系统梳理裸子植物系统分类历史,按标志性研究进展,将1753年以来裸子植物分类研究历史划分为6个发展阶段:第1阶段为混沌时期(1753—1827年),从Linnaeus至Brown提出裸子植物概念之前,裸子植物未被作为一个类群处理,以Linnaeus的性系统为代表;第2阶段为概念形成时期(1827—1843年),从Brown发现裸子植物至Brongniart正式将裸子植物处理为一个类群之前,是裸子植物发现和概念的确立时期;第3阶段为萌芽时期(1843—1864年),从Brongniart至Braun之前,这一时期人们虽然将裸子植物作为一个类群处理,但还是将它作为被子植物双子叶植物的一个部分,代表系统如Bentham & Hooker的分类系统;第4阶段为裸子植物系统分类的奠基时期(1864—1926年),从Braun至Pilger,这个阶段人们将裸子植物从被子植物中独立出来,发现了银杏精子具鞭毛这样的原始性状,将银杏从红豆杉类植物中分出,厘清了裸子植物的主要传代线,奠定了裸子植物系统分类的基础,但是该时期人们对裸子植物科的概念还比较模糊,以Eichler和Engler的系统为代表,为后续分类系统奠定基础;第5阶段为“百家争鸣”时期(1926—2011年),从Pilger至Christenhusz 等之前,该时期提出的分类系统很多,分类阶元的应用从目至门都有不同的观点,且从形态学、解剖学、胚胎学、植物化学、古植物学、分支分类学等方面,证据类型和方法十分多样。争论主要集中在银杏、红豆杉类和买麻藤类等类群的亲缘关系和系统归属,以及对罗汉松科、杉科和柏科的界定方面,以Pilger系统和郑万钧为代表;第6阶段是分子系统学和整合分类学时期(2007—),从Anderson等至今,该时期以分支分类学原理为准则,以系统发生树为骨架,综合分子系统学、解剖学、形态学、植物化学以及分子系统学等证据,建立和完善裸子植物分类系统,以Yang等提出的裸子植物分类系统为本阶段的代表性分类系统。笔者还展望了裸子植物未来在物种界定、整合分类学以及保护生物学等方面的重点研究发展趋势。

The botanical history of gymnosperm classification was reviewed and dividied into six major phases since the initiation of modern taxonomy in 1753 according to landmark concepts or creative classifications. The first phase lasted from Linnaeus to Brown during which the concept of gymnosperm was not conceived, and gymnosperms were not treated as a taxon. This phase was represented by the sexual system of Linnaeus. The second phase was from Brown to Brongniart. In this phase, the concept of gymnosperms was proposed and gradually accepted by botanists at that time, and Brongniart treated gymnosperms as a group in his classification system though the linear sequence, family delineation and relationships among taxa were ambiguous. The third phase was from Brongniart to Braun. In this phase, the gymnosperms were treated as a group within dicotyledons, the classification of Bentham & Hooker was considered as the representative classification systems. The fourth phase was from Braun to Pilger. During this period, the gymnosperms were treated as a separate taxon from angiosperms, the spermatozoids of cycads and ginkgo were discovered, the main lineages of gymnosperms became established, but the family delineation remains ambiguous, representative systems include Eichler and Engler. The fifth phase was from Pilger to Christenhusz et al.. Numerous classifications of gymnosperms were proposed in this phase, various categories were applied to lineages of gymnosperms, different lines of evidence were obtained and utilized including morphology, anatomy, embryology, phytochemistry, palaeobotany, cladistics, the focuses of debates were mainly on the relationships and systematic positions of ginkgo, taxads, and gnetophytes, and the taxonomic delineation of Podocarpaceae, Taxodiaceae and Cupressaceae, the classification systems of Pilger and Wanchun Cheng were representative in this phase. The sixth phase was from Anderson et al. till now. In this phase, people accepted the principles of cladistics, and conducted phylogenetic studies using DNA sequences, and also integrated different lines of evidences in taxonomy. The new classification of gymnosperms proposed by Yang et al. was introduced and the features of this new classification was discussed. Future studies should focus on species delimitation based on Genomic barcoding, integrative taxonomy, and conservation biology.