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 BioGeoBEARS. 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.
Full text
Available for:
BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
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.
Full text
Available for:
IZUM, KILJ, NUK, PILJ, PNG, SAZU, UL, UM, UPUK
Recent studies have provided evidence for pulses in the diversification of angiosperms, ferns, gymnosperms, and mosses as well as various groups of animals during the Cretaceous revolution of ...terrestrial ecosystems. However, evidence for such pulses has not been reported so far for liverworts. Here we provide new insight into liverwort evolution by integrating a comprehensive molecular dataset with a set of 20 fossil age constraints. We found evidence for a relative constant diversification rate of generalistic liverworts (Jungermanniales) since the Palaeozoic, whereas epiphytic liverworts (Porellales) show a sudden increase of lineage accumulation in the Cretaceous. This difference is likely caused by the pronounced response of Porellales to the ecological opportunities provided by humid, megathermal forests, which were increasingly available as a result of the rise of the angiosperms.
Full text
Available for:
IZUM, KILJ, NUK, PILJ, PNG, SAZU, UL, UM, UPUK
Amber is renowned for the exceptional preservation state of its inclusions, allowing detailed morphological analysis and providing relevant environmental, palaeoecological, geographical, and ...geological information. Amber deposits are predominantly known from North America, Europe, and Asia, and are considered to be rare on the continents that formed Gondwana. The recent discovery of fossiliferous amber deposits in Ethiopia, therefore, provides an inimitable opportunity to close gaps in the fossil record of African terrestrial biota and to study organisms which are otherwise rare in the fossil record. Here we show that diverse cryptogams are preserved in highest fidelity in Miocene Ethiopian amber. We describe gametophyte fragments of four liverworts: Thysananthus aethiopicus sp. nov. (Porellales, Lejeuneaceae), Lejeunea abyssinicoides sp. nov. (Porellales, Lejeuneaceae), Frullania shewanensis sp. nov. (Porellales, Frullaniaceae), and Frullania palaeoafricana sp. nov. (Porellales, Frullaniaceae). Furthermore, we describe a pleurocarpous moss of the extant genus Isopterygium (Hypnales, Pylaisiadelphaceae) and a lichen representing the order Lecanorales. These new specimens represent the first amber fossils of liverworts, mosses, and lichens from the African continent and render Ethiopian amber as one of the few worldwide amber deposits preserving bryophytes (mosses and liverworts) or lichens. Fossil species of Thysananthus were recorded in Eocene Baltic and Oligocene Bitterfeld as well as Miocene Dominican and probably also Miocene Mexican ambers. Fossils that can unequivocally be assigned to Lejeunea have only been found in Dominican amber so far. Neotropical ambers contain only one taxon of Frullania to date, while the genus is most diverse in Baltic, Bitterfeld, and Rovno ambers, formed in temperate regions. The new fossils support a tropical to subtropical origin of Ethiopian amber. The new African liverwort fossils are included in an updated list of leafy liverworts described from worldwide Cenozoic ambers to date.
The first African amber fossils of liverworts, mosses, and lichens are reported from Miocene Ethiopian amber. The new fossils show an exceptional life‐like preservation in crystal clear amber and four new species of liverworts are described. The cryptogam association, similar to that of Dominican amber, supports a tropical environment of Miocene Ethiopia.
Full text
Available for:
FZAB, GIS, IJS, IZUM, KILJ, NLZOH, NUK, OILJ, PILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
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.
Full text
Available for:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Lejeunea is a largely epiphytic, subcosmopolitan liverwort genus with a complex taxonomic history. Species circumscriptions and their relationships are subject to controversy; biogeographic history ...and diversification through time are largely unknown.
We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region of 332 accessions to explore the phylogeny of the Harpalejeunea-Lejeunea-Microlejeunea complex. Lejeunea forms a well-supported clade that splits into two main lineages corresponding to L. subg. Lejeunea and L. subg. Crossotolejeunea. Neotropical accessions dominate early diverging lineages of both main clades of Lejeunea. This pattern suggests an origin in the Neotropics followed by several colonizations from the Neotropics into the Paleotropics and vice versa. Most Afro-Madagascan clades are related to Asian clades. Several temperate Lejeunea radiations were detected. Eighty two of the 91 investigated Lejeunea species could be identified to species level. Of these 82 species, 54 were represented by multiple accessions (25 para- or polyphyletic, 29 monophyletic). Twenty nine of the 36 investigated species of L. subg. Lejeunea were monoicous and 7 dioicous. Within L. subg. Crossotolejeunea, 15 of the 46 investigated species were monoicous and 31 dioicous. Some dioicous as well as some monoicous species have disjunct ranges.
We present the first global phylogeny of Lejeunea and the first example of a Neotropical origin of a Pantropical liverwort genus. Furthermore, we provide evidence for the Neotropics as a cradle of Lejeunea lineages and detect post-colonization radiations in Asia, Australasia, Afro-Madagascar and Europe. Dioicy/monoicy shifts are likely non-randomly distributed. The presented phylogeny points to the need of integrative taxonomical studies to clarify many Lejeunea binomials. Most importantly, it provides a framework for future studies on the diversification of this lineage in space and time, especially in the context of sexual systems in Lejeuneaceae.
Full text
Available for:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Premise
Bryophytes form a major component of terrestrial plant biomass, structuring ecological communities in all biomes. Our understanding of the evolutionary history of hornworts, liverworts, and ...mosses has been significantly reshaped by inferences from molecular data, which have highlighted extensive homoplasy in various traits and repeated bursts of diversification. However, the timing of key events in the phylogeny, patterns, and processes of diversification across bryophytes remain unclear.
Methods
Using the GoFlag probe set, we sequenced 405 exons representing 228 nuclear genes for 531 species from 52 of the 54 orders of bryophytes. We inferred the species phylogeny from gene tree analyses using concatenated and coalescence approaches, assessed gene conflict, and estimated the timing of divergences based on 29 fossil calibrations.
Results
The phylogeny resolves many relationships across the bryophytes, enabling us to resurrect five liverwort orders and recognize three more and propose 10 new orders of mosses. Most orders originated in the Jurassic and diversified in the Cretaceous or later. The phylogenomic data also highlight topological conflict in parts of the tree, suggesting complex processes of diversification that cannot be adequately captured in a single gene‐tree topology.
Conclusions
We sampled hundreds of loci across a broad phylogenetic spectrum spanning at least 450 Ma of evolution; these data resolved many of the critical nodes of the diversification of bryophytes. The data also highlight the need to explore the mechanisms underlying the phylogenetic ambiguity at specific nodes. The phylogenomic data provide an expandable framework toward reconstructing a comprehensive phylogeny of this important group of plants.
Full text
Available for:
FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SBCE, SBMB, UL, UM, UPUK
A new species from Fiji, Lejeunea heinrichsii is described and illustrated herein based on morphological characters and molecular phylogenetic analyses. The taxonomic descriptions of L. heinrichsii ...and the comparison with their allies are presented, together with the Maximum Likelihood (ML) and Bayesian phylogenetic analyses based on sequences of two chloroplast regions (trnL‐trnF, rbcL) and the nuclear ribosomal ITS region. This delicate and rather minute species is characterized by its autoicy, eplicate perianths, the oblong‐rectangular leaf lobes, the distantly arranged underleaves with lobes to 1/2‐2/3 underleaf length, and the leaf cells without well‐developed trigones and intermediate thickenings. In the molecular phylogeny, L. heinrichsii is placed sister to the Asian L. compressiuscula from Indonesia and forms an independent lineage including accessions of Neotropical L. adpressa from the Dominican Republic and the widespread Palaeotropical L. anisophylla from São Tomé and Príncipe. Morphologically, L. heinrichsii is very similar to L. capensis, however the result based on the molecular phylogenetic analyses revealed that the two species are very distinct.
Full text
Available for:
FZAB, GIS, IJS, IZUM, KILJ, NLZOH, NUK, OILJ, PILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
Display omitted
•Species diversity within Australasian Plagiochilaceae is 29% higher than currently recognized.•36% of currently recognized Australasian species have circumscription ...issues.•Integrative taxonomy contributes to resolving robust species hypotheses within a challenging lineage of leafy liverwort.
As a framework for revisionary study of the leafy liverwort Plagiochila in Australia, two methods for species delimitation on molecular sequence data, General Mixed Yule Coalescence model (GMYC) and Automatic Barcode Gap Discovery (ABGD) were applied to a dataset including 265 individuals from Australia, New Zealand, and the Pacific. Groups returned by GMYC and ABGD were incongruent in some lineages, and ABGD tended to lump groups. This may reflect underlying heterogeneity in the history of diversification within different lineages of Plagiochila. GMYC from trees calculated using three different molecular clocks were compared, in some lineages different primary species hypotheses were returned by analyses of trees estimated under different clock models, suggesting clock model selection should be a routine component of phylogeny reconstruction for tree-based species delimitation methods, such as GMYC. Our results suggest that a minimum of 71 Plagiochilaceae species occur in Australasia, 16 more than currently accepted for the region, comprising 8 undetermined species and 8 synonyms requiring reinstatement. Despite modern taxonomic investigation over a four decade period, (1) real diversity is 29% higher than currently recognized; and (2) 12 of 33, or 36%, of currently accepted and previously untested Australasian species have circumscription issues, including polyphyly, paraphyly, internal phylogenetic structure, or combinations of two or more of these issues. These both reflect the many challenges associated with grouping decisions based solely on morphological data in morphologically simple yet polymorphic plant lineages. Our results highlight again the critical need for combined molecular-morphological datasets as a basis for resolving robust species hypotheses in species-rich bryophyte lineages.
Full text
Available for:
GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, UPUK, ZRSKP
Cambay amber originates from the warmest period of the Eocene, which is also well known for the appearance of early angiosperm-dominated megathermal forests. The humid climate of these forests may ...have triggered the evolution of epiphytic lineages of bryophytes; however, early Eocene fossils of bryophytes are rare. Here, we present evidence for lejeuneoid liverworts and pleurocarpous mosses in Cambay amber. The preserved morphology of the moss fossil is inconclusive for a detailed taxonomic treatment. The liverwort fossil is, however, distinctive; its zig-zagged stems, suberect complicate-bilobed leaves, large leaf lobules, and small, deeply bifid underleaves suggest a member of Lejeuneaceae subtribe Lejeuneinae (Harpalejeunea, Lejeunea, Microlejeunea). We tested alternative classification possibilities by conducting divergence time estimates based on DNA sequence variation of Lejeuneinae using the age of the fossil for corresponding age constraints. Consideration of the fossil as a stem group member of Microlejeunea or Lejeunea resulted in an Eocene to Late Cretaceous age of the Lejeuneinae crown group. This reconstruction is in good accordance with published divergence time estimates generated without the newly presented fossil evidence. Balancing available evidence, we describe the liverwort fossil as the extinct species Microlejeunea nyiahae, representing the oldest crown group fossil of Lejeuneaceae.
Full text
Available for:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
PDF
