以环地中海地区和北美葱属Allium的Amerallium亚属种类为外类群,基于36个形态学性状对葱属粗根组sect. Bromatorrhiza进行系统发育分析。简约分析获得的分支拓扑结构与分子系统学的研究结果相吻合,粗根组与环地中海地区的Amerallium亚属种类形成姐妹群关系,表明它们具有密切的亲缘关系。粗根组明显分为两支,粗根韭A. fasciculatum (x = 10,2胚珠)与染色体基数为11的种类(单胚珠)形成一支,另一支由染色体基数为7的大花韭A. macranthum与多星韭A. wallichii (2胚珠)组成,从而否定了粗根亚组subsect. Bromatorrhiza(粗根韭+大花韭+多星韭)的成立。综合已有研究结果,本研究认为粗根组内存在4个相对独立的演化支系,即多星韭及其变种、大花韭、粗根韭,以及染色体基数为11的种类(5种1变种),尽管前人建议的单胚珠亚组subsect. Uniovulatum是合理的,但没有必要在物种数量相对较少的粗根组下进一步细分。
Allium section Bromatorrhiza is endemic to the Hengduan Mountain Region and neighboring areas. Its taxonomic circumscription and systematic position have been clarified by previous systematic works, but the infra-sectional taxonomic classification and interspecific relationships within the section are as yet unclear. Here, section Bromatorrhiza is cladistically analyzed based on 36 morphological characters using subgenus Amerallium species of circle-Mediterranean Region and North America as outgroups. The topology of the tree obtained is consistent with the results derived from molecular data. Sect. Bromatorrhiza and subg. Amerallium species of circle-Mediterranean Region form a good clade, suggesting more closely relationships between them. Within the section, its species are arranged into two obvious clades. One consists of A. fasciculatum (x = 10, two ovules) and all x = 11 species (one ovule), and the other clade (x = 7, two ovules) includes A. macranthum and A. wallichii. The results suggest that subsection Bromatorrhiza previously presented is artificial and should be cancelled. Combining the existed results, the present study suggests that sect. Bromatorrhiza comprises four independent evolutionary clades, viz. A. wallichii and its variety, A. macranthum, A. fasciculatum and all x = 11 species (including five species and one variety). Although subsection Uniovulatum formerly proposed is reasonable, we argue that a subdivision of section Bromatorrhiza seems unnecessary for such a relatively small number of species.
[1] Li Q Q, Zhou S D, He X J, Yu Y, Zhang Y C, Wei X Q. Phylogeny and biogeography of Allium (Amaryllidaceae: Allieae) based on nuclear ribosomal internal transcribed spacer and chloroplast rps16 sequences, focusing on the inclusion of species endemic to China[J]. Annals of Botany, 2010,106: 709—733.
[2] Stearn W T. Allium and Milula in the Central and Eastern Nepal Himalaya[J]. Bulletin of the British Museum, Natural History (Botany), 1960,2(6): 159—191.
[3] 薛培凤. 国产葱属粗根组植物核型与亲缘关系的研究[D]. 成都: 四川大学硕士学位论文, 1994.
[4] Li Q Q, Zhou S D, He X J, Wei X Q. Phylogeny and character evolution in Allium Subgenus Amerallium (Amaryllidaceae)[J]. Plant Diversity and Resources, 2012,34: 107—119.
[5] Fritsch R. Anatomische untersuchungen an der blattspreite bei Allium L. (Alliaceae)—I. Arten mit einer einfachen Leitbün-delreihe [J]. Flora, Morphologie, Geobotanik, Oekophysiologie, 1988,181: 83—100.
[6] Friesen N, Fritsch R M, Pollner S, Blattner F R. Molecular and morphological evidence for an origin of the aberrant genus Milula within Himalayan species of Allium (Alliaceae)[J]. Molecular Phylogenetics and Evolution, 2000,17: 209—218.
[7] Hanelt P, Schulze-Motel J, Fritsch R M, Kruse J, Maass H I, Ohle H, Pistrick K. Infrageneric grouping of Allium-the Gatersleben approach[M]// Hanelt P, Hammer K, Knüpffer H. The Genus Allium: Taxonomic Problems and Genetic Resources. Germany: Gatersleben, 1992: 107—123.
[8] Fritsch R. Zur Wurzelanatomie in der Gattung Allium L. (Alliaceae)[J]. Beitr?ge zur Biologie der Pflanzen, 1992,67: 129—160.
[9] Huang D Q, Yang J T, Zhou C J, Zhou S D, He X J. Phylogenetic reappraisal of Allium subgenus Cyathophora (Amaryllidaceae) and related taxa, with a proposal of two new sections[J]. Journal of Plant Research, 2014,127(2): 275—286.
[10] Friesen N, Fritsch R M, Blattner F R. Phylogeny and new intrageneric classification of Allium L. (Alliaceae) based on nuclear rDNA ITS sequences[J]. Aliso, 2006,22: 372—395.
[11] Nguyen N H, Driscoll H E, Specht C D. A molecular phylogeny of the wild onions (Allium; Alliaceae) with a focus on the western North American center of diversity[J]. Molecular Phylogenetics and Evolution, 2008,47: 1157—1172.
[12] 杨蕾. 国产葱属粗根组植物系统演化研究[D]. 成都: 四川大学硕士学位论文, 1995.
[13] 何兴金. 国产葱属进化生物学研究[D]. 成都: 四川大学博士学位论文, 1999.
[14] 倪念春. 国产葱属粗根组植物分子系统学研究[D]. 成都: 四川大学硕士学位论文, 2002.
[15] Aedo C. Allium L.[M]// Talavera S, Andrés C, Arista M, Fernández Piedra M P, Rico E, Crespo M B, Quintanar A, Herrero A, Aedo C. Flora Iberica (Vol. 20), Liliaceae-Agavaceae. Madrid: CSIC. 2013: 220—273.
[16] McNeal D W, Jacobsen T D. Allium L.[M]// Editorial Committee of Flora of North America. Flora of North America (Vol. 26). New York: NYBG Press, 2002: 224—276.
[17] Choi H J, Cota-Sánchez J H. A taxonomic revision of Allium (Alliaceae) in the Canadian prairie provinces[J]. Botany, 2010,88(9): 787—809.
[18] 中国科学院中国植物志编辑委员会. 中国植物志(第14卷) [M]. 北京: 科学出版社, 1980: 202—213.
[19] Huang R F, Xu J M, Yu H. A study on karyotypes and their evolutionary trends in Allium Sect. Bromatorrhiza Ekberg (Liliaceae)[J]. Cathaya, 1995,7: 133—145.
[20] Wu Z Y, Raven P H. Flora of China (Vol. 24)[M]. Beijing: Science Press & St. Louis: Missouri Botanical Garden Press, 2000: 165—202.
[21] Swofford D L. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), version 4[M]. Sunderland: Sinauer Associates, 2003.
[22] McNeal D W, Ownbey M. Bulb morphology in some western North American species of Allium[J]. Madro?o, 1973,22: 10—24.
[23] Hanelt P. Taxonomy, evolution and history[M]// Rabinowitch H D, Brewster J L. Onions and Allied Crops (Vol. 1). Boca Raton: CRC Press, 1990: 1—26.
[24] Ekberg L. Studies in the genus Allium. II. A new subgenus and new sections from Asia[J]. Botaniska Notiser, 1969,122: 57—68.
[25] Li M J, Tan J B, Xie D F, Huang D Q, Gao Y D, He X J. Revisiting the evolutionary events in Allium subgenus Cyathophora (Amaryllidaceae): Insights into the effect of the Hengduan Mountains Region (HMR) uplift and Quaternary climatic fluctuations to the environmental changes in the Qinghai-Tibet Plateau[J]. Molecular Phylogenetics and Evolution, 2016,94: 802—813.
[26] Li M J, Guo X L, Li J, Zhou S D, Liu Q, He X J. Cytotaxonomy of Allium (Amaryllidaceae) subgenera Cyathophora and Amerallium sect. Bromatorrhiza[J]. Phytotaxa, 2017,331(2): 185—198.
[27] 张宇澄. 西南地区葱属系统分类学研究[D]. 成都: 四川大学硕士学位论文, 2008.
[28] Fritsch R. Septal nectaries in the genus Allium L.[M]// Hanelt P, Hammer K, Knüpffer H. The Genus Allium: Taxonomic Problems and Genetic Resources. Germany: Gatersleben, 1992: 77—85.