Morphological Comparison of the Epimedium franchetii Stearn Species Complex Based on Population Observation and Implications for Taxonomy
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摘要: 基于"居群"概念,对木鱼坪淫羊藿复合种(Epimedium franchetii speciescomplex)进行了系统的标本查阅,并于花期对该复合种9个居群的野外形态进行了观测和比较研究。结果显示,竹山淫羊藿(E.zhushanense K.F. Wuet S.X. Qian)花瓣为紫红色,易于与其它物种区分;时珍淫羊藿(E.lishihchenii Stearn)在模式产地(江西庐山)的JXLS居群中有1/5的个体根茎为结节状(并非像模式种一样根茎细长),其叶背被毛细长的性状较稳定,且与木鱼坪淫羊藿(E.franchetiiStearn)叶背被粗短伏毛的性状区别明显;JXJA居群(江西靖安)地理分布靠近江西庐山,叶背被毛也与时珍淫羊藿类似,但其典型的粗壮根茎则与木鱼坪淫羊藿一致,因此将其处理为木鱼坪淫羊藿-时珍淫羊藿过渡类型;保靖淫羊藿(E.baojingense Q.L.Chenet B.M. Yang)与木鱼坪淫羊藿主要区别在于前者小叶柄、叶柄、茎和节部密被柔毛,尤以节部明显,叶背被细长毛,而木鱼坪淫羊藿小叶柄、叶柄、茎和节部均光滑,叶背被粗短伏毛;HBFX居群(湖北房县)和HBMP居群(湖北神农架)小叶柄、叶柄、茎、节部和叶背均疏被毛,被毛特征介于木鱼坪淫羊藿和保靖淫羊藿之间,因此将HBFX和HBMP居群处理为木鱼坪淫羊藿-保靖淫羊藿过渡类型。聚类分析结果表明,9个居群可划分为3类,竹山淫羊藿与其它类群种间界限明显,建议将其从木鱼坪淫羊藿复合种中分离出来;木鱼坪淫羊藿与时珍淫羊藿和保靖淫羊藿的关系复杂;保靖淫羊藿与木鱼坪淫羊藿主要是被毛上的差异,地理上存在同域分布,推测其为微生境导致的生态宗,将其处理为木鱼坪淫羊藿的变种;时珍淫羊藿与木鱼坪淫羊藿地理分布相对隔离,是由地理隔离引起的地理宗,将其处理为木鱼坪淫羊藿亚种。Abstract: Many doubts and difficulties still exist in regards to the taxonomic treatment of the species complex in the genus Epimedium. For example, the E. franchetii species complex consists of four taxa difficult to distinguish, namely, E. franchetii Stearn, E. baojingense Q.L. Chen et B.M. Yang, E. zhushanense K.F. Wu et S.X. Qian, and E. lishihchenii Stearn. To understand the morphological characters and variation patterns of the E. franchetii species complex, a comprehensive herbarium specimen inspection and field investigation (during flowering periods) were carried out based on the "population" concept. Results showed that E. zhushanense was the most unique species due to its purple spur, making it the easiest to distinguish. However, the remaining taxa were very similar in flower and inflorescence characters, and key features among taxa were complex and unclear. For example, the contrast between the compact rhizome of E. franchetii and the long-creeping rhizome of E. lishihchenii appeared to be neither stable nor reliable. In the present study, 20% of individuals from the population collected from Lushan (type locality) had compact rhizomes. Although closely resembling E. franchetii in flower, inflorescence and rhizome, E. lishihchenii diverged by having different hairs on the underside of the leaflets. Specifically, E. franchetii had leaflets with short bristle-like hairs, while E. lishihchenii had multicellular long hairs. An interesting population (JXJA) distributed near Lushan had the hair of E. lishihchenii, but rhizome of E. franchetii. This population was defined as a transition type. In addition, E. baojingense was closely allied to E. franchetii, but differed in the petiolule, petiole and flowering stem, with pubescent hairs densest at the nodes and with long multicellular hairs on the leaflets. According to our field investigations, however, the density of hairs depended on the individuals and/or populations, and was often densest when young and gradually thinned in old age. The color of the pubescent hairs was not confined to dark yellow, but ranged from white on young twigs, to yellow when mature, and dark yellow when old. In addition, two populations (HBMP and HBFX), which were transition types between E. baojingense and E. franchetii, were observed with sparse hairs on the petioles, petiolules and flowering stems. A high-morphology-population structure, estimated by the unweighted pair group method with averaging (UPGMA) cluster analysis, was observed. This method allowed for the separation of the populations into three groups. The results suggested that E. zhushanense was a good and independent species, but the rest were complicated and close. Therefore, E. zhushanense should be removed from the E. franchetii species complex. Both E. franchetii and E. lishihchenii were originally described from cultivated plants by Stearn, which appeared to have fairly narrow species concepts. As far as E. franchetii and E. baojingense are concerned, they are geographically sympatrically distributed, with the fundamental difference being their leaf hairs. Therefore, E. baojingense should be treated as E. franchetii var. baojingense (ecological race). Both E. lishihchenii and E. franchetii have isolated distribution, with definite differences in leaf hair and rhizome styles. Thus, E. lishihchenii should be treated as E. franchetii ssp. lishihchenii (geographical race).
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