogenies:an approach using the bootstrap. Evolution 39,783-791.
Friesen. N., Fritsch, R.M., Pollner. S., Blattner. F.R.,2000. Molecular and morphological evidence for an origin of the aberrant genus Milula within Himalayan species of Allium (Alliaceae). Mol. Phylogenet. Evol.17,209-218.
Givnish, T.J.,1998. Adaptive radiation of plants on oceanic islands:classical patterns, molecular data, new insights. Pp.281-304 in P. Grant, ed. Evolution on islands. Oxford Univ. Press, Oxford, U.K.
Givnish, T.J., Kendra, M.C., Mast, A.R., Paterson, T.B., Theim, T.J., Hipp, A.L., Henss, J.M., Smith, J.F., Kenneth, R., Wood, K.R., Sytsma, K.J.,2009. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales:Campanulaceae). Proc. Roy. Soc. B 276, 407-416.
Guindon, S., Cascuel, O.,2003. A simple, fast, and accurate algorihm to estimate large phylogenies by maximum likelihood. Syst. Biol.52,696-704.
Harrison, TM, Copeland, P, Kidd, WSF, Yin, A.1992. Raising Tibet. Science 255,1663-1670.
Hong, S.P.,1995. Pollen morphology of Parapteropyrum and some putatively related genera (Polygonaceae- Atraphaxideae). Grana 34,153-159
Hong, S.P.,1998. Systematic significance of tepal surfac morphology in tribes Persicarieae and Polygoneae (Polygonaceae). Bot. J. Linn. Soc.127,91-116.
Huelsenbeck, J.P., Rannala, B.,1997. Phylogenetic Methods Come of Age:Testing Hypotheses in an Evolutionary Context. Science 276,227-232.
Huelsenbeck, J.P., Ronquist, F.,2001. MrBayes:Bayesian inference of phygeny. Bioinformatics 17,754-755.
Hughes, C., Eastwood, R.,2006. Island radiation on a continental scale:exceptional rates of plant diversification after uplift of the Andes. Proc. Natl. Acad. Sci. USA 103,10 334-10 339.
Kadereit, J.W.,1994. Molecules and morphology, phylogenetics and genetics. Bot. Acta.107, 369-373.
Lamb-Frye, A.S., Kron, K.A.,2003. rbcL phylogeny and character evolution in Polygonaceae. Syst. Bot.28(2),326-332.
Li, A.R.,1981. Parapteropyrum A. J. Li—Unum genus novum Polygonacearum sinicum. Acta Phytotax. Sin.19,330-331.
Li. A.R.,1998. Flora Republicae popularis Sinicae. Beijing:Science Press.
Li, J.J., Shi, Y.F., Li. B.Y.,1995. Uplift of the Qinghai-Xizang (Tibet) Plateau and Global Change. Lanzhou:Lanzhou University Press.
Liedo, M.D., Crespo, MB., Cameron. K.M., Fay, M.F., Chase, M.W.,1998. Systematics of Plumbaginaceae based upon cladistic aanlysis of rbcL sequence data. Syst. Bot.23,21-29.
Liu. J.Q., Chen, Z.D., Lu, A.M.,2000. The phylogenetic relationships of the Qinghai-Tibet endemic Sinadoxa, revealed by the ITS data. Acta Phytotax. Sin.42,656-658.
Liu, J.Q., Wang, Y.J., Wang, A.L., Hideaki, O., Abbott, R.J.,2006. Radiation and diversification within the Ligularia-Cremanlhodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau. Mol. Phylogenet. Evol.38,31-49.
Mao, Z.M., Yang. G., Wang C.G.,1983. Studies on chromosome number and anatomy of young branches of Calligonum of Xinjiang in relation to the evolution of some species of the genus. Acta Phytotaxon. Sin.21,44-48.
Ohnishi, O., Matsuoka, Y.,1996. Search for the wild ancestor of buckwheat Ⅱ. Taxonomy of Fagopyrum (Polygonaceae) species based on morphology, isozymes and cpDNA variability. Gene & Genet. Syst.71,383-390.
Ohnishi, O.,1998. Search for the wild ancestor of buckwheat I. Description of new Fagopyrum (Polygonaceae) species and their distribution in China and the Himalayan hills. Fagopyrum 15, 18-28.
Ohsako, T., Ohnishi, O.,2000. Intra- and interspecific phylogeny of wild Fagopyrum (Polygonaceae) species based on nucleotide sequences of noncoding regions in chloroplast DNA. Am. J. Bot.87,573-582.
Panero, J.L., Francisco-Ortega, J., Jansen, R.K., Santos-Guerra, A.,1999. Molecular evidence for multiple origins of woodiness and a New World biogeographic connection of the Macaronesian Island endemic Pericallis (Asteraceae:Senecioneae). Proc. Natl. Acad. Sci. USA 96 (24),13886-13891.
Posada, D., Crandall, K.A.,1998. Modeltest:testing the model of DNA substitution. Bioinformatics 14,817-81
Richardson, J.E., Pennington, R.T., Pennington, T.D., Hollingsworth, P.M.,2001. Rapid diversification of a species-rich genus of neotropical rain forest trees. Science 293,2242-2245.
Sanchez, A., Schuster, T.M., Kron, K.A.,2009. A large-scale phylogeny of Polygonaceae based on molecular data. Int J. Plant Sci.1708,1044-1055.
Shi. Y.F., Li, J.J., Li, B.Y.,1998. Uplift and environmental changes of Qinghai-Tibetan Plateau in the late Cenozoic. Guangzhou:Guangdong Science and Technology Press.
Spicer, RA, Harris, NBW, Widdowson, M, Herman, AB, Guo, S, Valdes, PJ, Wolfe, JA, Kelley, SP. 2003. Constant elevation of southern Tibet over the past 15 million years. Nature 421, 622-624.
Sun, H.,2002. Tethys retreat and Himalayas-Hengduanshan Mountains uplift and their significance on the origin and development of the Sino-Himalayan elements and alpine flora. Acta Bot. Yunnan.24,273-288.
Swofford, D.L.,2002. paup*:Phylogenetic Analyses Using Parsimony (* and Other Methods), Version 4. Sinauer & Associates, Sunderland, MA.
Tamura, K., Dudley, J., Nei, M., Kumar, S.,2007. MEGA4:Molecular Evolutionary Genetics Analysis (MEGA) v755 software version 4.0. Mol. Biol. Evol.24,1596-1599.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.,2008. Morphological cladistic analysis of Calligonum and Pteropyrum (Polygonaceae) in Iran. Iran. J. Biotechnol.14,117-125.
Tavakkoli, S., Osaloo, S.K., Maassoumi, A.A.,2010. The phylogeny of Calligonum and Pteropyrum (Polygonaceae) based on nuclear ribosomal DNA ITS and chloroplast trnL-F sequences. Iran. J. Biotechnol.8,7-15.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F., Higgins, D.G.,1997. Clustal-X windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucl. Acids Res.25,4876-4882.
Tian, X.M., Liu, R.R., Tian, B., Liu, J.Q.,2009. Karyological studies of Parapteropyrum and Atraphaxis (Polygonaceae). Caryologia 62 (4),261-266.
Tu, T.Y., Volis, S., Dilon, M.O., Sun, H., Wen, J.,2010. Dispersals of Hyoscyameae and Mandragoreae (Solanaceae) from the New World to Eurasia in the early Miocene and their biogeographic diversification within Eurasia. Mol. Phylogenet. Evol.57,1226-1237.
Wallace, A.R.,1878. Tropical Nature and Other Essays. Macmillan, London.
Wang, A.L., Yang, M.H., Liu, J.Q.,2005. Molecular phylogeny, recent Radiation and evolution of gross morphology of the rhubarb genus Rheum (Polygonaceae) inferred from chloroplast DNA trnL-F sequence. Ann. Bot.96,489-498.
Wang, Y.J., Susanna, A., Von, Raab-Straube., Milne, R., Liu, J.Q.2009. Island-like radiation of Saussurea (Asteraceae:Cardueae) triggered by uplift of the Qinghai-Tibet Plateau. Biol. J. Linn. Soc.97,893-903.
Wu, Y.,2004. The floristic characteristics in the region of Bayan Har Mountains. Acta Bot. Yunnan.26,587-603.
Yasui, Y., Ohnishi, O.,1998a. Interspecific relationships in Fagopyrum (Polygonaceae) revealed by the nucleotide sequences of the rbcL and accD genes and their intergenic region. Am. J. Bot.85,1134-1142.
Yasui, Y., Ohnishi, O.,1998b. Phylogenetic relationships among Fagopyrum species revealed by the nucleotide sequences of the ITS region of the nuclear rRNA gene. Genes Genet. Syst.73, 201-210.
Zhang, J.W., Nie,Z.L., Wen, J., Sun, H.,2011. Molecular phylogeny and biogeography of three closely related genera Soroseris, Stebbinsia,and Syncalathium (Asteraceae, Cichorieae) endemic to the Tibetan Plateau, SW China. Taxon 60,1-12.