中国荞麦属植物分类学的修订
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摘要
荞麦是一种蓼科(Polygonaceae family)荞麦属(Fagopyrum Mill)的古老作物,在全世界,尤其是东亚及东欧地区广泛种植。数千年以来,荞麦都是中国凉山等高海拔地区的主要粮食作物,其营养丰富,其中富含的芦丁等生物活性成分还具有抗氧化、抗肿瘤的功效。荞麦的栽培种甜荞和苦荞已经被广泛传播,但野生荞麦却只在特定区域分布。中国西南地区,包括四川、云南、贵州各省以及西藏自治区,是公认的世界荞麦属植物的起源中心、分布中心和多样化中心,荞麦种质资源非常丰富。目前已有文献报导的荞麦属植物超过20种,其绝大多数已在中国西南地区被发现,该地区的地理环境非常复杂,孕育了极具多样性的野生荞麦资源。但截至目前,这些已报导的种还没有经过科学的归类,同种异名的现象比较严重,本研究团队从2004年至今,在中国西南地区进行了多年多次的野外考察和标本采集工作,并在室内进行花和果实的解剖观察,获得了大量的荞麦属植物分类学数据。基于这些数据和荞麦染色体数量分析,经馆藏标本和模式照片的对比,并综合国内外相关文献,对中国荞麦属植物进行了分类学修订,将产自中国的荞麦属28个种归并为21个:即将Fagopyrum megaspartanium Q. F.Chen,F. pilus Q. F. Chen并入F. cymosum (Trrev)Meisn作为异名;F. pleioramosum Ohnishi,F. wenchuanense J.R.Shao并入F.caudatum (Samuelsson) A. J. Li作为异名;F. callianthum Ohnishi并入F.qiangcai D.Q.Ba作为异名;F. zuogongense Q.F.Chen并入F. homotropicum Ohnishi作为异名;F. longzhoushanense J.R.Shao并入F. pugense T.Yu作为异名。重新制定了21个种的分种检索表。本文将有助于重新认识荞麦属植物的进化地位,并为后续进一步探究属中各个种的亲缘关系奠定了基础。
Buckwheat(Fagopyrum ssp.) is an ancient crop belonging to Fagopyrum Mill,Polygonaceae family,which has been widely cultivated around the world,especially in East Asia and East Europe. Buckwheat is the major food crop in the high-attitude regions,including the Liangshan region of China for thousands of years. Buckwheat is rich in nutrition and considered as a functional food for the high contents of rutin and other bioactive compounds that are antioxidant and anti-tumor. Two species of cultivated buckwheat,common buckwheat(Fagopyrum esculentum) and tartary buckwheat(Fagopyrum tataricum),are widely spread,while the wild buckwheat resources only distribute in some certain regions. Southwest China,including Sichuan,Yunnan,Guizhou Province and Tibet Autonomous Region,is widely known as the center of origin,distribution and diversity of wild buckwheat plants. So far,more than 20 buckwheat species have been named and classi?ed,most of which have been found in Southwest China. The extremely complicated geographical environment of this area renders a number of diversi?ed wild buckwheat resources. However,the scienti?c classi?cation of Fagopyrum Mill species is de?cient,and some wild species are synonyms. Since 2004,our research team continually made field investigation and specimens collection works in Southwest China,anatomic observation of flowers and fruits were also made in our lab. Based on these morphological data as well as buckwheat chromosome number detecting results,comparing with other specimens,model photos and pervious articles,the Fagopyrum Mill was taxonomic revised and the number of species were adjusted from 28 to 21,namely,Fagopyrum megaspartanium Q.F. Chen and F. pilus Q. F. Chen were treated as synonyms of F. cymosum(Trrev)Meisn;F. pleioramosum Ohnishi and F. wenchuanense J.R.Shao were treated as synonyms of F. caudatum(Samuelsson) A. J. Li;F. callianthum Ohnishi was treated as a synonym of F. qiangcai D.Q.Bai;F. zuogongense Q.F.Chen was treated as a synonym of F. homotropicum Ohnishi;F. longzhoushanense J.R.Shao was treated as a synonym of F. pugense T.Yu. The key to species of Fagopyrum were also revised and presented. Our results provided a better understanding of the evolution position of Fagopyrum Mill and built a framework to learn the genetic relationships between buckwheat species.
Buckwheat(Fagopyrum ssp.) is an ancient crop belonging to Fagopyrum Mill,Polygonaceae family,which has been widely cultivated around the world,especially in East Asia and East Europe. Buckwheat is the major food crop in the high-attitude regions,including the Liangshan region of China for thousands of years. Buckwheat is rich in nutrition and considered as a functional food for the high contents of rutin and other bioactive compounds that are antioxidant and anti-tumor. Two species of cultivated buckwheat,common buckwheat(Fagopyrum esculentum) and tartary buckwheat(Fagopyrum tataricum),are widely spread,while the wild buckwheat resources only distribute in some certain regions. Southwest China,including Sichuan,Yunnan,Guizhou Province and Tibet Autonomous Region,is widely known as the center of origin,distribution and diversity of wild buckwheat plants. So far,more than 20 buckwheat species have been named and classi?ed,most of which have been found in Southwest China. The extremely complicated geographical environment of this area renders a number of diversi?ed wild buckwheat resources. However,the scienti?c classi?cation of Fagopyrum Mill species is de?cient,and some wild species are synonyms. Since 2004,our research team continually made field investigation and specimens collection works in Southwest China,anatomic observation of flowers and fruits were also made in our lab. Based on these morphological data as well as buckwheat chromosome number detecting results,comparing with other specimens,model photos and pervious articles,the Fagopyrum Mill was taxonomic revised and the number of species were adjusted from 28 to 21,namely,Fagopyrum megaspartanium Q.F. Chen and F. pilus Q. F. Chen were treated as synonyms of F. cymosum(Trrev)Meisn;F. pleioramosum Ohnishi and F. wenchuanense J.R.Shao were treated as synonyms of F. caudatum(Samuelsson) A. J. Li;F. callianthum Ohnishi was treated as a synonym of F. qiangcai D.Q.Bai;F. zuogongense Q.F.Chen was treated as a synonym of F. homotropicum Ohnishi;F. longzhoushanense J.R.Shao was treated as a synonym of F. pugense T.Yu. The key to species of Fagopyrum were also revised and presented. Our results provided a better understanding of the evolution position of Fagopyrum Mill and built a framework to learn the genetic relationships between buckwheat species.
引文
[1]Ye N G, Guo G Q. Classification, origin and evolution of genus Fagopyrum in China. Taiyuan:Agricultural Publishing House,1992:19-28
[2]李安仁·中国植物志:第25分册,寥科·北京:科学出版社,1998:108-117Li A R. Flora of China:Volume 25, Polygonaceae. Beijing:Science Press, 1998:108-117
[3] Ohnishi O.Discovery of new Fagopyrum species and its implication for the studies of evolution of Fagopyrum and of the origin of cultivated buckwheat. Japan:Shinshu University Press, 1995:175-190
[4] Ohnishi O-Search for the wild ancestor of buckwheat I Description of new Fagopyrum(Polygonaceae)species and their distribution in China. Fagopyrum, 1998, 15:18-28
[5] Ohsako T, Ohnishi O. New Fagopyrum species revealed by morphological and molecular analyses. Genes&Genetic Systems, 1998,73:85-94
[6] Ohsako T, Yamane K, Ohnishi O. Two new Fagopyrum(polygonaceae)species F.gracilipedoides and F. jinshaense from Yunnan-Genes&Genetic Systems, 2002, 77:399-408
[7] Chen Q F. A study of resources of Fagopyrum(Polygonaceae)native to China. Botanical Journal of Linnean Society, 1999,130(1):54-65
[8]刘建林,唐宇,夏明忠,邵继荣,蔡光泽,罗强,孙俊秀.中国四川寥科荞麦属一新种——皱叶野荞麦·植物分类学报,2008,46(6):929-932Liu J L, Tang Y, Xia M Z, Shao J R, Cai G Z, Luo Q, Sun J X. Fagopyrum crispatifolium J. L-Liu, a new species of Polygonaceae from Sichuan, China. Journal of Systematics and Evolution, 2008,46(6):929-932
[9] Tang Y, Zhou M L, Bai D Q, Shao J R, Zhu X M, Wang D Z, Tang Y X-Fagopyrum pugense(Polygonaceae), a new species from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2010, 20(2):239-242
[10] Shao J R, Zhou M L, Zhu X M, Wang D Z, Bai D Q.Fagopyrum wenchuanense and Fagopyrum qiangcai, Two new species of Polygonaceae from Sichuan, China. Novon, 2011,21(2):256-261
[11] Hou L L, Zhou M L, Zhang Q, Qi L P, Yang B X, Tang Y,Zhu X M, Shao J R-Fagopyrum luojishanense, a New Species of Polygonaceae from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2015,24(1):22-26
[12] Zhou M L, Zhang Q, Zheng Y D, Tang Y, Li F L, Zhu X M,Shao J R-Fagopyrum hailuogouense(Polygonaceae), one new species of Polygonaceae from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2015,24(2):222-224
[13] Wang C L, Li Z Q,Ding M Q,Tang Y U,Zhu X,Liu JFagopyrum longzhoushanense, a new species of Polygonaceae from Sichuan, China. Phytotaxa, 2017, 291(1):73-80
[14] Zhou M L, Kreft I, Suvorova G, Tang Y, Woo S H-Buckwheat germplasm in the world. ELSEVIER, San Diego, USALondon:Academic Press, 2018:61-95
[15] Miller P H. The gardeners dictionary. Abridged(edition 4)·London, 1754
[16] Meisner C F. Monographiae generis polygoni prodromusGenevae, 1826
[17] Gross M H-Remarques sur les polygonees del'Asie orientaleBulletin of the Torrey Botanical Club, 1913,23:7-32
[18] Stewad A N. The Polygoneae of eastern Asia. Contributions from the Gray Herbarium of Harvard University, 1930,88(88):1-129
[19]柏大全·四川西部地区荞麦资源细胞学及遗传标记研究·雅安:四川农业大学,2009Bo D Q. Research on Cytology and genetic mark among buckwheat species in west of China. Yaan:Sichuan Agricultural University, 2009
[20] Li A J,Bao B J,Grabovskaya-Borodina A E,Hong S,McNeill J,Mosyakin S L, Ohba H, Park C-Polygonaceae, Flora of China.Beijing:Science Press, 2003:277-350
[21] Ohnishi O. Cultivated buckwheat species and their relatives in the Himalaya and southern China. Proc. 4th International Symposium Buckwheat at Orel, 1989:562-571
[22]林汝法.中国荞麦.北京:中国农业出版社,1994:51-56Lin R F-Buckwheat in China. Beijing:China Agricultural Press,1994:51-56
[23]吴征镒·西藏植物志:第1卷·北京:科学出版社,1983:604-605Wu Z Y. Flora of Tibet:Volume 1. Beijing:Science Press,1983:604-605
[24] Samuelsson G, Polygonaceae H-Handel-Mazzetti:Symbolae sinicae. Wien,1992
[25] Graham S A. The genera of polygonaceae in the southeastern united States. Journal of the Arnold Arboretum, 1965,46(2):91-121
[26] Hedberg O. Pollen morphology in the genus polygonum L.(S.lat.)and its taxonomical significance. Svensk BotaniskTidskrift,1946,40:371-404
[27]Yukio D-Cytological studies in polygonum and related Genera I.Botanical Magazine(Tokyo), 1960, 37:337-340
[28]Tian X M,Luo J,Wang A L,Mao K S,Liu J Q. On the origin of the woody buckwheat Fagopyrum tibeticum(=Parapteropyrum tibeticum)in the Qinhai-Tibetan Plateau.Molecular Phylogenetics and Evolution, 2011,61:515-520
[29] Zhou M L, Wang C L, Wang D Z, Zheng Y D, Li F L, Zhu X M,Tang Y X, Shao J R, Tang Y, Wu Y M. Phylogenetic relationship of four new species related to southwestern Sichuan Fagopyrum based on morphological and molecular characterization.Biochemical Systematics and Ecology, 2014, 57:403-409
[30] Zhang L J, Li X X, Ma B,Gao Q, Du H L, Han Y H,Li Y,Cao Y H, Qi M, Zhu Y X, Lu H W, Ma M C, Liu L L, Zhou J P, Nan C H, Qin Y J, Wang J, Cui L, Lui H M, Liang C Z,Qiao Z J. The Tartary buckwheat genome provides Insights into rutin biosynthesis and abiotic stress tolerance-Molecular Plant,2017, 10(9):1224-1237
[31]Yasui Y,Hirakawa H,Ueno M,Matsui K,Katsube-Tanaka T,Yang S J, Ali J, Sato S, Mori M. Assembly of the draft genome of buckwheat and its applications in identifying agronomically useful genes. DNA Research, 2016,23(3):215-224
[2]李安仁·中国植物志:第25分册,寥科·北京:科学出版社,1998:108-117Li A R. Flora of China:Volume 25, Polygonaceae. Beijing:Science Press, 1998:108-117
[3] Ohnishi O.Discovery of new Fagopyrum species and its implication for the studies of evolution of Fagopyrum and of the origin of cultivated buckwheat. Japan:Shinshu University Press, 1995:175-190
[4] Ohnishi O-Search for the wild ancestor of buckwheat I Description of new Fagopyrum(Polygonaceae)species and their distribution in China. Fagopyrum, 1998, 15:18-28
[5] Ohsako T, Ohnishi O. New Fagopyrum species revealed by morphological and molecular analyses. Genes&Genetic Systems, 1998,73:85-94
[6] Ohsako T, Yamane K, Ohnishi O. Two new Fagopyrum(polygonaceae)species F.gracilipedoides and F. jinshaense from Yunnan-Genes&Genetic Systems, 2002, 77:399-408
[7] Chen Q F. A study of resources of Fagopyrum(Polygonaceae)native to China. Botanical Journal of Linnean Society, 1999,130(1):54-65
[8]刘建林,唐宇,夏明忠,邵继荣,蔡光泽,罗强,孙俊秀.中国四川寥科荞麦属一新种——皱叶野荞麦·植物分类学报,2008,46(6):929-932Liu J L, Tang Y, Xia M Z, Shao J R, Cai G Z, Luo Q, Sun J X. Fagopyrum crispatifolium J. L-Liu, a new species of Polygonaceae from Sichuan, China. Journal of Systematics and Evolution, 2008,46(6):929-932
[9] Tang Y, Zhou M L, Bai D Q, Shao J R, Zhu X M, Wang D Z, Tang Y X-Fagopyrum pugense(Polygonaceae), a new species from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2010, 20(2):239-242
[10] Shao J R, Zhou M L, Zhu X M, Wang D Z, Bai D Q.Fagopyrum wenchuanense and Fagopyrum qiangcai, Two new species of Polygonaceae from Sichuan, China. Novon, 2011,21(2):256-261
[11] Hou L L, Zhou M L, Zhang Q, Qi L P, Yang B X, Tang Y,Zhu X M, Shao J R-Fagopyrum luojishanense, a New Species of Polygonaceae from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2015,24(1):22-26
[12] Zhou M L, Zhang Q, Zheng Y D, Tang Y, Li F L, Zhu X M,Shao J R-Fagopyrum hailuogouense(Polygonaceae), one new species of Polygonaceae from Sichuan, China. Novon:A Journal for Botanical Nomenclature, 2015,24(2):222-224
[13] Wang C L, Li Z Q,Ding M Q,Tang Y U,Zhu X,Liu JFagopyrum longzhoushanense, a new species of Polygonaceae from Sichuan, China. Phytotaxa, 2017, 291(1):73-80
[14] Zhou M L, Kreft I, Suvorova G, Tang Y, Woo S H-Buckwheat germplasm in the world. ELSEVIER, San Diego, USALondon:Academic Press, 2018:61-95
[15] Miller P H. The gardeners dictionary. Abridged(edition 4)·London, 1754
[16] Meisner C F. Monographiae generis polygoni prodromusGenevae, 1826
[17] Gross M H-Remarques sur les polygonees del'Asie orientaleBulletin of the Torrey Botanical Club, 1913,23:7-32
[18] Stewad A N. The Polygoneae of eastern Asia. Contributions from the Gray Herbarium of Harvard University, 1930,88(88):1-129
[19]柏大全·四川西部地区荞麦资源细胞学及遗传标记研究·雅安:四川农业大学,2009Bo D Q. Research on Cytology and genetic mark among buckwheat species in west of China. Yaan:Sichuan Agricultural University, 2009
[20] Li A J,Bao B J,Grabovskaya-Borodina A E,Hong S,McNeill J,Mosyakin S L, Ohba H, Park C-Polygonaceae, Flora of China.Beijing:Science Press, 2003:277-350
[21] Ohnishi O. Cultivated buckwheat species and their relatives in the Himalaya and southern China. Proc. 4th International Symposium Buckwheat at Orel, 1989:562-571
[22]林汝法.中国荞麦.北京:中国农业出版社,1994:51-56Lin R F-Buckwheat in China. Beijing:China Agricultural Press,1994:51-56
[23]吴征镒·西藏植物志:第1卷·北京:科学出版社,1983:604-605Wu Z Y. Flora of Tibet:Volume 1. Beijing:Science Press,1983:604-605
[24] Samuelsson G, Polygonaceae H-Handel-Mazzetti:Symbolae sinicae. Wien,1992
[25] Graham S A. The genera of polygonaceae in the southeastern united States. Journal of the Arnold Arboretum, 1965,46(2):91-121
[26] Hedberg O. Pollen morphology in the genus polygonum L.(S.lat.)and its taxonomical significance. Svensk BotaniskTidskrift,1946,40:371-404
[27]Yukio D-Cytological studies in polygonum and related Genera I.Botanical Magazine(Tokyo), 1960, 37:337-340
[28]Tian X M,Luo J,Wang A L,Mao K S,Liu J Q. On the origin of the woody buckwheat Fagopyrum tibeticum(=Parapteropyrum tibeticum)in the Qinhai-Tibetan Plateau.Molecular Phylogenetics and Evolution, 2011,61:515-520
[29] Zhou M L, Wang C L, Wang D Z, Zheng Y D, Li F L, Zhu X M,Tang Y X, Shao J R, Tang Y, Wu Y M. Phylogenetic relationship of four new species related to southwestern Sichuan Fagopyrum based on morphological and molecular characterization.Biochemical Systematics and Ecology, 2014, 57:403-409
[30] Zhang L J, Li X X, Ma B,Gao Q, Du H L, Han Y H,Li Y,Cao Y H, Qi M, Zhu Y X, Lu H W, Ma M C, Liu L L, Zhou J P, Nan C H, Qin Y J, Wang J, Cui L, Lui H M, Liang C Z,Qiao Z J. The Tartary buckwheat genome provides Insights into rutin biosynthesis and abiotic stress tolerance-Molecular Plant,2017, 10(9):1224-1237
[31]Yasui Y,Hirakawa H,Ueno M,Matsui K,Katsube-Tanaka T,Yang S J, Ali J, Sato S, Mori M. Assembly of the draft genome of buckwheat and its applications in identifying agronomically useful genes. DNA Research, 2016,23(3):215-224
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