前言
马兜铃属(Aristolochia L. sensu lato)是马兜铃科中种类最多和分布最广的属, 约有600种(González, 2012; Zhu et al, 2019), 广布于热带和亚热带地区, 少数分布于温带地区(马金双, 1989)。基于经典分类学和分子系统学研究, 目前对广义马兜铃属的分类主要存在两种观点: (1) 3亚属系统: 马兜铃亚属(subgen. Aristolochia): 合蕊柱顶部6(5)裂, 花药6(5), 单一的与蕊柱裂片对生, 蒴果由基部向上开裂, 分布于地中海区域及亚洲、非洲和美洲的热带和亚热带地区; 对药马兜铃亚属(subgen. Siphisia): 合蕊柱顶部3裂, 花药两两成对且与蕊柱裂片对生, 蒴果由上而下开裂, 间断分布于亚洲(东亚和南亚)和美洲(北美洲和中美洲); 多药马兜铃亚属(subgen. Pararistolochia): 合蕊柱顶部多裂, 花药多数, 浆果不开裂, 分布于非洲、澳洲的热带地区和亚洲的马来西亚地区(Duchartre, 1854, 1864; Schmidt, 1935; 马金双, 1989; González, 1999; González & Stevenson, 2000; Wanke et al, 2006; Do et al, 2015a, b)。(2) 2亚族4属系统: 马兜铃亚族(subtrib. Aristolochiinae)包含狭义马兜铃属(Aristolochia L. sensu stricto)和闭果兜铃属(Pararistolochia); 关木通亚族(subtrib. Isotrematinae)包含关木通属(Isotrema)和蛇木通属(Endodeca) (González & Stevenson, 2002; Kelly & González, 2003; Neinhuis et al, 2005; Ohi-Toma et al, 2006; Wanke et al, 2007)。Zhu等(2019)基于形态性状和分子系统学证据, 正式将对药马兜铃亚属从广义马兜铃属中恢复成关木通属(包括传统分类中的关木通属和蛇木通属)。中国所产广义马兜铃属种类皆属于现今的狭义马兜铃属和关木通属, 因两者关系密切且关木通属刚恢复不久, 故本文将两者同时撰稿。
黄淑美(1988)在《中国植物志》中首次对中国的马兜铃属(广义)进行了全面系统的整理, 共记载39种2变种3变型; 随后, 马金双(1989, 1994)进行了东亚和南亚马兜铃属(广义)的修订和补订; Hwang等(2003)在Flora of China中又对其进行了一次修订, 共记载45种, 另有金江马兜铃(Aristolochia jingiangensis Hao Zhang & C. K. Hsieh)和斑喉马兜铃(A. faucimaculata Hao Zhang & C. K. Hsien)两个存疑种未作处理。近年来, 随着调查和研究的深入, 不时有中国马兜铃属(广义)新类群发表, 截至投稿时, 共计15种1亚种(Xu et al, 2011; Huang et al, 2013, 2015; Wu et al, 2013, 2015; Lu & Wang, 2014; Zhu et al, 2015, 2017a, b, 2018a, b; Gong et al, 2018; Yang et al, 2018); 另有1变种即小花马兜铃(A. yunnanensis var. meionantha Handel-Mazzetti)被提升到了种级水平(Zhu et al, 2016); 同时有11个物种即斑喉马兜铃、扁茎马兜铃(A. compressicaulis Z. L. Yan)、川南马兜铃(A. austroszechuanica C. P. Tsien & C. Y. Cheng ex C. Y. Cheng & J. L. Wu)、川西马兜铃(Aristolochia thibetica Franch.)、大别山马兜铃(A. dabieshanensis C. Y. Cheng & W. Yu)、滇南马兜铃(A. austroyunnanensis S. M. Hwang)、柔叶马兜铃(A. mollis Dunn)、台湾马兜铃(A. shimadae Hayata)、吴氏马兜铃(A. wuana Zhen W. Liu & Y. F. Deng)、线叶马兜铃(A. neolongifolia J. L. Wu & Z. L. Yang)和异叶马兜铃(A. heterophylla Hemsl.)得到了确认、恢复或重新发表(Murata, 2006; Liu & Deng, 2009; Ohi-Toma et al, 2014; Do et al, 2015c; Zhu et al, 2016, 2017a); 长叶马兜铃(A. championii Merr. & Chun)、翅茎马兜铃(A. caulialata C. Y. Wu ex J. S. Ma & C. Y. Cheng)和偏花马兜铃(A. obliqua S. M. Hwang)得到了修订(Do et al, 2015c; Zhu et al, 2017b, 2018a); 排除了大叶马兜铃(A. kaempferi Willd.)在国内的分布(Murata, 2006; Ohi-Toma et al, 2014; Zhu et al, 2017a)。Ohi-Toma等(2014)提出了在 Flora of China 中被并入大叶马兜铃的大别山马兜铃、柔叶马兜铃、台湾马兜铃、线叶马兜铃和异叶马兜铃都应独立成种, Do等(2015a)也在检索表中承认了川西马兜铃、怒江马兜铃(A. salweenensis C. Y. Cheng & J. S. Ma)和线叶马兜铃的种级地位。除了Murata (2006)、Ohi-Toma等(2014)和Zhu等(2017a)已经确认的大别山马兜铃、台湾马兜铃和异叶马兜铃外, 本文赞同Ohi-Toma等(2014)和Do等(2015a)的观点, 认为川西马兜铃、怒江马兜铃、柔叶马兜铃、台湾马兜铃和线叶马兜铃都应独立成种。但由于所获取野外材料的限制, 本文仅对柔叶关木通(柔叶马兜铃)和线叶关木通(线叶马兜铃)的种级地位进行进一步确认。Zhu等(2019)基于形态性状和分子系统学证据将对药马兜铃亚属提升为关木通属(Isotrema Raf.), 并进行了大量的新组合, 同时将奇异马兜铃(A. kaempferi f. mirabilis S. M. Hwang)提升到了种级水平即奇异关木通(I. mirabile (S. M. Hwang) X. X. Zhu, S. Liao & J. S. Ma), 另外承认了大寒药(I. feddei (H. Lév.) X. X. Zhu, S. Liao & J. S. Ma)和云南关木通(I. yunnanense (Franch.) X. X. Zhu, S. Liao & J. S. Ma)的种级地位, 并将扁茎马兜铃(A. compressicaulis Z. L. Yan)并入凉山关木通(I. liangshanense (Z. L. Yang) X. X. Zhu, S. Liao & J. S. Ma)。由此可见, 国内本属植物已发生了巨大变化, 而且仍在持续变动中(新物种的暴发式发现、部分物种重新确认或修订、复合群的物种界定等)。截至投稿, 国内马兜铃属(狭义)植物仅剩17种, 而关木通属植物已多达58种1亚种。基于国内此两属植物已发生巨变, 为了更清晰地呈现它们在中国的种类和分布情况, 本文在Zhu等(2019)的基础上重新编制了最新的名录和检索表(附录1), 并对物种进行一一论述,同时对一些疑难和存疑类群进行了探讨(附录2)。
结语
由于国内本属在分类学方面缺乏足够的野外观察与采集, 标本量少而且花或果特征不完整, 加之很多重要花部特征在制成标本后消失或变化, 致使本属仍有很多经典分类问题没有得到妥善解决。本文的工作仅是对中国马兜铃属和关木通属的阶段性总结, 随着野外调查的深入, 网络交流的增速, 偏远地区交通的不断便利, 信息资源获取更加便捷, 越来越多的马兜铃属和关木通属新类群将持续被发现, 而随着数码照相技术的发展, 以往在标本制作过程中容易丢失的花部颜色、形态等性状也能被及时地记录下来, 为物种的准确界定提供了更多信息, 也将为这两属植物的分类修订提供有力的佐证。相信不久的将来, 国内这两属植物有望突破百种大关, 极大地丰富马兜铃科植物的多样性。
附录 Supplementary Material
附录1 中国马兜铃属和关木通属检索表
Appendix 1 Key to species of Aristolochia and Isotrema from China
附录2 中国马兜铃属和关木通属分种论述(依据检索表顺序排列)
Appendix 2 Discussion of each species of Aristolochia and Isotrema from China (order by the rank in the key)
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