In recent years, the use of extensive molecular and morphological datasets has clarified the phylogenetic relationships among the orders of complex thalloid liverworts (Marchantiidae). However, previous studies excluded extinct taxa; thereby, undersampling the actual taxonomic diversity of the group. Here, we conducted a total-evidence analysis of Marchantiidae incorporating fossils. The combined dataset consisted of 11 genes—sampled from the nuclear, mitochondrial and plastid genomes—and 128 morphological characters. Sixty-two species, representing all classes and orders within Marchantiophyta and genera within Marchantiidae were included in the analyses. Six fossils were scored from literature: two assigned to the outgroup (Metzgeriothallus sharonae and Pallaviciniites sandaolingensis) and four to the ingroup (Marchantites cyathodoides, M. huolinhensis, Ricciopsis ferganica and R. sandaolingensis). Tree searches were conducted using parsimony as the optimality criterion. Clade sensitivity was assessed across a wide range of weighting regimes. Also, we evaluated the influence of fossils on the inferred topologies and branch support. Our results were congruent with previously inferred clades above the order level: Neohodgsoniales was sister to a clade formed by Sphaerocarpales and Marchantiales. However, relationships among families within Marchantiales contradicted recent studies. For instance, a clade consisting of Monosoleniaceae, Wiesnerellaceae and Targioniaceae was sister to the morphologically simple taxa instead of being nested within them as in previous studies. Novel synapomorphies were found for several clades within Marchantiales. Outgroup fossils were more influential than Marchantiidae fossils on overall topologies and branch support values. Except for a single weighting scheme, sampling continuous characters and down-weighting characters improved fossil stability. Ultimately, our results challenge the widespread notion that bryophyte fossils are problematic for phylogenetic inference.

中文翻译：

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不畏死亡：将化石纳入复杂的海苔（Marchantiidae，Marchantiophyta）的系统发育证实了高阶进化枝，但揭示了家庭层面关系的差异

近年来，广泛的分子和形态学数据集的使用已经阐明了复杂的海苔（Marchantiidae）目之间的系统发育关系。然而，以前的研究排除了灭绝的分类群；因此，欠采样该组的实际分类多样性。在这里，我们对包含化石的马钱蒂科进行了全证据分析。组合数据集由 11 个基因（从核、线粒体和质体基因组中采样）和 128 个形态特征组成。分析中包括了 62 个物种，代表了 Marchantiophyta 内的所有类别和目以及 Marchantiidae 内的属。从文献中对六种化石进行评分：两种属于外群（Metzgeriothallus sharonae和Pallaviciniites sandaolingensis) 和四个内群 ( Marchantites cyathodoides , M. huolinhensis , Ricciopsis ferganica和R. sandaolingensis )）。使用简约性作为最优标准进行树搜索。在广泛的加权方案中评估了进化枝敏感性。此外，我们评估了化石对推断拓扑和分支支持的影响。我们的结果与先前推断的高于订单水平的进化枝一致：Nehodgsoniales 是由 Sphaerocarpales 和 Marchantiales 形成的进化枝的姐妹。然而，Marchantiales 内的家庭之间的关系与最近的研究相矛盾。例如，由 Monosoleniaceae、Wiesnerellaceae 和 Targioniaceae 组成的进化枝是形态简单分类群的姐妹，而不是像以前的研究那样嵌套在它们之中。在马钱蒂亚目中发现了几个进化枝的新突触。外群化石在整体拓扑结构和分支支持值上的影响比马鹞科化石更具影响力。除了单一的加权方案外，采样连续特征和降低加权特征提高了化石的稳定性。最终，我们的结果挑战了苔藓植物化石对系统发育推断有问题的普遍观念。