The timescale of early land plant evolution Morris, Jennifer L.; Puttick, Mark N.; Clark, James W. ...
Proceedings of the National Academy of Sciences - PNAS,
03/2018, Letnik:
115, Številka:
10
Journal Article
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Establishing the timescale of early land plant evolution is essential for testing hypotheses on the coevolution of land plants and Earth’s System. The sparseness of early land plant megafossils and ...stratigraphic controls on their distribution make the fossil record an unreliable guide, leaving only the molecular clock. However, the application of molecular clock methodology is challenged by the current impasse in attempts to resolve the evolutionary relationships among the living bryophytes and tracheophytes. Here, we establish a timescale for early land plant evolution that integrates over topological uncertainty by exploring the impact of competing hypotheses on bryophyte−tracheophyte relationships, among other variables, on divergence time estimation. We codify 37 fossil calibrations for Viridiplantae following best practice. We apply these calibrations in a Bayesian relaxed molecular clock analysis of a phylogenomic dataset encompassing the diversity of Embryophyta and their relatives within Viridiplantae. Topology and dataset sizes have little impact on age estimates, with greater differences among alternative clock models and calibration strategies. For all analyses, a Cambrian origin of Embryophyta is recovered with highest probability. The estimated ages for crown tracheophytes range from Late Ordovician to late Silurian. This timescale implies an early establishment of terrestrial ecosystems by land plants that is in close accord with recent estimates for the origin of terrestrial animal lineages. Biogeochemical models that are constrained by the fossil record of early land plants, or attempt to explain their impact, must consider the implications of a much earlier, middle Cambrian–Early Ordovician, origin.
During the 1920s, the botanist W. H. Lang set out to collect and investigate some very unpromising fossils of uncertain affinity, which predated the known geological record of life on land. His ...discoveries led to a landmark publication in 1937, ‘On the plant-remains from the Downtonian of England and Wales’, in which he revealed a diversity of small fossil organisms of great simplicity that shed light on the nature of the earliest known land plants. These and subsequent discoveries have taken on new relevance as botanists seek to understand the plant genome and the early evolution of fundamental organ systems. Also, our developing knowledge of the composition of early land-based ecosystems and the interactions among their various components is contributing to our understanding of how life on land affects key Earth Systems (e.g. carbon cycle). The emerging paradigm is one of early life on land dominated by microbes, small bryophyte-like organisms and lichens. Collectively called cryptogamic covers, these are comparable with those that dominate certain ecosystems today. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.
The evolutionary emergence of land plant body plans transformed the planet. However, our understanding of this formative episode is mired in the uncertainty associated with the phylogenetic ...relationships among bryophytes (hornworts, liverworts, and mosses) and tracheophytes (vascular plants). Here we attempt to clarify this problem by analyzing a large transcriptomic dataset with models that allow for compositional heterogeneity between sites. Zygnematophyceae is resolved as sister to land plants, but we obtain several distinct relationships between bryophytes and tracheophytes. Concatenated sequence analyses that can explicitly accommodate site-specific compositional heterogeneity give more support for a mosses-liverworts clade, “Setaphyta,” as the sister to all other land plants, and weak support for hornworts as the sister to all other land plants. Bryophyte monophyly is supported by gene concatenation analyses using models explicitly accommodating lineage-specific compositional heterogeneity and analyses of gene trees. Both maximum-likelihood analyses that compare the fit of each gene tree to proposed species trees and Bayesian supertree estimation based on gene trees support bryophyte monophyly. Of the 15 distinct rooted relationships for embryophytes, we reject all but three hypotheses, which differ only in the position of hornworts. Our results imply that the ancestral embryophyte was more complex than has been envisaged based on topologies recognizing liverworts as the sister lineage to all other embryophytes. This requires many phenotypic character losses and transformations in the liverwort lineage, diminishes inconsistency between phylogeny and the fossil record, and prompts re-evaluation of the phylogenetic affinity of early land plant fossils, the majority of which are considered stem tracheophytes.
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•Early land plant relationships are extremely uncertain•We resolve the “Setaphyta” clade of liverworts plus mosses•The simple body plan of liverworts results from loss of ancestral characters•The ancestral land plant was more complex
Puttick et al. resolve a “Setaphyta” clade uniting liverworts and mosses and support for bryophyte monophyly. Their results indicate that the ancestral land plant was more complex than has been envisaged based on phylogenies recognizing liverworts as the sister lineage to all other embryophytes.
Lichenization is assumed to be a very ancient mode of fungal nutrition, but fossil records are rare. Here we describe two fragments of exceptionally preserved, probably charred, lichen thalli with ...internal stratification. Cyanolichenomycites devonicus has a cyanobacterial and Chlorolichenomycites salopensis a unicellular, presumably green algal photobiont. Fruiting bodies are missing. Cyanolichenomycites devonicus forms asexual spores in a pycnidium.
All specimens were examined with scanning electron microscopy techniques. The fossils were extracted by maceration. Extant lichens and free-living cyanobacteria were either experimentally charcoalified for comparison or conventionally prepared.
Based on their septate hyphal structure, both specimens are tentatively interpreted as representatives of the Pezizomycotina (Ascomycota). Their presence in 415 million yr (Myr) old rocks from the Welsh Borderland predates existing Late Cretaceous records of pycnidial conidiomata by some 325 Myr and Triassic records of lichens with broadly similar organization by some 195 Myr.
These fossils represent the oldest known record of lichens with symbionts and anatomy as typically found in morphologically advanced taxa today. The latter does not apply to Winfrenatia reticulata, the enigmatic crustose lichen fossil from the Lower Devonian, nor to presumed lichen-like organisms such as the Cambrian Farghera robusta or to the Lower Devonian Spongiophyton minutissimum.
50 I. 50 II. 52 III. 53 IV. 66 V. 71 VI. 72 VII. 74 75 References 75 SUMMARY: Cryptospores, recovered from Ordovician through Devonian rocks, differ from trilete spores in possessing distinctive ...configurations (i.e. hilate monads, dyads, and permanent tetrads). Their affinities are contentious, but knowledge of their relationships is essential to understanding the nature of the earliest land flora. This review brings together evidence about the source plants, mostly obtained from spores extracted from minute, fragmented, yet exceptionally anatomically preserved fossils. We coin the term ‘cryptophytes’ for plants that produced the cryptospores and show them to have been simple terrestrial organisms of short stature (i.e. millimetres high). Two lineages are currently recognized. Partitatheca shows a combination of characters (e.g. spo‐rophyte bifurcation, stomata, and dyads) unknown in plants today. Lenticulatheca encompasses discoidal sporangia containing monads formed from dyads with ultrastructure closer to that of higher plants, as exemplified by Cooksonia. Other emerging groupings are less well characterized, and their precise affinities to living clades remain unclear. Some may be stem group embryophytes or tracheophytes. Others are more closely related to the bryophytes, but they are not bryophytes as defined by extant representatives. Cryptophytes encompass a pool of diversity from which modern bryophytes and vascular plants emerged, but were competitively replaced by early tracheophytes. Sporogenesis always produced either dyads or tetrads, indicating strict genetic control. The long‐held consensus that tetrads were the archetypal condition in land plants is challenged.
The charcoalified fragment of the dorsiventrally organized, internally stratified presumed green algal lichen Chlorolichenomycites salopensis from the Lower Devonian Lochkovian strata in the Welsh ...Borderland carries bacterial colonies on the upper surface, i.e. the cortex, and actinobacterial filaments in the medulla underneath the photobiont layer. Moreover relatively thin hyphae of presumed endolichenic fungi were found. As in extant lichens, which are best regarded as consortia with an unknown number of participants, this internally stratified, fossil thallus fragment of a presumed green algal lichen harbours a diverse microbial community.
•Chlorolichenomycites salopensis was recovered from Lower Devonian Lochkovian strata.•It is the oldest known green algal lichen with stratified, dorsiventrally organized thallus.•We found bacteria on its surface and actinobacteria and nonlichenized fungi in its interior.•Endolichenic actinobacteria and fungi are very common and widespread in extant lichens.•Their occurrence in a 415 Myr old thallus fragment supports the view of a long cohabitation.
The ca. 1.38 billion years (Ga) old Roper Group of the McArthur Basin, northern Australia, is one of the most extensive Proterozoic hydrocarbon‐bearing units. Organic‐rich black siltstones from the ...Velkerri Formation were deposited in a deep‐water sequence and were analysed to determine their organic geochemical (biomarker) signatures, which were used to interpret the microbial diversity and palaeoenvironment of the Roper Seaway. The indigenous hydrocarbon biomarker assemblages describe a water column dominated by bacteria with large‐scale heterotrophic reworking of the organic matter in the water column or bottom sediment. Possible evidence for microbial reworking includes a large unresolved complex mixture (UCM), high ratios of mid‐chained and terminally branched monomethyl alkanes relative to n‐alkanes—features characteristic of indigenous Proterozoic bitumen. Steranes, biomarkers for single‐celled and multicellular eukaryotes, were below detection limits in all extracts analysed, despite eukaryotic microfossils having been previously identified in the Roper Group, albeit largely in organically lean shallower water facies. These data suggest that eukaryotes, while present in the Roper Seaway, were ecologically restricted and contributed little to export production. The 2,3,4‐ and 2,3,6‐trimethyl aryl isoprenoids (TMAI) were absent or in very low concentration in the Velkerri Formation. The low abundance is primary and not caused by thermal destruction. The combination of increased dibenzothiophene in the Amungee Member of the Velkerri Formation and trace metal redox geochemistry suggests that degradation of carotenoids occurred during intermittent oxygen exposure at the sediment–water interface and/or the water column was rarely euxinic in the photic zone and likely only transiently euxinic at depth. A comparison of this work with recently published biomarker and trace elemental studies from other mid‐Proterozoic basins demonstrates that microbial environments, water column geochemistry and basin redox were heterogeneous.
Descriptions of Lower Devonian plants from Yunnan, South China, have revolutionized concepts of diversity and disparity in tracheophytes soon after they became established on land. Sichuan ...assemblages have received little attention since their discovery almost 25 years ago and require revision. With this objective, fieldwork involving detailed logging and collection of fossils was undertaken in the Longmenshan Mountain Region, Jiangyou County and yielded the two new taxa described here. They are preserved as coalified compressions and impressions that allowed morphological but not anatomical analyses. Yanmenia (Zosterophyllum) longa comb nov is based on numerous rarely branching shoots with enations resembling lycophyte microphylls, without evidence for vasculature. The presence of sporangia is equivocal making assignation to the Lycopsida conjectural. The plant was recently described as a zosterophyll, but lacks strobili. These are present in the second plant and comprise bivalved sporangia. The strobili terminate aerial stems which arise from a basal axial complex displaying diversity in branching including H- and K- forms. These features characterise the Zosterophyllopsida, although the plant differs from Zosterophyllum in valve shape. Comparisons indicate greatest similarities to the Lower Devonian Guangnania cuneata, from Yunnan, but differences, particularly in the nature of the sporangium border, require the erection of a new species, G. minor. Superficial examination of specimens already published indicate a high degree of endemism at both species and generic level, while this study shows that Yanmenia is confined to Sichuan and Guangnania is one of the very few genera shared with Yunnan, where assemblages also show a high proportion of further endemic genera. Such provincialism noted in the Chinese Lower Devonian is explained by the palaeogeographic isolation of the South China plate, but this cannot account for differences/endemism between the Sichuan and Yunnan floras. Such an enigma demands further integrated geological, palaeobotanical and palynological studies.
Asphaltites, long known to strand along the coastline of southern Australia and as distantly as New Zealand and Macquarie Island, are widely regarded as artefacts of submarine oil seepage. Their ...remarkably uniform composition suggests a common source: marine shale containing sulphur-rich Type II kerogen, probably deposited during an Early Cretaceous oceanic anoxic event (OAE). Suitable hydrocarbon kitchens may exist in the offshore Bight and Otway basins. The physical character of the asphaltites, including laminations and flow structures, and their degree of alteration, which is not the result of biodegradation or extensive water washing, suggest an origin from subsurface tar mats subsequently exposed by the incision of submarine canyons, with the possible formation of asphaltic volcanoes. API gravities of 4–18° impart quasi-neutral buoyancy, implying many asphaltites were submerged drifters prior to stranding, their degree of weathering reflecting, at least in part, the residence time in the marine environment. For any individual asphaltite specimen, this will depend on the proximity of the seafloor seep to the stranding site, an important consideration when attempting to locate their point of origin.
This study investigates the hydrocarbon biomarker signatures and n-alkane δ13C profiles of asphaltite specimens from stranding sites on the Eyre Peninsula (n = 2), Kangaroo Island (n = 4) and the Limestone Coast (n = 3), South Australia, and the south island of New Zealand (n = 2). Sub-samples of the interior and weathered surface of each specimen were analysed. No distinction could be made between strandings based on their source-dependent molecular and isotopic signatures, confirming their common origin. Comparison of the interior and exterior sub-samples revealed subtle although consistent differences. Given their degree of degradation and isotopic variance, these Australasian asphaltites seem to be products of low intensity seeps in the Ceduna Sub-basin of the Bight Basin and/or Morum Sub-basin of the Otway Basin.
•Study of biomarker and δ13C profiles of asphaltites from 4 disparate localities.•Degradation patterns imply subsurface alteration probably due to tar mat formation.•Analyses confirm their common origin despite their disparate stranding localities.•Biomarker and δ13C signatures show consistent variations with spatial distribution.•Probably sourced from Ceduna Sub-basin (Bight) and/or Morum Sub-basin (Otway) seeps.
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