He-Nygrén Xiaolan, Juslén Aino, Ahonen Inkeri, Glenny David, Piippo Sinikka
Finnish Museum of Natural History, Botanical Museum, PO Box 7, FI-00014 University of Helsinki.
University of Helsinki, Department of Biological and Environmental Sciences, Plant Biology, PO Box 65, FI-00014 University of Helsinki.
Cladistics. 2006 Feb;22(1):1-31. doi: 10.1111/j.1096-0031.2006.00089.x.
The phylogenetic relationships of liverworts were reconstructed using the sequence data of four genome regions including rbcL, rps4 and trnL-F of the chloroplast and 26S large subunit ribosomal rRNA gene of the nucleus, and 90 characters of morphological, ultrastructural and developmental aspects. The taxa sampled consisted of 159 species including 135 liverworts (108 genera, 54 families and 29 suborders), 13 mosses, two hornworts, seven vascular plants and two charophyte algae. Analyses based on maximum parsimony using both direct optimization (POY) and static alignment (NONA), as well as Bayesian inference (MrBayes) were done. All the data sets were analyzed simultaneously. Our study confirms that liverworts compose a monophyletic group which consists of three classes. The class Treubiopsida including both Treubia and Haplomitrium is resolved as the earliest diverging liverwort lineage. Blasia and the complex thalloids are assigned to the Marchantiopsida, under which Blasiidae and Marchantiidae are divided. Marchantiidae include Sphaerocarpales and Marchantiales. The simple thalloid and leafy liverworts form the Jungermanniopsida, which is further divided to subclasses Pelliidae subclassis nov., Metzgeriidae and Jungermanniidae. Metzgeriidae here is defined to include only Metzgeriaceae, Aneuraceae and Vandiemeniaceae, and is the sister group to the leafy liverworts. The leafy liverworts Jungermanniidae include the orders Pleuroziales, Porellales and Jungermanniales. It is assumed that the Porellales and the Jungermanniales have split early, at least in the Jurassic period. In the Porellales, the diversification rate may have remained relatively constant for long periods of time but speeding up only recently within some of the families, associated with an explosive radiation of angiosperms. The Jungermanniales are most probably a recently diversified group which has attained the greatest profusion of structure and the most remarkable diversity of leaf development and protective devices for maturing sporophytes. A detailed classification scheme for liverworts is presented.
利用叶绿体的rbcL、rps4和trnL-F以及细胞核的26S大亚基核糖体RNA基因这四个基因组区域的序列数据，以及形态学、超微结构和发育方面的90个特征，重建了苔类植物的系统发育关系。所采样的分类群包括159个物种，其中有135种苔类植物（108属、54科和29亚目）、13种藓类植物、2种角苔、7种维管植物和2种轮藻。使用直接优化（POY）和静态比对（NONA）的最大简约法以及贝叶斯推断（MrBayes）进行了分析。所有数据集同时进行分析。我们的研究证实，苔类植物构成一个单系类群，由三个纲组成。包括Treubia和Haplomitrium的Treubiopsida纲被确定为最早分化的苔类植物谱系。Blasia和复杂叶状体苔类被归入地钱纲，在地钱纲下又分为Blasiidae科和地钱科。地钱科包括球蒴目和地钱目。简单叶状体苔类和叶状苔类构成叶苔纲，叶苔纲进一步分为新的Pelliidae亚纲、美苔亚纲和叶苔亚纲。这里的美苔亚纲被定义为仅包括美苔科、无轴藓科和范迪门藓科，是叶状苔类的姐妹群。叶状苔类的叶苔亚纲包括侧蒴苔目、光萼苔目和叶苔目。据推测，光萼苔目和叶苔目很早就已经分化，至少在侏罗纪时期就已分化。在光萼苔目中，多样化速率可能在很长一段时间内保持相对恒定，但只是最近在一些科内加快，这与被子植物的爆发式辐射有关。叶苔目很可能是一个最近才多样化的类群，其结构最为丰富，叶的发育以及成熟孢子体的保护装置的多样性最为显著。文中给出了苔类植物的详细分类方案。
