广义齿缘草属 (Eritrichium Schrad. ex Gaudin) 隶属于紫草科齿缘草族 (Trib. Eritrichieae Benth. et Hook.)，全世界约有95种，北温带分布，集中分布于中亚至泛喜马拉雅和北美西部，少数生长在欧洲和南美洲。虽然许多国家和地区的植物志都已编写完成，然而对于广义齿缘草属下的狭义齿缘草属和假鹤虱属 (Hackelia Opiz ex Berchtold) 的分合以及一些种归属问题上的争论仍然没有结束，而且研究范围内狭义齿缘草属的野外居群调查严重不足，近30年来馆藏标本增加甚微，在名实问题、属下系统和物种划分以上仍存有诸多问题。因此，对中国及泛喜马拉雅地区狭义齿缘草属和假鹤虱属进行全面系统研究势在必行，这既有助于完成《泛喜马拉雅植物志》的编写，同时也为世界狭义齿缘草属和假鹤虱属的修订提供资料。 本文在文献考证的基础上，研究了国内外24个标本馆1000余份标本，采集了狭义齿缘草属植物38种1亚种1变种以及假鹤虱属植物3种，收集标本材料1100余份，观察了105个居群，对研究范围内狭义齿缘草属和假鹤虱属植物形态性状的变异幅度及变异规律进行了归纳总结，对此2属进行了分类修订；利用分子系统学，探讨了中国及泛喜马拉雅地区广义齿缘草属的系统发育关系，同时结合外部形态、叶表皮毛微形态、花粉微形态、小坚果微形态等分类学证据对狭义齿缘草属进行了全面的分类学研究。本研究的主要结论如下： 1) 在宏观形态性状中，首次提出花序类型以及花序上苞片的有无、果实着生面有无白色增厚在狭义齿缘草属中具有重要的分类价值。详细总结了狭义齿缘草属的四种花序类型：仅下部花具苞片的单蝎尾状聚伞花序、全具苞片的单蝎尾状聚伞花序、二歧式蝎尾状聚伞花序以及三歧式蝎尾状聚伞花序。形态上，假鹤虱属植物与狭义齿缘草属植物极为接近，但仍有着明显的不同：茎粗壮；叶宽大，叶基多为心形；二歧式蝎尾状聚伞花序无苞片；果梗自基部强烈弯折；雌蕊基金字塔形，高超过2 mm。 2) 狭义齿缘草属植物共有两种毛被类型：糙毛和柔毛。其中糙毛根据开展与否可分为开展糙毛和贴伏糙毛；同时根据其长度的不同，又可将其划分为长糙毛和短糙毛。毛被类型在同一类群的不同居群中极为稳定，可为属下组的划分提供依据；但毛被密度受环境的影响较大，不能作为物种划分的依据。 3) 狭义齿缘草属植物的花粉形态较一致，多数为小型茧形花粉，三孔沟和三假沟相间排列；极面观多为圆角三角形；表面纹饰简单，具有不规则孔穴。主坐标分析结果显示，根据每个孔沟内萌发孔的个数可将研究种类的花粉划分为2种类型：单萌发孔型和双萌发孔型。 4) 狭义齿缘草属植物具有两种果实类型：背腹压扁的二面体型和陀螺型。数量分析显示，这两种果实类型支持将本属划分为拟假鹤虱亚属 (Subgen. Pseudohackelia (Popov) Ovczinnikova) 和齿缘草亚属 (Subgen. Eritrichium)。 5) 分子系统学研究结果表明：狭义齿缘草属和假鹤虱属均是单系群；狭义齿缘草属属下可划分为2大枝5小组；无梗齿缘草 (E. sessilifructum Y. S. Lian & J. Q. Wang)、云南齿缘草 (E. echinocaryum (I. M. Johnst.) Y. S. Lian & J. Q. Wang) 和假鹤虱齿缘草 (E. thymifolium (A. DC.) Y. S. Lian & J. Q. Wang) 应归属于狭义齿缘草属。 6) 确认中国及泛喜马拉雅地区共有狭义齿缘草属植物40种1亚种4变种以及假鹤虱属植物7种。其中发现并描述3个新物种：库地齿缘草 (E. kudiense Q. R. Liu & J .C. Hao)、喀什齿缘草 (E. kashgaricum Q. R. Liu & J .C. Hao) 以及温泉齿缘草 (E. wenquanense Q. R. Liu & J .C. Hao)。将6个名称处理为异名：E. petiolare var. villosum W. T. Wang处理为E. longifolium Decne.的异名；阿克陶齿缘草 (E. aktoense Y. S. Lian & J. Q. Wang) 处理为新疆齿缘草 (E. subjacquemontii Popov) 的异名；异果齿缘草 (E. heterocarpum Y. S. Lian & J. Q. Wang) 处理为云南齿缘草的异名；垂果齿缘草 (E. pendulifructum Y. S. Lian & J. Q. Wang) 处理为三角刺齿缘草 (E. deltodentum Y. S. Lian & J. Q. Wang) 的异名；E. lasicarpum W. T. Wang处理为E. minimum (Brand) H. Hara的异名；篦毛齿缘草 (E. pectinatociliatum Y. S. Lian & J. Q. Wang) 处理为疏花齿缘草 (E. laxum I. M. Johnst.) 的异名。同时发现小灌齿缘草 (E. canum var. fruticulosum (Klotzsch) Y. J. Nasir) 和匙叶齿缘草 (E. canum var. spathulatum (Benth.) Y. J. Nasir) 在中国没有分布。此外，由于研究材料缺乏，将互助齿缘草 (E. huzhuense X. F. Lu & G. R. Zheng) 和短梗齿缘草 (E. fetisovii Regel) 暂作存疑处理。同时综合宏观形态学、微形态学以及分子系统学提出了新的属下分类系统，将狭义齿缘草属划分为两个亚属：拟假鹤虱亚属和齿缘草亚属。其中拟假鹤虱亚属分为三个组：拟假鹤虱组 (Sect. Pseudohackelia Popov)、青藏组 (Sect. Qing-zangnica (Y. S. Lian & J. Q. Wang) J. C. Hao & Q. R. Liu) 以及假鹤虱齿缘草组 (Sect. Thymifolia J. C. Hao & Q. R. Liu)。齿缘草亚属下划分为两个组：密丛组 (Sect. Coloboma A. DC.) 和齿缘草组 (Sect. Eritrichium)。

The genus Eritrichium Schrader ex Gaudin s. l. (Boraginaceae: Eritrichieae) consists about 95 species in the north temperate zone, with most species distributed in Central Asia to the Pan-Himalaya area and western North America, and several species in Europe and South America. Although the Flora of many countries and regions have been completed successively, the debate about the treatment of Eritrichium s. l. regarding the independence of Eritrichium s. s. and Hackelia Opiz ex Berchtold and some controversial species is still not over. Also field population investigations of the genus is still seriously insufficient and the increment of Eritrichium s. s. specimens in herbarium is few in recent 30 years. Furthermore, there are still many problems in nomenclature, species division, relationship with closely related genus. Therefore, it is imperative to carry out comprehensive revision of Eritrichium s. l. in China and Pan-Himalaya area. And it is hoped that the result of the present study will contribute to compile Flora of Pan-Himalaya and also lay foundation to worldwide revision of Eritrichium s. s. and Hackelia. Based on examination literature, the author has checked more than 1000 specimens from 24 domestic and abroad herbarium; collected about 1100 specimens including 38 species, 1 subspecies and 1 variety of Eritrichium s. s. and 3 species of Hackelia; observed 105 populations and generalized variation ranges and patterns of morphological characters in the genus Eritrichium s. s. and Hackelia; carried out taxonomic revision of Eritrichium s. s. and Hackelia; discussed the phylogenetic relationship between these two genus; meanwhile, combining with leaf epidermal hair micromorphology, palynology and nutlet micromorphology carried out a comprehensive taxonomic study to the genus Eritrichium s. s.. The results are summarized as follows: 1) For the first time, proposes the inflorescence types, bract presence or not, attachment scar of nutlet with white appendages or not are highly valuable in taxonomic revision in the genus Eritrichium s. s.. Four inflorescence types are concluded: scorpioid cymes (only basal flowers with bracts), scorpioid cymes (all flowers with bracts), dich-scorpioid cymes and trich-scorpioid cyme. The genus Hackelia is very similar with Eritrichium s. s. , but differs in having brawny stems, broad leaves with cordate bases, dich-scorpioid cymes without bracts, Fruiting pedicel deflexed from the base, and the gynobase pyramidal, more than 2 mm in height. 2) Leaf epidermal indumentum of 38 species, 1 subspecies and 1 variety of Eritrichium s. s. were investigated. Two kinds of indumentum are found in the genus: strigose hair and villous hair. Herein, strigose hair can be divided into appressed strigose hair and erected strigose hair according to whether it is erected or not; and also it can be divided into long strigose hair and shortly strigose hair according to the length. Besides, the indumentums type is very stable within the genus, but density of indumentums is greatly influenced by environment. In addition, different types of indumentums can provide evidence for section division. 3) The pollen micromorphology characteristics of 36 species, 1 subspecies and 1 variety are carefully observed and compared under SEM. Meanwhile, 5 quantitative characters along with 1 qualitative character have been analyzed by Principal Coordinate Analysis (PCoA). It is showed that the shape of pollen grains in the genus are relatively consistent and are mostly small-size cocoon-shaped with three colporate apertures and three pseudocolpus. Most taxa are round triangle in polar view, while few species are suborbiculate in polar view. Irregular holes are found on the pollen grains surface. The result of PCoA shows that the pollen grains in this research can be divided into 2 types: one type with only one germinal aperture pollen in every colporate aperture and the other one with two germinal aperture pollen in every colporate aperture. 4) Nutlet micromorphology of 38 species, 1 subspecies and 1 variety of Eritrichium s. s. in China and Pan-Himalaya area are investigated by photomicrography and numerical taxonomy. Observation result shows that there are two kinds of nutlet type in Eritrichium s. s.: dorsiventrally compressed nutlet and turbinate nutlet. According to the two nutlet types, numerical taxonomy shows that the genus Eritrichium s. s. can be divided into two subgenus: Subgen. Pseudohackelia (Popov) Ovczinnikova and Subgen. Eritrichium. 5) Based on the sequence data of the nuclear ITS and plastid rps16, rpl32-trnL and trnL-F markers of 56 taxa including 40 taxa of Eritrichium s. s., 11 species of Hackelia and 5 species of Lappula Moench, the relationships and monophyly of Eritrichium s. s. and Hackelia were assessed, with special reference to the infrageneric phylogeny of Eritrichium s. s.. The result shows that Eritrichium s. s. and Hackelia are two monophyletic genus; two clade and five group are proposed in the genus Eritrichium s. s.; E. sessilifructum Y. S. Lian & J. Q. Wang, E. echinocaryum (I. M. Johnst.) Y. S. Lian & J. Q. Wang and E. thymifolium (A. DC.) Y. S. Lian & J. Q. Wang belong to the genus Eritrichium s. s.. 6) This research confirms that there are 40 species, 1 subspecies and 4 varieties of Eritrichium s. s. and 7 species of Hackelia in China and Pan-Himalaya region. Among them, 3 new species are described: E. kudiense Q. R. Liu & J .C. Hao, E. kashgaricum Q. R. Liu & J .C. Hao and E. wenquanense Q. R. Liu & J .C. Hao; 6 names are firstly reduced as synonyms: E. petiolare var. villosum W. T. Wang is reduced as the synonym of E. longifolium Decne.; E. aktoense Y. S. Lian & J. Q. Wang is reduced as the synonym of E. subjacquemontii Popov; E. heterocarpum Y. S. Lian & J. Q. Wang is reduced as the synonym of E. echinocaryum; E. pendulifructum Y. S. Lian & J. Q. Wang is reduced as the synonym of E. deltodentum Y. S. Lian & J. Q. Wang; E. lasicarpum W. T. Wang is reduced as the synonym of E. minimum (Brand) H. Hara; E. pectinatociliatum Y. S. Lian & J. Q. Wang is reduced as the synonym of E. laxum I. M. Johnst. Also, it is found out that E. canum var. fruticulosum (Klotzsch) Y. J. Nasir and E. canum var. spathulatum (Benth.) Y. J. Nasir are not distributed in China. Besides, E. huzhuense X. F. Lu & G. R. Zheng and E. fetisovii Regel remains to be further investigated due to lacking of research materials. Meanwhile, based on macromorphology, micromorphology and molecular data, 2 subgenus and 5 sections are proposed in the genus Eritrichium s. s.: Subgen. Pseudohackelia including Sect. Pseudohackelia Popov, Sect. Qing-zangnica (Y. S. Lian & J. Q. Wang) J. C. Hao & Q. R. Liu and Sect. Thymifolia J. C. Hao & Q. R. Liu; Subgen. Eritrichium including Sect. Coloboma A. DC. and Sect. Eritrichium.