2024, Vol. 32
刺蕊草属(Pogostemon Desf.)隶属于唇形科(Lamiaceae)野芝麻亚科(Lamioideae)刺蕊草族(Pogostemoneae)[1]，是该族之中最大的属，约80种，主要分布于热带至亚热带亚洲地区，其物种多样性中心位于印度次大陆，少数物种扩散至温带地区, 另外非洲有5种且为其所特有[2‒3]。中国产29种2变种[4‒6]。目前所接受的刺蕊草属为其广义概念, 包含狭义刺蕊草属(Pogostemon s.s.)和水蜡烛属(Dysophylla Blume)[2‒3]。广藿香[P. cablin (Blanco) Bentham; 图 1: A]作为我国著名的南药之一[7‒8]，是该属重要的代表性物种，其提取物广藿香油在医药、化妆品以及杀虫剂等方面应用非常广泛，具有重要的经济价值[2‒3,7‒9]。该属刺蕊草亚属(subgen. Pogostemon)其他物种同样含有藿香油[2‒3]，因此对该属开展系统分类学研究对其资源的充分认识、合理利用及有效保护极为重要。近年来，关于刺蕊草属在物种水平的分类学处理、属下类群划分等方面都取得了重要进展，故在此对其系统分类学研究进展进行归纳整理，为该属后续相关研究指明方向，并为该属植物资源的可持续利用提供分类学参考。
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图 1 刺蕊草属植物。A: 广藿香; B: 香薷状刺蕊草; C: 刺蕊草; D: 北刺蕊草; E: 黑刺蕊草; F: 长柄水珍珠菜; G: 四叶水蜡烛; H: 齿叶水蜡烛; I: 水蜡烛。 Fig. 1 Pogostemon species. A: P. cablin; B: P. elsholtzioides; C: P. glaber; D: P. septentrionalis; E: P. brachystachyus; F: P. barbatus; G: P. quadrifolius; H: P. sampsonii; I: P. yatabeanus. |
刺蕊草属分类学研究历史悠久，但其范围的界定历来争议很大。关于该属分类学研究，最早可追溯至Hermann[10]于1717年的研究，他曾将水蜡烛属描述为“Veronica hirsute latifolia Zeylanica aquatic”。之后Linnaeus[11]于1747年首次发表了水蜡烛属的正式学名Alopecuro-veronica L.，但他于1767年又将这一属名处理为薄荷属(Mentha L.)的异名[12]。Desfontaines[13]于1815年基于物种Pogostemon plectranthoides Desf.正式建立刺蕊草属(Pogostemon Desf.; 叶对生)，并认为花丝被念珠状的毛是其重要形态鉴别特征(图 1: C)。另外，Blume[14]于1826年基于水珍珠菜[Dysophylla auricularia (L.) Blume; 叶对生]建立水蜡烛属(Dysophylla Blume)，认为花萼齿在果期闭合、花盘肉质肿胀以及花丝也明显被念珠状的毛等一系列综合特征是该属重要的鉴别性状。之后，水蜡烛属的界定范围被不断扩大，不少叶轮生类群也被纳入其中，但叶轮生类群于1837年又被描述为独立的水虎尾属(Eusteralis Raf.)[15]，不过这一处理在之后相当长的时期未被采纳，相关物种依旧归于水蜡烛属之中。El-Gazzar等[16]在1967年对水蜡烛属的研究中则根据叶序特征将其分为叶对生(图 1: F)和叶轮生(图 1: G~I) 2个主要类群，并认为叶对生类群与刺蕊草属可能更为接近，因此建议将其纳入刺蕊草属的范畴。这一处理直接导致剩下的水蜡烛属(Dysophylla Blume)叶轮生类群缺少模式种，Airy Shaw[17]为解决这一问题重新对水蜡烛属进行了范围界定，重新界定后的水蜡烛属仅包含叶轮生类群，四叶水蜡烛(Dysophylla quadrifolia Bentham; 图 1: G)被指定为其模式种，同时将属名修订为Dysophylla El-Gazzar & L. Watson，该观点在《中国植物志》[18]等著作中被采用。然而在Flora of China[19]，尽管水蜡烛属仅含叶轮生物种，但依然采用属名Dysophylla Blume。不过，考虑到叶轮生类群实际在1837年就已有正式发表的属名：水虎尾属(Eusteralis Raf.)[15]，该属名在上世纪又被部分学者重新采用[20‒21]。整体来看，传统界定的狭义刺蕊草属与水蜡烛属在生境、植株生活型、花萼大小等方面都存在较为明显的差异。其中前者为陆生类群(图 1: A~E)，常为多年生灌木或亚灌木，稀草本，花萼较大(长通常超过3 mm)；而后者常为水生类群(图 1: F, H~I)，多为一年生草本或稀多年生草本，花萼较小(长通常明显不足3 mm)，但有极少数物种似乎处于这个属的中间过渡状态(图 1: G; 陆生，叶轮生，花萼较小)，将二者紧密联系在一起，也暗示了这2属间的近缘关系。
另一方面，考虑到水蜡烛属与刺蕊草属物种的花丝基本上都具有念珠状的毛这一形态特征, Hasskårl早在1842年就曾建议将二者合并[22]，成立广义刺蕊草属(Pogostemon s.l.)，并将花丝被念珠状的毛这一特征作为其形态鉴别特征(图 1: C)，这一观点被后来众多学者所接受[2,23‒26]，且在Bendiksby等[27]的研究中首次得到分子系统学研究结果的支持，在近期对该属进行广泛取样的分子系统学研究中进一步获得支持[3]。于此，刺蕊草属界定范围被最终澄清，即刺蕊草属应接受为其广义概念，包含传统界定的狭义刺蕊草属与水蜡烛属所有物种，这一观点在最近唇形科分类系统中也被采纳[1]。
2 刺蕊草属属下类群划分研究历史由于刺蕊草属形态特征变异较大，使得该属属下类群的划分历来争议很大。Bentham于1833年最早对狭义刺蕊草属和水蜡烛属进行过属下类群划分[28]，基于花序形态将刺蕊草属分为2组：锥序组(sect. Paniculatae Benth.; 花序多分支，呈圆锥花序状；图 1: B, D)和总序组(sect. Racemosae Benth.; 花序不分支，呈总状花序；图 1: E)，基于叶序特征将水蜡烛属分为2组：叶对生组(sect. Oppositifoliae Benth.；图 1: F)和叶轮生组(sect. Verticillata Benth.；图 1: H, I)。Briquet则基于花萼特征将水蜡烛属分为2组[29]：sect. Rhabdocalicinae Briq. (花萼圆柱形)和sect. Goniocalicinae Briq. (花萼明显五棱形)，并进一步将前者分为2系：叶对生系(§1. Oppositifoliae Benth.)和叶轮生系(§2. Verticillatae Benth.)。Kudô则对茎被毛情况、叶形及叶边缘的齿等性状进行综合考虑, 将水蜡烛属界定为2组：sect. Chotekia Kudo和sect. Eudysophylla Kudo[30]。
另一方面，在广义刺蕊草属框架下，Keng曾基于叶形将其分为2组：叶对生组(sect. Pogostemon)和叶轮生组[sect. Eusteralis (Rafin.) Keng][25]。在基于大量形态特征对刺蕊草属的分类修订研究中, Bhatti等[2]对广义刺蕊草属重新进行了属下类群划分, 基于花萼大小、花萼脉的数目、花序特征等一系列特征将其界定为3亚属：刺蕊草亚属(subgen. Pogostemon)、异刺蕊草亚属(subgen. Allopogostemon Bhatti & Ingr.)和水蜡烛亚属[subgen. Dysophyllus (Blume) Bhatti & Ingr.]，并对相关亚属进一步进行了组及亚组水平类群的划分。然而基于分子系统学研究结果，Yao等[3]发现前人所提出的刺蕊草属相关属下分类系统并未能获得支持，揭示出该属内部复杂的系统进化关系。因此，基于所得分子系统发育框架并结合性状演化分析结果，Yao等[3]对刺蕊草属进行了新的属下类群划分，将该属分为2亚属：刺蕊草亚属(subgen. Pogostemon)和水蜡烛亚属(subgen. Dysophyllus)，其中前者包含了狭义刺蕊草属中的锥序组物种(图 1: A~D)，而后者包含了传统水蜡烛属所有物种和狭义刺蕊草属总序组物种(图 1: E~I)。形态特征上，新界定的刺蕊草亚属为多年生灌木或亚灌木，稀草本；花序通常具有多余2个侧生分支，呈圆锥状(图 1: B, D)；苞片和小苞叶较宽大，通常呈宽卵形、卵形或稀披针形；水蜡烛亚属则为一年生草本，或稀为多年生草本或亚灌木，花序单一无分支或稀具2个侧生分支，常呈总状花序状(图 1: E, F, H)；苞片和小苞叶较狭窄且小，通常呈披针形, 线性或丝状。另外，这一属下类群划分观点同样获得了植物化学成分方面证据的支持，刺蕊草亚属类群含广藿香油而水蜡烛亚属类群不含广藿香油[3]。刺蕊草属不同时期的属下类群划分观点见表 1。
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表 1 刺蕊草属(含水蜡烛属)属下分类处理观点比较 Table 1 Comparison of related infrageneric classifications in Pogostemon (including Dysophylla) |
尽管对刺蕊草属分类学研究较多，但该属中不少物种在分类学处理上争议较大，物种数目也不断变动。Press[26]于1982年采用广义刺蕊草属的观点，对其物种进行过整理，认为该属约71种。在其基础上，Bhatti等[2]于1997年对刺蕊草属物种进行了重新整理，收录79种，其中刺蕊草亚属收录21种；不过，他们同时也认为刺蕊草亚属物种界定较难, 将该亚属所有21种全部处理为广布且形态多变的Pogostemon plectranthoides Desf.也未尝不可。在这2项研究中，由于查阅的中国标本有限或部分物种未能查阅到任何标本，部分中国物种未被收录或对部分种的处理存在问题，例如Dysophylla lythroides Diels、D. tsiangii Y. Z. Sun等早已被处理为其他种的异名，依然被Press接受并给予新的组合名称Pogostemon lythroides (Diels) Press、P. tsiangii (Y. Z. Sun) Press[26]，P. lythroides (Diels) Press同样被Bhatti等[2]接受。
近年来，随着野外考察工作和物种水平分类学研究的深入开展，刺蕊草属物种水平的分类学研究不断完善，多个新种发表，如印度的Pogostemon jaitapurensis Chandore & S.R. Yadav[31]和P. raghavendranii R. Murugan et Livingst.[32]，泰国的P. nudus Bongcheewin & Pramali[33]，关岛地区的关岛刺蕊草(P. guamensis Lorence & W.L. Wagner)[34]，中国台湾地区的深山刺蕊草(P. monticola T. C. Hsu, S. W. Chung, S. H. Liu & W. J. Huang)[5]和海南省的海南刺蕊草(P. hainanensis L.X. Yuan & Gang Yao)[6]等。此外，曾被认为是中国特有种的长苞刺蕊草(P. chinensis C. Y. Wu et Y. C. Huang)被发现在印度东北部地区也有分布[35]；中国特有种狭叶刺蕊草(P. dielsianus Dunn)在其模式标本于1905年被采集过之后的100多年间未在野外见其踪影，但近期在其模式产地重新发现其野生居群[36]。由此不难看出，刺蕊草属物种水平的多样性还有待深入挖掘，对该属类群开展广泛的野外考察工作还有待加强，相关工作将进一步深化对于该属生物多样性构成的认识。
对于中国刺蕊草属物种的分类学研究，《中国植物志》和Flora of China处理结果较为一致[18‒19]，均采用狭义观点将其处理为狭义刺蕊草属与水蜡烛属，且认为中国产刺蕊草属约16种，水蜡烛属约7种。然而中国所产部分物种，尤其是部分特有种，在其作为新种发表时大多仅与中国本土物种进行形态特征比较，缺少在全属层面的仔细研究，因此其分类学地位是否可靠缺乏深入研究。基于前人对中国刺蕊草属物种处理所存在的诸多问题，Yao等[4] 2015年对其进行了分类学修订，基于广泛的野外考察及标本查阅确认中国产刺蕊草属27种2变种, 其中发表一新种：河南水蜡烛(Pogostemon henanensis Gang Yao)；接受Rehder[37] 1935年的观点，将膜叶刺蕊草[P. esquirolii (H. Lév.) C.Y. Wu & Y.C. Huang]处理为刺蕊草(P. glaber Bentham)的异名，并将膜叶刺蕊草的金平变种组合为刺蕊草的变种：P. glaber var. tsingpingensis (C.Y. Wu & Y.C. Huang) Gang Yao；将长柱刺蕊草宽叶变种提升到种的水平并处理为宽叶刺蕊草：P. latifolius (C.Y. Wu & Y.C. Huang) Gang Yao；将P. brevicorollus Y.Z. Sun、P. nigrescens Dunn、P. tsiangii (Y.Z. Sun) Press等6个名称处理为其他种的异名；确认了四叶水蜡烛[P. quadrifolius (Benth.) Kuntze]和小穗水蜡烛[P. fauriei (H. Lév.) Press]在中国的分布。综合中国刺蕊草属近期研究结果[4]，本研究确认中国产该属29种2变种，种间识别特征可参考本文物种水平检索表。
4 形态解剖特征研究花萼脉的特征在刺蕊草属类群中多样化程度较大，Bhatti等[2]曾对此进行过广泛研究，结果表明, 在刺蕊草属植物花萼在主脉特征上大致可分为具5脉和10脉2种类型，在其所接受的属下类群划分框架下，刺蕊草亚属与水蜡烛亚属均具5脉，而异刺蕊草亚属则具10脉。另外，陆生类群刺蕊草亚属和异刺蕊草亚属花萼均具明显的二级脉，但二者在二级脉的生长方向上又存在明显差异，而水生类群水蜡烛亚属花萼通常无明显的二级脉。刺蕊草亚属和异刺蕊草亚属花萼二级脉在形态上的差异意味着这一特征在这2个亚属中可能并非同源，这一点在基于分子系统框架的该属性状演化分析中获得了支持[3]。
小坚果形态特征在唇形科分类处理上被认为具有重要的分类学价值[38‒40]。Bhatti等[2]曾对刺蕊草属小坚果形态特征进行过广泛研究，认为水蜡烛亚属类群小坚果大多呈长圆球形，而刺蕊草亚属与异刺蕊草亚属小坚果形态则变异较大，如呈卵形、长圆球形、椭圆体、披针状、线性披针状、圆球形、D-形等多种形状。在表面纹饰上，在不同类群的鉴定上似乎具有较好的分类学价值，如水蜡烛亚属小坚果表面通常较为平滑，异刺蕊草亚属中的subsection Glabriusculus类群小坚果表面则具有明显腺体。
孢粉形态特征研究通常被认为具有重要的分类学价值，然而关于刺蕊草属花粉形态特征的研究较少。Abu-Asab等[41]的研究曾涉及了刺蕊草属10种，认为刺蕊草属花粉具3沟，外壁纹饰呈双网状，且部分种的网眼包含有被多个小孔包围着的1至几个大孔，花粉壁基柱不分支。另外，刺蕊草属在花粉形态上与广防风属(Anisomeles R. Br.)相似，这与分子系统学研究结果相一致，分子系统学研究结果支持二者为支持率很高的姐妹群关系[3,27]。不过, 尽管Abu-Asab等[41]对刺蕊草属物种的取样涉及到不同的进化分支，但该属不同物种花粉形态相似性很大，在属下类群划分方面意义不大。
Yao等[3]对刺蕊草属进行了较为广泛的性状演化研究，认为该属可能经历过复杂的进化历史，传统分类处理中用于该属属下类群划分所依据的众多形态性状可能是趋同进化所致，并不具有很好的系统学价值，如花萼较大、花萼具5脉或10脉、花萼具二级脉、叶对生等。相比之下，花序特征以及苞片与小苞叶形态特征则具有很好的系统学价值，可用于该属之中重新界定的刺蕊草亚属与水蜡烛亚属的划分[3]。
5 分子系统学研究进展Bendiksby等[27]基于4个序列片段(matK、rps16、trnL、trnL-F)所构建的唇形科野芝麻亚科系统发育关系中，对刺蕊草属取样8种，结果揭示出这8种形成1个单系分支并与广防风属呈姐妹群关系，而传统界定的水蜡烛属形成一个分支并嵌套在狭义刺蕊草属之中。Yao等[3]对30种刺蕊草属植物，基于6个序列片段(matK、rbcL、rps16、trnH-psbA、trnL-F、nrITS)构建该属系统发育关系，结果表明Bhatti和Ingrouille [2]所界定的刺蕊草属所有3亚属均非单系；刺蕊草亚属在包含水蜡烛亚属中的短冠刺蕊草(P. amaranthoides Benth.)的情况下形成1个单系分支并与狭义刺蕊草属中的锥序组相对应，而异刺蕊草属形成2个分支混在水蜡烛亚属中；水蜡烛亚属叶轮生类群构成1个单系分支深深嵌套在叶对生的物种内部，意味着其独立的属级地位未能获得支持。根据系统发育研究初步结果，Yao等[3]将刺蕊草属界定为刺蕊草亚属与水蜡烛亚属2个主要类群，与该属中2个主要的进化分支相对应。不过, 由于该属类群可能发生过物种辐射分化，种间关系(尤其刺蕊草亚属)的解析度还不够；此外在水蜡烛亚属之中，尽管被揭示出几个独立的进化分支，但由于相关分支物种取样不够充分，对其在组水平的划分还未展开。
系统发育基因组学研究近年来广泛开展，为众多被子植物类群疑难系统关系的解析提供了极大帮助[1,42‒43]。然而目前涉及到刺蕊草属物种的相关研究主要为叶绿体系统发育基因组学分析，且数量还较少，对该属物种取样也极少[1,44‒45]，目前Zhang等[45]在该属取样最多也仅涉及5种。
6 展望到目前为止，尽管刺蕊草属在系统分类学方面取得了重要进展，不少问题被澄清，但依然存在部分问题有待解决。首先，属下类群划分尚不完善, 目前基于系统发育研究结果新提出的属下类群划分仅到亚属层面，尤其对于目前所界定的水蜡烛亚属，所包含类群涉及进化分支较多且形态特征变异丰富，有必要基于更加广泛取样的系统发育框架, 对其进一步完善组以及亚组水平的类群划分。其次，物种水平系统发育关系的解决有深入开展，前期系统发育关系研究对该属物种水平取样有限，到目前为止该属有近半数物种未被取样；在前期已取样的物种中大量种间关系也未能得到很好解决，尤其是具有重要药用价值的刺蕊草亚属。再者，物种水平的分类学修订有待进一步开展，到目前为止该属中有不少物种仅有模式标本或除模式标本之外仅有极少数几份非模式标本，这明显不利于对相关物种的形态变异式样较为全面的把握，其独立的种级地位有待深入研究加以阐明；广泛的野外调查并加大标本采集工作，对该属物种地理分布区信息的完善以及可能还存在的新种或分布新记录种的发现同样至关重要。综上所述，在后期工作中，进一步加强野外考察并不断完善物种水平甚至同种不同居群的研究工作，在此基础上完善对该属物种水平的取样用于系统发育分析，尤其需采用系统发育基因组学手段来提高物种水平系统关系的解析度很有必要。在针对刺蕊草亚属的研究中，基于同种不同居群的广泛采样，挖掘有效序列数据用于DNA条形码物种鉴定分析，结合形态特征数据澄清物种之间的界限；基于叶绿体基因组数据与核基因组序列数据，从不同角度揭示相关物种之间的进化关系，尤其揭示出广藿香这一重要物种的野生近缘种，在此基础上完善对于该属类群植物资源的开发、利用与评价等工作，将是后期关于该属系统分类学研究工作的重点。
中国刺蕊草属物种检索表
1. 多年生灌木或亚灌木；花序常具有超过2个侧生分枝，呈圆锥花序状；小苞叶和苞片较宽大，常呈宽卵形、卵形或稀
披针状································································································································· 2 (刺蕊草亚属)
1. 常为一年生草本；花序常单一无分枝，或极稀具2个侧生分支，呈总状花序状；小苞叶和苞片狭长，呈披针形、线性
或丝状······························································································································· 15 (水蜡烛亚属)
2. 花萼长不足3 mm; 花冠比花萼短或稍长···································································· 短冠刺蕊草P. amaranthoides
2. 花萼长常超过3 mm (深山刺蕊草2.5~3.5 mm); 花冠长于花萼且明显伸出····························································· 3
3. 栽培种；花萼长6~8 mm········································································································· 广藿香P. cablin
3. 野生种；花萼长3~5 mm··························································································································· 4
4. 叶狭椭圆形或披针形································································································································ 5
4. 叶卵形，椭圆形，或宽卵形······················································································································· 6
5. 花萼管状，长3.5~4.5 mm，花萼齿长为花萼筒的1/5~1/4；花冠长7~8 mm······························ 狭叶刺蕊草P. dielsianus
5. 花萼钟状，长3~3.5 mm，花萼齿长为花萼筒的1/3~1/2；花冠长约4.5 mm······················· 香薷状刺蕊草P. elsholtzioides
6. 叶边缘缺刻-锯齿状或重缺刻-锯齿状············································································································ 7
6. 叶边缘具重锯齿，重圆齿-锯齿，或重齿······································································································· 8
7. 茎、叶近无毛或毛被非刚毛状；花萼齿长约为花萼筒的1/2··············································· 台湾刺蕊草P. formosanus
7. 茎、叶明显被刚毛；花萼齿长约为花萼筒的1/3························································· 苍耳叶刺蕊草P. xanthiiphyllus
8. 花穗较短，长0.7~3.5 cm···························································································· 短穗刺蕊草P. parviflorus
8. 花穗较长，通常可超过5 cm，有时超过10 cm······························································································· 9
9. 花萼齿长为花萼筒的2/3或与之近等长········································································· 海南刺蕊草P. hainanensis
9. 花萼齿长不足花萼筒的2/3，一般不超过花萼筒长的1/2················································································· 10
10. 花萼内具腺体······································································································································ 11
10. 花萼内无腺体······································································································································ 13
11. 花萼近管状，萼齿狭三角形························································································· 长苞刺蕊草P. chinensis
11. 花萼漏斗状，萼齿三角形······················································································································· 12
12. 花萼齿长、宽均为1.3~1.5 mm；中国台湾地区特有·························································· 深山刺蕊草P. monticola
12. 花萼齿长、宽均为0.8~1 mm；中国大陆特有································································ 北刺蕊草P. septentrionalis
13. 叶片侧脉3对；花萼齿长为花萼筒的1/3~1/2；花柱长2~2.5 mm····································· 宽叶长柱刺蕊草P. latifolius
13. 叶侧脉5对；花萼齿长为花萼筒的1/3；柱头裂片长不超过1.5 mm································································· 14
14. 茎、叶、小苞片以及花萼外面近无毛或被稀疏微柔·············································· 刺蕊草原变种P. glaber var. glaber
14. 茎、叶、小苞片以及花萼外面被长柔毛················································ 刺蕊草金平变种P. glaber var. tsingpingensis
15. 叶完全对生········································································································································· 16
15. 叶轮生或对生与轮生共存······················································································································· 20
16. 花穗疏松；花萼具10脉；花萼齿通常呈钻形，果期尤为明显········································································ 17
16. 花穗紧密；花萼具5脉；花萼齿呈三角形或宽三角形，稀狭三角形································································· 19
17. 花萼长4~6 mm；花冠可长达7.5 mm················································································ 小刺蕊草P. fraternus
17. 花萼长不超过4 mm；花冠长不超过5 mm·································································································· 18
18. 叶侧脉3~5对；花萼在果期呈管状至钟形；花丝中部稀疏被毛········································ 黑刺蕊草P. brachystachyus
18. 叶侧脉5~9对；花萼果期常呈壶状；花丝无毛·························································· 刚毛萼刺蕊草P. hispidocalyx
19. 茎、叶具黑色或深褐色糙伏毛；叶柄长通常超过1 cm；叶边缘具圆齿至重圆齿，基部宽楔形至楔形····················
·· ····························································································································· 长柄水珍珠菜P. barbatus
19. 茎、叶具黄色糙伏毛；叶近无柄或柄长不超过1 cm；叶边缘具锯齿或重锯齿，基部圆形或浅心形，稀楔形···········
·· ······························································································································· 水珍珠菜P. auricularius
20. 叶对生与轮生都存在，具短柄················································································································· 21
20. 叶完全轮生，无柄································································································································ 22
21. 叶片镰状················································································································ 镰叶水珍珠菜P. falcatus
21. 叶片线性至披针形·································································································· 四叶水蜡烛P. quadrifolius
22. 茎、叶明显被毛··································································································································· 23
22. 茎、叶无毛········································································································································· 25
23. 茎被锈色微柔毛；花萼长2.5~2.8 mm；花冠长4~4.2 mm······················································ 线叶水蜡烛P. linearis
23. 茎被糙伏毛；花萼长不足2 mm；花冠长不足3 mm······················································································· 24
24. 茎被向外开展的黄色糙硬毛；叶边缘全缘，边缘极外卷····················································· 毛茎水蜡烛P. cruciatus
24. 茎被平伏糙硬毛；叶边缘具较远的浅锯齿，边缘平展或稍外卷········································ 思茅水蜡烛P. szemaoensis
25. 花萼长超过2 mm；花冠长超过3 mm········································································································ 26
25. 花萼长不足2 mm；花冠长不足3 mm········································································································ 27
26. 叶片长1~3.5 cm；花穗长0.8~2.5 cm，宽不超过1 cm··························································· 小穗水蜡烛P. fauriei
26. 叶片长3.5~7 cm；花穗长2.8~7 cm，宽可达1.5 cm······························································· 水蜡烛P. yatabeanus
27. 叶片长不超过3 cm；叶边缘全缘，稍外卷；花穗长不超过4 cm········································ 五棱水蜡烛P. pentagonus
27. 叶片长超过3 cm；叶边缘具细锯齿，或稀全缘并外卷；花穗长7~9 cm···························································· 28
28. 叶片倒卵状长圆形至长圆状披针形或线性披针形，边缘除基部1/3全缘外具明显的锯齿··········· 齿叶水蜡烛P. sampsonii
28. 叶片线性，边缘具较远的锯齿、细锯齿或全缘···························································································· 29
29. 叶4~8枚轮生；花萼长约1 mm；花冠长约1.5~1.8 mm····························································· 水虎尾P. stellatus
29. 叶3枚轮生；花萼长约1.6 mm；花冠长约2.5 mm·························································· 河南水蜡烛P. henanensis
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