We evaluated the species richness and beta diversity of epiphyllous assemblages from three selected localities in Sabah, i.e. Mt. Silam in Sapagaya Forest Reserve, and Ulu Senagang and Mt. Alab in Crocker Range Park. A total of 98 species were found and a phytosociological survey was carried out based on the three study areas. A detailed statistical analysis including standard correlation and regression analyses, ordination of species and leaves using centered principal component analysis, and the SDR simplex method to evaluate the beta diversity, was conducted. Beta diversity is very high in the epiphyllous liverwort assemblages in Sabah, with species replacement as the major component of pattern formation and less pronounced richness difference. The community analysis of the epiphyllous communities in Sabah makes possible their detailed description and comparison with similar communities of other continents.

A floristic exploration was carried out of the epiphyllous bryophytes in Mount Tebu Forest Reserve, in the state of Terengganu, Peninsular Malaysia. Epiphyllous liverworts are reported for the first time from Terengganu. A total of 54 species from 18 genera and seven families of epiphyllous bryophytes were discovered, and of these, 43 liverworts are newly reported from Terengganu. The largest family is represented by Lejeuneaceae, with 44 species (81%) from 12 genera, and the largest genus is Cololejeunea, with 15 species (28%). Most of the species (43 spp., 69%) have an Indo-Pacific distribution.

The evolutionary history and classification of epiphyllous cryptogams are still poorly known. Leptolejeunea is a largely epiphyllous pantropical liverwort genus with about 25 species characterized by deeply bilobed underleaves, elliptic to narrowly obovate leaf lobes, the presence of ocelli, and vegetative reproduction by cladia. Sequences of three chloroplast regions (rbcL, trnL‐F, psbA) and the nuclear ribosomal ITS region were obtained for 66 accessions of Leptolejeunea and six outgroup species to explore the phylogeny, divergence times, and ancestral areas of this genus. The phylogeny was estimated using maximum‐likelihood and Bayesian inference approaches, and divergence times were estimated with a Bayesian relaxed clock method. Leptolejeunea likely originated in Asia or the Neotropics within a time interval from the Early Eocene to the Late Cretaceous (67.9 Ma, 95% highest posterior density [HPD]: 47.9–93.7). Diversification of the crown group initiated in the Eocene or early Oligocene (38.4 Ma, 95% HPD: 27.2–52.6). Most species clades were established in the Miocene. Leptolejeunea epiphylla and L. schiffneri originated in Asia and colonized African islands during the Plio‐Pleistocene. Accessions of supposedly pantropical species are placed in different main clades. Several monophyletic morphospecies exhibit considerable sequence variation related to a geographical pattern. The clear geographic structure of the Leptolejeunea crown group points to evolutionary processes including rare long‐distance dispersal and subsequent speciation. Leptolejeunea may have benefitted from the large‐scale distribution of humid tropical angiosperm forests in the Eocene.

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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 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 (Parmeliaceae, Lecanoromycetes) and described as P. magna Kaasalainen, with open areas, and were rich in epiphytes (Grimaldi, 1996). Based on its animal and plant inclusions, the Dominican amber forest has been characterized as a moist tropical forest, which was not remarkably different from the modern Neotropical lowland rainforest (Grimaldi 1996; Poinar & Poinar 1999). Three species of lichens, 22 species of liverworts, and 29 species of mosses have so far been described from Dominican amber, of which the liverworts clearly represent extinct taxa (Poinar et al. 2000;Rikkinen & Poinar 2008; Heinrichs et al. 2015a). Also the fossil lichens have been described as representatives of extinct species, however, the absence of several important chemical and apothecial characters hinders a reliable differentiation between the fossil and morphologically similar extant species (Poinar et al. 2000;Rikkinen & Poinar 2008 DNA sequence variation of extant species provides certain likelihood for a species age that is in accordance with the age of the amber (Heinrichs et al. 2013b, Villarreal & Renner 2014 Heinrichs et al. 2015b;Váňa et al. 2015).Here, we present a piece of Dominican amber that provides a window on a Miocene Caribbean epiphyte community. The virtually in situ preserved cryptogamic community includes a new species of the extant lichen genus Phyllopsora, a moss that is similar but clearly distinguished from the modern genus Aptychella and therefore assigned to the new fossil genus Aptychellites, as well as three extinct species of the extant leafy liverwort genera Cheilolejeunea, Lejeunea, and Radula. The members of this community are distinct from extant Caribbean species, further substantiating the notion of a stasis in morphotype diversity but a certain turnover of species in the Caribbean since the early Miocene as recently suggested by Schneider et al. (2015). It is also in line with the hypothesis of Poinar and Poinar (1999) that many of the plants and animals preserved in the Miocene Dominican amber do not have immediate extant descendants. Material & MethodsThe fossilized epiphytes are preserved in a ca 3 cm x 2 cm x 0.8 cm sized piece of Dominican amber which is housed in the amber collection of the American Museum of Natural History in New York (AMNH DR-15-3). The amber specimen originated from the amber mines in the Cordillera of the Dominican Republic.The mines in the Northern, Eastern and Western parts of the Dominican Republic yield ambers with an...

Understanding the biogeographical and diversification processes explaining current diversity patterns of subcosmopolitan-distributed groups is challenging. We aimed at disentangling the historical biogeography of the subcosmopolitan liverwort genus Lejeunea with estimation of ancestral areas of origin and testing if sexual system and palaeotemperature variations can be factors of diversification. We assembled a dense taxon sampling for 120 species sampled throughout the geographical distribution of the genus. Lejeunea diverged from its sister group after the paleocene-eocene boundary (52.2 Ma, 95% credibility intervals 50.1-54.2 Ma), and the initial diversification of the crown group occurred in the early to middle Eocene (44.5 Ma, 95% credibility intervals 38.5-50.8 Ma). The DEC model indicated that (1) Lejeunea likely originated in an area composed of the Neotropics and the Nearctic, (2) dispersals through terrestrial land bridges in the late Oligocene and Miocene allowed Lejeunea to colonize the Old World, (3) the Boreotropical forest covering the northern regions until the late eocene did not facilitate Lejeunea dispersals, and (4) a single long-distance dispersal event was inferred between the Neotropics and Africa. Biogeographical and diversification analyses show the Miocene was an important period when Lejeunea diversified globally. We found slight support for higher diversification rates of species with both male and female reproductive organs on the same individual (monoicy), and a moderate positive influence of palaeotemperatures on diversification. Our study shows that an ancient origin associated with a dispersal history facilitated by terrestrial land bridges and not long-distance dispersals are likely to explain the subcosmopolitan distribution of Lejeunea. By enhancing the diversification rates, monoicy likely favoured the colonisations of new areas, especially in the Miocene that was a key epoch shaping the worldwide distribution. Intercontinental biogeographical disjunctions, in which species span several continents, have long been a main focus of evolutionary biologists and biogeographers. Since its inception, continental drift theory has provided exemplar systems in support of vicariance biogeography. However, the importance of vicariance in biogeography has been reconsidered since evidence for dispersal, and even long-distance dispersal (LDD), has accumulated