We present phylogenetic analyses of the fruticose Ramalinaceae based on extensive collections from many parts of the world, with a special focus on the Vizcaíno deserts in north-western Mexico and the coastal desert in Namibia. We generate a four-locus DNA sequence dataset for accessions of Ramalina and two additional loci for Niebla and Vermilacinia. Four genera are strongly supported: the subcosmopolitan Ramalina, the new genus Namibialina endemic to SW Africa, and a duo formed by Niebla and Vermilacinia, endemic to the New World except the sorediate V. zebrina that disjunctly occurs in Namibia. The latter three genera are restricted to coastal desert and chaparral where vegetation depends on moisture from ocean fog. Ramalina is subcosmopolitan and much more diverse in its ecology. We show that Ramalina and its sister genus Namibialina diverged from each other at c. 48 Myrs, whereas Vermilacinia and Niebla split at c. 30 Myrs. The phylogeny of the fruticose genera remains unresolved to their ancestral crustose genera. Species delimitation within Namibialina and Ramalina is rather straightforward. The phylogeny and taxonomy of Vermilacinia are fully resolved, except for the two youngest clades of corticolous taxa, and support current taxonomy, including four new taxa described here. Secondary metabolite variation in Niebla generally coincides with major clades which are comprised of species complexes with still unresolved phylogenetic relationships. A micro-endemism pattern of allopatric species is strongly suspected for both genera, except for the corticolous taxa within Vermilacinia. Both Niebla and saxicolous Vermilacinia have chemotypes unique to species clades that are largely endemic to the Vizcaíno deserts. The following new taxa are described: Namibialinagen. nov. with N. melanothrix (comb. nov.) as type species, a single new species of Ramalina (R. krogiae) and four new species of Vermilacinia (V. breviloba, V. lacunosa, V. pustulata and V. reticulata). The new combination V. granulans is introduced. Two epithets are re-introduced for European Ramalina species: R. crispans (= R. peruviana auct. eur.) and R. rosacea (= R. bourgeana auct. p.p). A lectotype is designated for Vermilacinia procera. A key to saxicolous species of Vermilacinia is presented.

中文翻译：

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Ramalinaceae 中的果子属（子囊菌门、Lecanoromycetes）：其多样性和进化历史


我们基于来自世界许多地方的大量收集品，对多果的拉玛林科 (Ramalinaceae) 进行系统发育分析，特别关注墨西哥西北部的比斯卡伊诺沙漠和纳米比亚的沿海沙漠。我们为 Ramalina 种质生成了一个四基因座 DNA 序列数据集，并为 Niebla 和 Vermilacinia 生成了两个额外基因座。四个属得到大力支持：次世界性的 Ramalina 属、西南非洲特有的新属 Namibialina 以及由 Niebla 和 Vermilacinia 组成的二属（除了在纳米比亚不连贯出现的 sorediate V. zebrina 外，新世界特有的属）。后三个属仅限于沿海沙漠和丛林，那里的植被依赖于海雾的水分。拉马利纳是亚国际大都市，其生态更加多样化。我们表明 Ramalina 和它的姐妹属 Namibialina 在 c 世纪时彼此分歧。 48 Myrs，而 Vermilacinia 和 Niebla 在 c. 分裂。 30 密尔。果果属的系统发育仍未解决其祖先壳果属的问题。纳米比亚利纳和拉马利纳内的物种划界相当简单。除两个最年轻的皮质分类群外，Vermilacinia 的系统发育和分类学已完全解决，并支持当前的分类学，包括此处描述的四个新分类群。 Niebla 的次生代谢物变异通常与主要进化枝一致，这些进化枝由尚未解决系统发育关系的物种复合体组成。除了 Vermilacinia 内的皮质类群外，强烈怀疑这两个属都存在异域物种的微观特有模式。 Niebla 和 Saxicolous Vermilacinia 都具有物种分支所独有的化学型，这些物种主要是比斯卡伊诺沙漠特有的。描述了以下新分类单元：Namibiainagen。 十一月以 N. melanothrix (comb. nov.) 为模式种，还有一个 Ramalina 新种 (R. krogiae) 和 4 个 Vermilacinia 新种 (V. breviloba、V. lacunosa、V. pustulata 和 V. reticulata)。推出新组合 V. 颗粒。欧洲 Ramalina 物种重新引入了两个绰号：R. Cripans（= R. peruviana auct. eur.）和 R. rosacea（= R. bourgeana auct. pp）。为 Vermilacinia procera 指定了选型。提出了 Vermilacinia 的伞形物种的关键信息。