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•We evaluated the oldest available fossil ascomycetes from amber and chert.•We solely used fossil ascomycetes to model a molecular clock with BEAST.•Our results suggest an initial ...diversification of ascomycetes in the Ordovician.•Continuous diversification within ascomycete classes occurred during the Phanerozoic.•Taxa of each lineage probably survived crises due to their ecological diversity.
The phylum Ascomycota is by far the largest group in the fungal kingdom. Ecologically important mutualistic associations such as mycorrhizae and lichens have evolved in this group, which are regarded as key innovations that supported the evolution of land plants. Only a few attempts have been made to date the origin of Ascomycota lineages by using molecular clock methods, which is primarily due to the lack of satisfactory fossil calibration data. For this reason we have evaluated all of the oldest available ascomycete fossils from amber (Albian to Miocene) and chert (Devonian and Maastrichtian). The fossils represent five major ascomycete classes (Coniocybomycetes, Dothideomycetes, Eurotiomycetes, Laboulbeniomycetes, and Lecanoromycetes). We have assembled a multi-gene data set (18SrDNA, 28SrDNA, RPB1 and RPB2) from a total of 145 taxa representing most groups of the Ascomycota and utilized fossil calibration points solely from within the ascomycetes to estimate divergence times of Ascomycota lineages with a Bayesian approach. Our results suggest an initial diversification of the Pezizomycotina in the Ordovician, followed by repeated splits of lineages throughout the Phanerozoic, and indicate that this continuous diversification was unaffected by mass extinctions. We suggest that the ecological diversity within each lineage ensured that at least some taxa of each group were able to survive global crises and rapidly recovered.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, 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.
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IZUM, KILJ, NUK, PILJ, PNG, SAZU, 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.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Lepidolejeunea is a largely epiphytic, pantropical genus of leafy liverworts. While the phylogenetic position of Lepidolejeunea has been the subject of recent molecular investigations, the ...relationships within this genus have been little studied. We employed sequences of two chloroplast regions (trnL-trnF, rbcL) and the nuclear ribosomal ITS region from 32 accessions of Lepidolejeunea to reconstruct its phylogeny. Two accessions of Rectolejeunea were used as outgroup. Maximum parsimony and likelihood analyses indicate that the diversity of Lepidolejeunea is underestimated by current morphological classification, and that Lepidolejeunea involuta consists of two independent entities, Lepidolejeunea involuta s.str. and Lepidolejeunea cuspidata comb. nov. Despite being treated as synonymous in earlier studies, both species nest in different main clades of Lepidolejeunea. Lepidolejeunea involuta belongs to L. subg. Kingiolejeunea which includes species with ocelli of the same size as the surrounding leaf cells. Lepidolejeunea cuspidata instead belongs to the recircumscribed L. subg. Perilejeunea which includes species having ocelli of at least partly smaller size than the surrounding leaf cells. Lepidolejeunea auriculata is introduced as a new species based on molecular and morphological evidence. Our results highlight the need for integrative taxonomic studies to clarify the status of many binomials within Lejeuneaceae. They also suggest that as integrative approaches are applied across liverworts estimates of global diversity will be revised upwards.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
• Premise of the study: Closing gaps in the fossil record and elucidating phylogenetic relationships of mostly incomplete fossils are major challenges in the reconstruction of the diversification of ...fern lineages through time. The cosmopolitan family Dryopteridaceae represents one of the most species-rich families of leptosporangiate ferns, yet its fossil record is sparse and poorly understood. Here, we describe a fern inclusion in Miocene Dominican amber and investigate its relationships to extant Dryopteridaceae.• Methods: The morphology of the fossil was compared with descriptions of extant ferns, resulting in it being tentatively assigned to the bolbitidoid fern genus Elaphoglossum. This assignment was confirmed by reconstructing the evolution of the morphological characters preserved in the inclusion on a molecular phylogeny of 158 extant bolbitidoid ferns. To assess the morphology-based assignment of the fossil to Elaphoglossum, we examined DNA-calibrated divergence time estimates against the age of the amber deposits from which it came.• Key results: The fossil belongs to Elaphoglossum and is the first of a bolbitidoid fern. Its assignment to a particular section of Elaphoglossum could not be determined; however, sects. Lepidoglossa, Polytrichia, and Setosa can be discounted because the fossil lacks subulate scales or scales with acicular marginal hairs. Thus, the fossil might belong to either sects. Amygdalifolia, Wrightiana, Elaphoglossum, or Squamipedia or to an extinct lineage.• Conclusions: The discovery of a Miocene Elaphoglossum fossil provides remarkable support to current molecular clock-based estimates of the diversification of these ferns.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
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► First global phylogeny of an epiphyllous liverwort genus, Diplasiolejeunea. ► Molecular phylogenies are not congruent with current supraspecific classifications. ► Topologies ...support a deep split into a Neotropical and a Paleotropical clade. ► Paleotropical clade structured into Australasian, Asian and Afromadacascan lineages. ► Evidence for speciation along altitudinal gradients and pantropical species ranges.
Diplasiolejeunea is a pantropical, epiphytic genus of leafy liverworts that occurs from the lowlands to more than 4000m altitude. Phylogenetic analyses of a molecular dataset consisting of three markers (nuclear ribosomal ITS region, plastidic trnL-F region and rbcL gene) and 122 accessions (plus two outgroups, Colura and Cololejeunea) indicate that the evolutionary diversity of Diplasiolejeunea is underestimated by current morphology-based classification. Four morphologically semi-cryptic species have been recovered. The molecular phylogenies support a deep split into a Neotropical and a Paleotropical clade, the latter structured into Australasian, Asian and Afromadacascan lineages. Presented results confirm the ranges of two pantropical species (D. cavifolia, D. rudolphiana), provide evidence for dispersal from the Neotropics into the Paleotropics, indicate speciation along altitudinal gradients and demonstrate extensive morphological homoplasy. We propose a revised supraspecific classification of Diplasiolejeunea into a predominantly Paleotropical subgenus Physolejeunea and predominantly Neotropical subgenera Austrolejeuneopsis and Diplasiolejeunea, the former containing mainly epiphytic species, the latter mainly epiphylls. Several clades are supported by combinations of morphological character states, and could be assigned to sections at some later point. This is the first comprehensive phylogeny of a largely epiphyllous genus of liverworts.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, UPUK
More than 200 research papers on the molecular phylogeny and phylogenetic biogeography of bryophytes have been published since the beginning of this millenium. These papers corroborated assumptions ...of a complex genetic structure of morphologically circumscribed bryophytes, and raised reservations against many morphologically justified species concepts, especially within the mosses. However, many molecular studies allowed for corrections and modifications of morphological classification schemes. Several studies reported that the phylogenetic structure of disjunctly distributed bryophyte species reflects their geographical ranges rather than morphological disparities. Molecular data led to new appraisals of distribution ranges and allowed for the reconstruction of refugia and migration routes. Intercontinental ranges of bryophytes are often caused by dispersal rather than geographical vicariance. Many distribution patterns of disjunct bryophytes are likely formed by processes such as short distance dispersal, rare long distance dispersal events, extinction, recolonization and diversification.
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FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
In this study, we evaluate the impact of fossil assignments and different models of calibration on divergence time estimates carried out as Bayesian analyses. Estimated ages from preceding studies ...and liverwort inclusions from Baltic amber are used as constraints on a molecular phylogeny of Cephaloziineae (Jungermanniopsida) obtained from sequences of two chloroplast coding regions: rbcL and psbA. In total, the comparison of 12 different analyses demonstrates that an increased reliability of the chronograms is linked to the number of fossils assigned and to the accuracy of their assignments. Inclusion of fossil constraints leads to older ages of most crown groups, but has no influence on lineage through time plots suggesting a nearly constant accumulation of diversity since the origin of Cephaloziineae in the early to Middle Jurassic. Our results provide a note of caution regarding the interpretation of chronograms derived from DNA sequence variation of extant species based on a single calibration point and/or low accuracy of the assignment of fossils to nodes in the phylogeny.
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BFBNIB, DOBA, EMUNI, FIS, FZAB, GEOZS, GIS, IJS, IMTLJ, IZUM, KILJ, KISLJ, MFDPS, NLZOH, NMLJ, NUK, OILJ, PILJ, PNG, SAZU, SBCE, SBJE, SBMB, SBNM, UILJ, UKNU, UL, UM, UPUK, VKSCE, ZAGLJ
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► Shared haplotypes of
Ptilidium ciliare and
Ptilidium pulcherrimum between Europe and North America. ► Presence of Southern Hemisphere haplotypes of
Ptilidium ciliare in the Northern ...Hemisphere. ► Recent long distance dispersal rather than Gondwanan vicariance explains the bipolar disjunct range of
Ptilidium ciliare. ► Morphologically cryptic speciation in several
Ptilidium lineages.
The small, phylogenetically isolated liverwort genus
Ptilidium has been regarded as of cool-Gondwanic origin with the bipolar, terrestrial
Ptilidium ciliare giving rise to the Northern Hemisphere epiphytes
Ptilidium pulcherrimum and
Ptilidium californicum. This hypothesis is examined using a dataset including three chloroplast DNA regions from 134
Ptilidium accessions and one accession each of its closest relatives
Trichocoleopsis and
Neotrichocolea. Maximum likelihood and parsimony analyses point to a close relationship between
P. ciliare and
P. pulcherrimum, whereas
P. californicum is placed sister to the remainder of the genus, separated by a long branch. Haplotype analysis and our phylogeny indicate the presence of Southern Hemisphere haplotypes of
P. ciliare in the Northern Hemisphere, and shared haplotypes of
P. ciliare and
P. pulcherrimum between Europe and North America. Based on our findings, we reject the Gondwana-scenario and propose recent long distance dispersal as an explanation for the bipolar disjunct range.
Ptilidium ciliare is resolved as paraphyletic with
P. pulcherrimum nested within it. An isolated
Ptilidium lineage with the morphology of
P. ciliare from the Himalaya region likely represents a hitherto unrecognized cryptic species.
Ptilidium pulcherrimum splits into a Japanese clade and a clade with accessions from Europe and North America.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, UPUK
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► We report the first global phylogeny of Scapania based on 184 accessions and three molecular markers. ► Molecular phylogenies are not congruent with current supraspecific ...classifications. ► We propose a classification into six subgenera and rearrangements within numerous sections. ► Scapania irrigua is paraphyletic with several species nested in it. ► Southern lineages are nested in northern hemispheric clades.
Scapania is a northern temperate genus with a few disjunctions in the south. Despite receiving considerable attention, the supraspecific classification of this genus remains unsatisfactorily solved. We use three molecular markers (nrITS, cpDNA trnL-F region, atpB-rbcL spacer) and 175 accessions belonging to 50 species (plus eight outgroup taxa) to estimate the phylogeny and to test current classification systems. Our data support the classification of Scapania into six rather than three subgenera, rearrangements within numerous sections, and inclusion of Macrodiplophyllum microdontum. Scapania species with a plicate perianth form three early diverging lineages; the most speciose subgenus, Scapania s.str., represents a derived clade. Most morphological species concepts are supported by the molecular topologies but classification of sect. Curtae requires further study. Southern lineages are nested in northern hemispheric clades. Palearctic–Nearctic distribution ranges are supported for several species.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UL, UM, UPCLJ, UPUK
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