Preservation of liverworts in amber, a fossilized tree resin, is often exquisite. Twenty-three fossil species of liverworts have been described to date from Eocene (35–50 Ma) Baltic amber. In addition, two inclusions have been assigned to the extant species Ptilidium pulcherrimum (Ptilidiales or Porellales). However, the presence of the boreal P. pulcherrimum in the subtropical or warm-temperate Baltic amber forest challenges the phytogeographical interpretation of the Eocene flora. A re-investigation of one of the fossils believed to be P. pulcherrimum reveals that this specimen in fact represents the first fossil evidence of the genus Tetralophozia, and thus is re-described here as Tetralophozia groehnii sp. nov. A second fossil initially assigned to P. pulcherrimum is apparently lost, and can be reassessed only based on the original description and illustrations. This fossil is morphologically similar to the extant North Pacific endemic Ptilidium californicum, rather than P. pulcherrimum. Divergence time estimates based on chloroplast DNA sequences provide evidence of a Miocene origin of P. pulcherrimum, and thus also argue against the presence of this taxon in the Eocene. Ptilidium californicum originated 25–43 Ma ago. As a result, we cannot rule out that the Eocene fossil belongs to P. californicum. Alternatively, the fossil might represent a stem lineage element of Ptilidium or an early crown group species with morphological similarities to P. californicum.
Abstract Aim How disjunct distributions arise and why organisms differ in diversification patterns remain some of the most compelling fundamental questions in biogeography. We carry out phylogeographical analyses of the pantropical liverwort genus Ceratolejeunea to identify its geographical origin and the dispersal routes by which it gained its Neotropical‐African disjunction. Furthermore, we investigate whether there is geographical structure in Ceratolejeunea on islands and whether island diversity is a result of radiations or recurrent migration. Location America, Africa and Australasia. Methods Sequences of two chloroplast regions ( trnL – F , rbcL ) and the nuclear ribosomal ITS region were obtained for 55 accessions of 20 species to explore the phylogeny, divergence times and ancestral areas of Ceratolejeunea . The phylogeny was reconstructed using maximum likelihood and Bayesian inference approaches, and divergence times were estimated with a Bayesian relaxed clock method and fossil and secondary calibrations. Ancestral areas were estimated using BioGeo BEARS. Results Ceratolejeunea likely originated in the Neotropics during the Palaeogene (42.2 Ma, 95% HPD : 22.2–64.8) and the initial diversification of its crown group took place between the Eocene and Miocene (25.5 Ma, 95% HPD : 16.6–35.8). Although boreotropical migration and subsequent extinction in northern regions cannot be rejected, the observed disjunctions are best explained by four transoceanic dispersal events from the Neotropics to Africa during the late Oligocene to Pleistocene. Geographical structure is prevalent on islands, particularly in the C . cornuta complex. Three species and the subgenus Ceratolejeunea are recovered as paraphyletic. Main conclusions Widespread paraphyly and sister group relationships between disjunct taxa indicate an important role of cryptic speciation and transoceanic dispersal with subsequent genetic differentiation in the evolution of Ceratolejeunea . On islands, recurrent migration, rather than radiation, has shaped bryophyte diversity.
ABSTRACT Fossil tree resins preserve a wide range of animals, plants, fungi and microorganisms in microscopic fidelity. Fossil organisms preserved in an individual piece of amber lived at the same time in Earth history and mostly even in the same habitat, but they were not necessarily parts of the same interacting community. Here, we report on an in situ preserved corticolous community from a piece of Miocene Dominican amber which is composed of a lichen, a moss and three species of leafy liverworts. The lichen is assigned to the extant genus Phyllopsora (Ramalinaceae, Lecanoromycetes) and is described as P. magna Kaasalainen, Rikkinen & A. R. Schmidt sp. nov. The moss, Aptychellites fossilis Schäf.-Verw., Hedenäs, Ignatov & Heinrichs gen. & sp. nov., closely resembles the extant genus Aptychella of the family Pylaisiadelphaceae. The three leafy liverworts comprise the extinct Lejeuneaceae species Cheilolejeunea antiqua (Grolle) Ye & Zhu, 2010 and Lejeunea miocenica Heinrichs, Schäf.-Verw., M. A. M. Renner & G. E. Lee sp. nov. and the extinct Radulaceae species Radula intecta M. A. M. Renner, Schäf.-Verw. & Heinrichs sp. nov. The presence of five associated extinct cryptogam species, four of which belong to extant genera, further substantiates the notion of a stasis in morphotype diversity, but a certain turnover of species, in the Caribbean since the early Miocene.