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The origin of plants and their colonization of land fundamentally transformed the terrestrial environment. Here we elucidate the basis of this formative episode in Earth history through patterns of lineage, gene and genome evolution. We use new fossil calibrations, a relative clade age calibration (informed by horizontal gene transfer) and new phylogenomic methods for mapping gene family origins. Distinct rooting strategies resolve tracheophytes (vascular plants) and bryophytes (non-vascular plants) as monophyletic sister groups that diverged during the Cambrian, 515–494 million years ago. The embryophyte stem is characterized by a burst of gene innovation, while bryophytes subsequently experienced an equally dramatic episode of reductive genome evolution in which they lost genes associated with the elaboration of vasculature and the stomatal complex. Overall, our analyses reveal that extant tracheophytes and bryophytes are both highly derived from a more complex ancestral land plant. Understanding the origin of land plants requires tracing character evolution across a diversity of modern lineages.

Molecular time trees indicating that embryophytes originated around 500 million years ago (Ma) during the Cambrian are at odds with the record of fossil plants, which first appear in the mid-Silurian almost 80 million years later. This time gap has been attributed to a missing fossil plant record, but that attribution belies the case for fossil spores. Here, we describe a Tremadocian (Early Ordovician, about 480 Ma) assemblage with elements of both Cambrian and younger embryophyte spores that provides a new level of evolutionary continuity between embryophytes and their algal ancestors. This finding suggests that the molecular phylogenetic signal retains a latent evolutionary history of the acquisition of the embryophytic developmental genome, a history that perhaps began during Ediacaran-Cambrian time but was not completed until the mid-Silurian (about 430 Ma).

The relationships among the four major embryophyte lineages (mosses, liverworts, hornworts, vascular plants) and the timing of the origin of land plants are enigmatic problems in plant evolution. Here, we resolve the monophyly of bryophytes by improving taxon sampling of hornworts and eliminating the effect of synonymous substitutions. We then estimate the divergence time of crown embryophytes based on three fossil calibration strategies, and reveal that maximum calibration constraints have a major effect on estimating the time of origin of land plants. Moreover, comparison of priors and posteriors provides a guide for evaluating the optimal calibration strategy. By considering the reliability of fossil calibrations and the influences of molecular data, we estimate that land plants originated in the Precambrian (980–682 Ma), much older than widely recognized. Our study highlights the important contribution of molecular data when faced with contentious fossil evidence, and that fossil calibrations used in estimating the timescale of plant evolution require critical scrutiny.

— Separate cladistic analyses of the green algae, liverworts, and hornworts are presented. Classificatory and evolutionary implications of these analyses, in addition to our previously published cladistic analyses of mosses and the embryophytes as a whole, are discussed. The embryophytes are monophyletic, and are part of a larger monophyletic group that includes some of the green algae (the “charophytes”). Important evolutionary transformations in the early phylogeny of the land plants include: (1) retention of the zygote on the haploid plant (gametophyte), with the sporophyte generation arising de novo by delaying meiosis, (2) independent elaboration of an elongate sporophyte in some liverworts, some hornworts, and in the moss-tracheophyte clade, (3) independent origin of radial (axial) symmetry in the gametophyte in some liverworts and in the moss-tracheophyte clade, (4) independent origin of leaves on the gametophyte in some liverworts and in mosses, and (5) the unique development of a branching sporophyte with multiple sporangia in the tracheophytes.

Summary ●Cells are continuously exposed to chemical signals that they must discriminate between and respond to appropriately. In embryophytes, the leucine‐rich repeat receptor‐like kinases (LRR‐RLKs) are signal receptors critical in development and defense. LRR‐RLKs have diversified to hundreds of genes in many plant genomes. Although intensively studied, a well‐resolved LRR‐RLK gene tree has remained elusive.●To resolve the LRR‐RLK gene tree, we developed an improved gene discovery method based on iterative hidden Markov model searching and phylogenetic inference. We used this method to infer complete gene trees for each of the LRR‐RLK subclades and reconstructed the deepest nodes of the full gene family.●We discovered that the LRR‐RLK gene family is even larger than previously thought, and that protein domain gains and losses are prevalent. These structural modifications, some of which likely predate embryophyte diversification, led to misclassification of some LRR‐RLK variants as members of other gene families. Our work corrects this misclassification.●Our results reveal ongoing structural evolution generating novel LRR‐RLK genes. These new genes are raw material for the diversification of signaling in development and defense. Our methods also enable phylogenetic reconstruction in any large gene family.

Premise Phylogenetic trees of bryophytes provide important evolutionary context for land plants. However, published inferences of overall embryophyte relationships vary considerably. We performed phylogenomic analyses of bryophytes and relatives using both mitochondrial and plastid gene sets, and investigated bryophyte plastome evolution. Methods We employed diverse likelihood‐based analyses to infer large‐scale bryophyte phylogeny for mitochondrial and plastid data sets. We tested for changes in purifying selection in plastid genes of a mycoheterotrophic liverwort (Aneura mirabilis) and a putatively mycoheterotrophic moss (Buxbaumia), and compared 15 bryophyte plastomes for major structural rearrangements. Results Overall land‐plant relationships conflict across analyses, generally weakly. However, an underlying (unrooted) four‐taxon tree is consistent across most analyses and published studies. Despite gene coverage patchiness, relationships within mosses, liverworts, and hornworts are largely congruent with previous studies, with plastid results generally better supported. Exclusion of RNA edit sites restores cases of unexpected non‐monophyly to monophyly for Takakia and two hornwort genera. Relaxed purifying selection affects multiple plastid genes in mycoheterotrophic Aneura but not Buxbaumia. Plastid genome structure is nearly invariant across bryophytes, but the tufA locus, presumed lost in embryophytes, is unexpectedly retained in several mosses. Conclusions A common unrooted tree underlies embryophyte phylogeny, [(liverworts, mosses), (hornworts, vascular plants)]; rooting inconsistency across studies likely reflects substantial distance to algal outgroups. Analyses combining genomic and transcriptomic data may be misled locally for heavily RNA‐edited taxa. The Buxbaumia plastome lacks hallmarks of relaxed selection found in mycoheterotrophic Aneura. Autotrophic bryophyte plastomes, including Buxbaumia, hardly vary in overall structure.

The AVRPPHB SUSCEPTIBLE1 (PBS1) and RESISTANCE TO PSEUDOMONAS SYRINGAE 5 (RPS5) proteins are involved in signal transduction to evoke innate plant immune response. In Arabidopsis, PBS1 is cleaved by the AvrPphB (Pseudomonas phaseolicola Avirulence protein B) protease, activating RPS5 and turning in a hypersensitive response (HR). We searched for PBS1 orthologs to trace their origin and evolution. PBS1 orthologs were found in embryophytes and in other plant taxa but with lower similarity. PBS1 phylogenetic analysis indicates high divergence, suggesting that the decoy function described for Arabidopsis PBS1 might be associated with a small fraction of orthologs. Ancestral reconstruction analysis suggests an elevated diversity in the amino acid sequence within the described motifs. All the orthologs contain the conserved PBS1 kinase subdomains, whereas the cleavage motif is present in several embryophyte orthologs but absent in most other taxa. The putative resistance recognition motifs in PBS1 orthologs are highly diverse. PBS1 cleavage site motif is exposed in some 3D structure predictions, whereas it is not in others, suggesting different modes of regulation and functions in PBS1 orthologs. Our findings suggest that PBS1 originated in the lineage that gave rise to embryophytes, with the angiosperm sequences forming a separate clade from pteridophyte proteins.

Over the last 470 million years, plant evolution has seen major evolutionary transitions such as the move from water to land and the origins of vascular tissues, seeds, and flowers[1]. These have resulted in the evolution of terrestrial flora that has shaped modern ecosystems, and the diversification of the Plant Kingdom, Viridiplantae, into over 374,000 described species[2]. Each of these transitions was accompanied by the gain and loss of genes in plant genomes. For example, whole genome duplications are known to be fundamental to the origins of both seed and flowering plants [3,4]. With the ever-increasing quality and quantity of whole genome data, evolutionary insight into origins of distinct plant groups using comparative genomic techniques is now feasible. Here, using an evolutionary genomics pipeline to compare 208 complete genomes, we analyse the gene content of the ancestral genomes of the last common ancestor of land plants and all other major groups of plant. This approach reveals an unprecedented level of fundamental genomic novelties in two nodes related to the origin of land plants, the first in the origin of streptophytes during the Ediacaran and another in the ancestor of land plants in the Ordovician. Our findings highlight the biological processes that evolved with the origin of land plants and emphasise the importance of conserved gene novelties in plant diversification. Comparisons to other eukaryotic studies suggest a separation of the genomic origins of multicellularity and terrestralisation in plants.

ECERIFERUM1 (CER1) and ECERIFERUM3 (CER3) are key genes in synthesis of alkanes, a major component of cuticular waxes in land plants. The genes share extensive similarity, including the N-terminal (ERG3/FAH) and C-terminal (WAX2) domains. This study, traces the origin, evolutionary history, phylogenetic relationships and variation in copy number of the two genes within and beyond the Viridiplantae (green plants). Protein homologs of both CER1 and CER3 were identified across most Embryophyta (land plants), a single homolog (CER1/3) in charophytes and prasinophytes, and none in the other green, red or brown algae. Ancestral state reconstructions in 100 sequenced Archaeplastida using presence/absence of CER1/3 family genes revealed that the CER1/3 gene probably originated in the common ancestor of Viridiplantae. Phylogenetic analysis of CER1 and CER3 protein sequences from 146 plant species strongly suggests that the two genes originated by duplication of CER1/3 in the ancestral embryophyte. The evolution of CER1 and CER3 genes involved differential divergence of the two domains. Outside Embryophyta, CER1/3 similar sequences identified in diatoms and a cryptophyte, were the closest relatives of the CER1/3 family proteins. Proteins harbouring WAX2-wxAR (WAX2 associated region) similar regions were identified in proteins of bacteria, Archaea, cryptophytes, dinoflagellates and Stramenopiles. The independent existence of both ERG3/FAH and WAX2-wxAR domains in diverse lineages strongly points to the origin of CER1/3 gene in green plants by the fusion of pre-existing domains.

We hereby review the perception and responses to the stress hormone Abscisic acid (ABA), along the trajectory of 500M years of plant evolution, whose understanding may resolve how plants acquired this signaling pathway essential for the colonization of land. ABA levels rise in response to abiotic stresses, coordinating physiological and metabolic responses, helping plants survive stressful environments. In land plants, ABA signaling cascade leads to growth arrest and large-scale changes in transcript levels, required for coping with environmental stressors. This response is regulated by a PYRABACTIN RESISTANCE 1-like (PYL)-PROTEIN PHOSPHATASE 2C (PP2C)-SNF1-RELATED PROTEIN KINASE 2 (SnRK2) module, that initiates phosphor-activation of transcription factors and ion channels. The enzymatic portions of this module (phosphatase and kinase) are functionally conserved from streptophyte algae to angiosperms, whereas the regulatory component -the PYL receptors, putatively evolved in the common ancestor of Zygnematophyceae and embryophyte as a constitutive, ABA-independent protein, further evolving into a ligand-activated receptor at the embryophyta. This evolutionary process peaked with the appearance of the strictly ABA-dependent subfamily III stress-triggered angiosperms' dimeric PYL receptors. The emerging picture is that the ancestor of land plants and its predecessors synthesized ABA, as its biosynthetic pathway is conserved between ancestral and current day algae. Despite this ability, it was only the common ancestor of land plants which acquired the hormonal-modulation of PYL activity by ABA. This raises several questions regarding both ABA's function in ABA-non-responsive organisms, and the evolutionary aspects of the ABA signal transduction pathway.

A relatively rich association of six species of embryophyte plants is known from the upper Silurian of the Prague Basin (Bohemian Massif, Czech Republic). All stratigraphically controlled specimens come from the Neocolonograptus parultimus-Neocolonograptus ultimus Zone of the Přidoli. A new genus and species, Tichavekia grandis Psenicka, Sakala et Kraft, is established. The new combination Aberlemnia bohemica (Schweitzer) Sakala, Psenicka et Kraft, comb. nov. illustrates its Lycophytina affinity. The distribution and taphonomy of these plants in the Prague Basin indicates the proximity of exposed land interpreted to be islands of volcanic origin. Two local associations show primary differences in vegetated areas of the islands and apparent ecological responses of the primitive vascular plants to their habitats. Suitable environments in the coastal zones of volcanic islands in the Prague Basin, which was situated at the outer periphery of a broad Gondwanan shelf, likely represent significant transfer points for an initial dispersion and a first expansion of land plants. Repeated vegetation of the islands is inferred from the discrete distribution of land plants at different stratigraphic levels. These changes are explained partly in context of local changes of the island relief, and partly as responses to global climatic and sea level fluctuations, and fit environmental dynamics of islands of volcanic origin in general. This Initial Plant Diversification and Dispersal Event (IPDDE) is of considerable significance in the evolution of plants documented from the Bohemian Massif.

Significance The evolution of land plants from algae is an age-old question in biology. The entire terrestrial flora stems from a grade of algae, the streptophyte algae. Recent phylogenomic studies have pinpointed the Zygnematophyceae as the modern-day streptophyte algal lineage that is most closely related to the algal land plant ancestor. Here, we provide insight into the biology of this ancestor that might have aided in its conquest of land. Specifically, we uncover the existence of stress-signaling pathways and the potential for intimate plastid-nucleus communication. Plastids act as environmental sensors in land plants; our data suggest that this feature was present in a common ancestor they shared with streptophyte algae., Streptophytes are unique among photosynthetic eukaryotes in having conquered land. As the ancestors of land plants, streptophyte algae are hypothesized to have possessed exaptations to the environmental stressors encountered during the transition to terrestrial life. Many of these stressors, including high irradiance and drought, are linked to plastid biology. We have investigated global gene expression patterns across all six major streptophyte algal lineages, analyzing a total of around 46,000 genes assembled from a little more than 1.64 billion sequence reads from six organisms under three growth conditions. Our results show that streptophyte algae respond to cold and high light stress via expression of hallmark genes used by land plants (embryophytes) during stress–response signaling and downstream responses. Among the strongest differentially regulated genes were those associated with plastid biology. We observed that among streptophyte algae, those most closely related to land plants, especially Zygnema, invest the largest fraction of their transcriptional budget in plastid-targeted proteins and possess an array of land plant-type plastid-nucleus communication genes. Streptophyte algae more closely related to land plants also appear most similar to land plants in their capacity to respond to plastid stressors. Support for this notion comes from the detection of a canonical abscisic acid receptor of the PYRABACTIN RESISTANCE (PYR/PYL/RCAR) family in Zygnema, the first found outside the land plant lineage. We conclude that a fine-tuned response toward terrestrial plastid stressors was among the exaptations that allowed streptophytes to colonize the terrestrial habitat on a global scale.

Cryptospores from the Dapingian–Darriwilian Kanosh Shale at Fossil Mountain, Utah, USA, occur as tetrads, dyads, irregular clusters, or planar sheets of spore dyads. These spore ‘thalli’ are placed into the new taxon Grododowon orthogonalis gen. et sp. nov. based on the nature of division patterning and gross overall shape. The antithetic hypothesis of embryophyte origins dictates that spores evolved first and that the vegetative sporophyte evolved later via mitotic cell divisions that preceded meiosis and spore formation. We interpret the planar spore sheets of Grododowon gen. nov. to have formed via the co-option of a prior vegetative gametophytic developmental pattern which was expressed in the zygote, resulting in a two-dimensional, thalloid bauplan. However, the ploidy of the resultant ‘spores’ as haploid is necessarily conjectural, and this pattern of growth is clearly not the ancestral condition in the streptophyte lineage that gave rise to the first axial plant sporophyte.

Cryptospores from the Hanadir Member, Qasim Formation (Darriwilian) examined with transmission electron microscopy (TEM), show two distinct kinds of walls: homogeneous and lamellated. A recently described tetrad from the Hanadir Member, Cryptotetras erugata Strother, Traverse & Vecoli, and the dyad, Pseudodyadospora sp. cf. P. laevigata Johnson, possess the homogeneous wall type, matching that seen in younger, Silurian species of Tetrahedraletes from Laurentia. The lamellated wall structure of a single Hanadir Member monad is also described. Homogeneous walls are characteristic of extant embryophyte sporoderm, which is actively secreted by a tapetum. This implies that these Ordovician cryptospores developed within a sporangium, although fossilized sporangia of this antiquity are not known. Lamellated walls occur in cryptospore dyads found in Siluro-Devonian cryptophytes (as well as fragments of presumed cryptophytes as old as the Ordovician), but this type of wall is also common in Cambrian cryptospores. Therefore, lamellate spore walls appear to be have evolved in algal (charophytic) lineages well in advance of the first plants that possessed an upright axial sporophyte. Isometric, tetrahedral tetrads, like Cryptotetras and Tetrahedraletes, which dominate the Hanadir Member assemblage, express a geometry that reflects simultaneous cytokinesis during sporogenesis. Thus, the Hanadir Member specimens present what we interpret as the earliest evidence of simultaneous meiosis occurring inside a sporangium that possessed an active tapetum.

The origin of land plants from algae is a long-standing question in evolutionary biology. It is becoming increasingly clear that many characters that were once assumed to be 'embryophyte specific' can in fact be found in their closest algal relatives, the streptophyte algae. One such case is the phenylpropanoid pathway. While biochemical data indicate that streptophyte algae harbor lignin-like components, the phenylpropanoid core pathway, which serves as the backbone of lignin biosynthesis, has been proposed to have arisen at the base of the land plants. Here we revisit this hypothesis using a wealth of new sequence data from streptophyte algae. Tracing the biochemical pathway towards lignin biogenesis, we show that most of the genes required for phenylpropanoid synthesis and the precursors for lignin production were already present in streptophyte algae. Nevertheless, phylogenetic analyses and protein structure predictions of one of the key enzyme classes in lignin production, cinnamyl alcohol dehydrogenase (CAD), suggest that CADs of streptophyte algae are more similar to sinapyl alcohol dehydrogenases (SADs). This suggests that the end-products of the pathway leading to lignin biosynthesis in streptophyte algae may facilitate the production of lignin-like compounds and defense molecules. We hypothesize that streptophyte algae already possessed the genetic toolkit from which the capacity to produce lignin later evolved in vascular plants.

The broadly accepted definition of algae is ‘oxygenic photosynthesisers other than embryophyte land plants’, and multicellular algae are generally termed macroalgae. The numbers of green, brown, and red algal species are estimated to be approximately 1100, 1500, and 4000–6000, respectively, worldwide. Embryophytes living in marine environments are mainly seagrasses, which are angiosperms adapted to nearshore areas with soft sediment. Seagrasses are less diverse than macroalgae; there are less than 60 species worldwide. The three groups of macroalgae have different subcellular structures and evolutionary histories. Green and red algae evolved from a common ancestral photosynthetic eukaryote, which acquired their plastids through primary endosymbiosis, in which a cyanobacterium was engulfed by an ancestral eukaryotic host cell. Brown algae evolved by secondary endosymbiosis, in which a red alga was engulfed by a eukaryotic host protist. Because of this, green and red algae have plastids with two envelope membranes, while brown algae have plastids with four envelope membranes. The green and red algae differ in their photosynthetic pigment composition, thylakoid structure and storage products. The three macroalgal groups became multicellular independently of each other from different unicellular ancestors. Both macroalgal and seagrass beds are very productive ecosystems as their photosynthesis per unit area can be the highest in the world. These macrophytes support biodiverse coastal ecosystems by providing energy and structural complexity.

The interiors of plants are colonized by diverse microorganisms that are referred to as endophytes. Endophytes have received much attention over the past few decades, yet many questions remain unanswered regarding patterns in their biodiversity at local to global scales. To characterize research effort to date, we synthesized results from ∼600 published studies. Our survey revealed a global research interest and highlighted several gaps in knowledge. For instance, of the seventeen biomes encompassed by our survey, seven were understudied and together composed only 7% of the studies that we considered. We found that fungal endophyte diversity has been characterized in at least one host from 30% of embryophyte families, while bacterial endophytes have been surveyed in hosts from only 10.5% of families. We complimented our survey with a vote counting procedure to determine endophyte richness patterns among plant tissue types. We found that variation in endophyte assemblages in above-ground tissues varied with host growth habit. Stems were the richest tissue in woody plants, whereas roots were the richest tissue in graminoids. For forbs, we found no consistent differences in relative tissue richness among studies. We propose future directions to fill the gaps in knowledge we uncovered and inspire further research.Originality-Significance StatementMuch remains to be learned regarding the biodiversity and distribution of the microbes that colonize the interiors of plants. Here, we surveyed approximately 600 publications to characterize gaps in knowledge and provide a roadmap for future research. We compared biomes, plant families, and geographical regions in terms of the research interest that they have garnered. Additionally, we synthesized published results and report that variation in endophyte richness among plant tissue types is a function of host growth habit. Stems were the richest tissue in woody plants, whereas roots were the richest tissue in graminoids. We hope to inspire research to fill the gaps in knowledge that we uncovered.

Adaptor protein complexes and the related complexes COPI and TSET function in packaging vesicles for transport among endomembrane compartments in eukaryotic cells. Differences in the complement of these complexes in lineages such as yeast and mammals as well as apicomplexan and kinetoplastid parasites via loss or duplication of subunits appears to reflect specialization in their respective trafficking systems. The model plant Arabidopsis thaliana possesses multiple paralogues for adaptor protein complex subunits, raising questions as to the timing and extent of these duplications in embryophytes (land plants). However, adaptor protein complex evolution in embryophytes is unexplored. Therefore, we analyzed genomes of diverse embryophytes and closely related green algae using extensive homology searches and phylogenetic analysis of 35 complex subunit proteins. The results reveal numerous paralogues, the vast majority of which, approximately 97%, arose from recent duplication events. This suggests that specialization of these protein complexes may occur frequently but independently in embryophytes.

SummaryMosses (Bryophyta) are a key group occupying important phylogenetic position for understanding land plant (embryophyte) evolution. The class Bryopsida represents the most diversified lineage and contains more than 95% of the modern mosses, whereas the other classes are by nature species-poor. The phylogeny of mosses remains elusive at present.Recurrent whole genome duplications have shaped the evolution trajectory of angiosperms, but little is known about the genome evolutionary history in mosses. It remains to be answered if there existed a historical genome duplication event associated with the species radiation of class Bryopsida.Here, the high-confidence moss phylogeny was generated covering major moss lineages. Two episodes of ancient genomic duplication events were elucidated by phylogenomic analyses, one in the ancestry of all mosses and another before the separation of the Bryopsida, Polytrichopsida and Tetraphidopsida, with estimated ages of the gene duplications clustered around 329 and 182 million year ago, respectively.The third episode of polyploidy event (termed ψ) was tightly associated with the early diversification of Bryopsida with an estimated age of ~87 million years. By scrutinizing the phylogenetic timing of duplicated syntelogs in Physcomitrella patens, the WGD1 and WGD2 events were confidently re-recognized as the ψ event and the Funarioideae duplication event (~65 mya), respectively. Together, our findings unveiled four episodes of polyploidy events in the evolutionary past of Physcomitrella patens.

Biomass allocation patterns have received substantial consideration, leading to the recognition of several 'universal' interspecific trends. Despite efforts to understand biomass partitioning among embryophytes, few studies have examined macroalgae that evolved independently, yet function ecologically in much the same ways as plants. Kelps allocate photosynthate among three organs (the blade(s), stipe(s) and holdfast) that are superficially convergent with organs of land plants, providing a unique opportunity to test the limits of 'universal' trends. In this study, we used an allometric approach to quantify interspecific biomass partitioning patterns in kelps and assess whether embryophyte-based predictions of biomass scaling can be applied to marine macrophytes that lack root-to-leaf hydraulic transport. Photosynthetic area and dry mass were found to scale to approximately the ¾ power and kelp biomass allocation patterns were shown to match closely to empirical measures of allometric scaling among woody plants. Larger kelp species were found to have increased relative stipe and holdfast mass than smaller species, highlighting important consequences of size for marine macroalgae. Our study provides insights into the evolution of size in the largest marine macrophytes and corroborates previous work suggesting that the morphology of divergent lineages of photoautotrophs may reflect similar selective pressures.

Premise of research. Climatically favorable conditions correspond with fossil evidence for dramatic Ordovician marine biodiversification, but coeval terrestrial biodiversity is less well understood. Although diverse Middle and Late Ordovician microfossils are interpreted as reproductive remains of early bryophyte-like land plants (consistent with molecular data indicating pre-Ordovician embryophyte origin), the vegetative structure of Ordovician plants remains mysterious, as do relationships to modern groups. Because distinctive fungal microfossils indicating land plant presence were previously reported from Ordovician carbonate deposits in Wisconsin, we examined another nearby outcrop for additional evidence of terrestrial biodiversification.Methodology. Replicate collections were made from well-understood 455–454 Ma Platteville Formation carbonates of relatively low porosity and hydraulic conductivity. We employed measures to avoid contamination, and organic remains extracted by acid maceration were cha...

International audience; This study aims to approximate an illustrative vision of the apparent temporal dynamics of land plant diversity during the terrestrialization process (i.e., the invasion of the land by plants). This problem has often been addressed through studying the Silurian–Devonian megafossil-based diversity patterns of land plants (embryophytes). However, the inclusion of the dispersed spore fossil record is essential for characterizing the first steps of this process (i.e., Eoembryophytic flora). Consequently, this new study includes both spore taxa and plant megafossils in order to discern the diversity trajectories from the first phases of early land plant diversification until the stabilization of early Carboniferous forests (i.e., Palaeophytic flora). The diversity patterns of the primary recognized plant lineages (i.e., Lycophyta and Euphyllophyta) have been also traced in order to discern the origin of the major fluctuations of plant megafossil-based diversity. Results reveal that while the dispersed spore diversity curve shows a sustained increase towards the end of the Devonian Period – with a first maximum towards the late Silurian – the megafossil embryophyte diversity curve is characterized by a set of sequential ascending peaks at the Pragian (Early Devonian), Givetian (Mid-Devonian), and Visean (Mississippian), with a single significant depletion in the Eifelian (Mid-Devonian). The comparative analysis of the lycophyte and euphyllophyte diversity patterns suggests that the substantial embryophytic diversity changes are driven by diversification within lineages rather than by the rhythm of the appearance of key morphological traits. This evidence (1) advocates for an intrinsic ecological control on the apparent changes in taxic richness and (2) implies that the observed plant diversity dynamics respond to the overlapping of the various expansion phases of the plant lineages involved.

Terrestrialization probably began more than one billion years ago and irreversibly altered biogeochemical processes at planetary scale. In this paper, we focus on the terrestrialization process of the Streptophyta, the division that includes charophytes and land plants (embryophytes) and whose members are today ecologically dominant in all terrestrial environments. The timing and the phylogenetic context of the early evolution of land plants are reviewed. The available information on the relationships within embryophytes and related organisms is compiled in two informal consensus trees based either on morphological/anatomical or on molecular data. We also consider the algal/embryophyte transition through the analysis of the evidence provided by microfossils (cryptospores and spores). The ongoing debate about the definition of the term cryptospores, but more importantly about the biological affinities of these microfossils that are possibly derived from early land plants, is discussed. All important clades of embryophytes, with a focus on their Palaeozoic representatives, are described; the significance of several embryophyte key characters is evaluated. The terrestrialization of land plants evolved in different steps. The new term “proembryophytic phase” is introduced to define the very long period of time during which the green algae ancestor of land plants acquired all the evolutionary characters that ultimately allowed their terrestrialization since the late Precambrian. An “eoembryophytic phase”, spanning the Middle-Upper Ordovician, is defined based on the occurrence of the earliest evidence of liverwort-like plants. The inception of the trilete spores in Late Ordovician times is then taken to define the start of the eo/eutracheophytic phase, which lasts until the first occurrence of vascular plant macrofossils in the Silurian.

Photorespiration is a process that is crucial for the survival of oxygenic phototrophs in environments that favour the oxygenation reaction of Rubisco. While photorespiration is conserved among cyanobacteria, algae, and embryophytes, it evolved to different levels of complexity in these phyla. The highest complexity is found in embryophytes, where the pathway involves four cellular compartments and respective transport processes. The complexity of photorespiration in embryophytes raises the question whether a simpler system, such as cyanobacteria, may serve as a model to facilitate our understanding of the common key aspects of photorespiration. In this study, we conducted a meta-analysis of publicly available metabolite profiles from the embryophyte Arabidopsis thaliana and the cyanobacterium Synechocystis sp. PCC 6803 grown under conditions that either activate or suppress photorespiration. The comparative meta-analysis evaluated the similarity of metabolite profiles, the variability of metabolite pools, and the patterns of metabolite ratios. Our results show that the metabolic signature of photorespiration is in part conserved between the compared model organisms under conditions that favour the oxygenation reaction. Therefore, our findings support the claim that cyanobacteria can serve as prokaryotic models of photorespiration in embryophytes.

We (1) attempted to establish an evolutionary timescale for land plant evolution utilizing available genome-scale data and a new set of calibrations constraining the age of clades based on critical analysis of paleontologic, phylogenetic (2), and geologic evidence. We explored many factors, such as the inclusion or exclusion of a calibration on the crown embryophyte node and concluded that the living clade of land plants emerged in a middle Cambrian–Early Ordovician interval. Hedges et al. (3) argue that the results of our study are not robust to dating strategies since removal of maximum constraints (maxima) results in significantly older clade age estimates. They … [↵][1]1To whom correspondence may be addressed. Email: h.schneider{at}nhm.ac.uk or phil.donoghue{at}bristol.ac.uk. [1]: #xref-corresp-1-1

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. 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.

Bioinformatics analyses of caleosin/peroxygenases (CLO/PXG) demonstrated that these genes are present in the vast majority of Viridiplantae taxa for which sequence data are available. Functionally active CLO/PXG proteins with roles in abiotic stress tolerance and lipid droplet storage are present in some Trebouxiophycean and Chlorophycean green algae but are absent from the small number of sequenced Prasinophyceaen genomes. CLO/PXG-like genes are expressed during dehydration stress in Charophyte algae, a sister clade of the land plants (Embryophyta). CLO/PXG-like sequences are also present in all of the >300 sequenced Embryophyte genomes, where some species contain as many as 10–12 genes that have arisen via selective gene duplication. Angiosperm genomes harbour at least one copy each of two distinct CLO/PX isoforms, termed H (high) and L (low), where H-forms contain an additional C-terminal motif of about 30–50 residues that is absent from L-forms. In contrast, species in other Viridiplantae taxa, including green algae, non-vascular plants, ferns and gymnosperms, contain only one (or occasionally both) of these isoforms per genome. Transcriptome and biochemical data show that CLO/PXG-like genes have complex patterns of developmental and tissue-specific expression. CLO/PXG proteins can associate with cytosolic lipid droplets and/or bilayer membranes. Many of the analysed isoforms also have peroxygenase activity and are involved in oxylipin metabolism. The distribution of CLO/PXG-like genes is consistent with an origin >1 billion years ago in at least two of the earliest diverging groups of the Viridiplantae, namely the Chlorophyta and the Streptophyta, after the Viridiplantae had already diverged from other Archaeplastidal groups such as the Rhodophyta and Glaucophyta. While algal CLO/PXGs have roles in lipid packaging and stress responses, the Embryophyte proteins have a much wider spectrum of roles and may have been instrumental in the colonisation of terrestrial habitats and the subsequent diversification as the major land flora.

The arbuscular mycorrhizal (AM) symbiosis is a nearly ubiquitous association formed by most land plants. Numerous insights into the molecular mechanisms governing this symbiosis have been obtained in recent years leading to the identification of a core set of plant genes essential for successful formation of the AM symbiosis by angiosperm hosts. Recent phylogenetic analyses indicate that while the origin of some of these symbiotic genes predated the first land plants, the rest appeared through processes including de novo evolution and gene duplication that occurred specifically in the land plants. Purifying selection on this core gene set has been maintained over millions of years of plant evolution to conserve the AM symbiosis. However, several independent losses of this association have been recorded in numerous embryophyte lineages. In these lineages, potential compensatory mechanisms have been identified that could have helped these plants overcome the adversities imposed by the loss of the AM symbiosis. This review will focus on the processes governing the conservation of the AM symbiosis in the land plant lineage.

Motivation Genome plotting is performed using a wide range of visualizations tools each with emphasis on a different informative dimension of the genome. These tools can provide a deeper insight into the genomic structure of the organism. Results Here, we announce a new visualization tool that is specifically designed for chloroplast genomes. It allows the users to depict the genetic architecture of up to ten chloroplast genomes in the vicinity of the sites connecting the inverted repeats to the short and long single copy regions. The software and its dependent libraries are fully coded in R and the reflected plot is scaled up to realistic size of nucleotide base pairs in the vicinity of the junction sites. We introduce a website for easier use of the program and R source code of the software to be used in case of preferences to be changed and integrated into personal pipelines. The input of the program is an annotation GenBank (.gb) file, the accession or GI number of the sequence or a DOGMA output file. The software was tested using over a 100 embryophyte chloroplast genomes and in all cases a reliable output was obtained. Availability and implementation Source codes and the online suit available at https://irscope.shinyapps.io/irapp/ or https://github.com/Limpfrog/irscope.

Compression fossils from the Silurian and Devonian of southern Britain, composed of cuticles and tubes, were described by W. H. Lang as the genus Nematothallus and placed, with Prototaxites, in Nematophytales, related neither to algae nor tracheophytes. Dispersed cuticles of Nematothallus and perforated forms assigned to Cosmochlaina were frequently recovered in macerates, their affinities being unresolved. New collections from a Lochkovian locality in the Welsh Borderland permitted the reconstruction of the stratified thalli of these nematophytes; they comprise a superficial cortex (which produced the cuticles) overlying a palisade zone composed of septate, parallel tubes, presumed to be hyphae, and a basal zone comprising wefts of randomly interwoven hyphae. Excellent three dimensional preservation allows the erection of a new species of Nematothallus, N. williamii. A similar anatomy is seen in a new group of fossils with either circular incisions in the cortex or complete separation of thickened cortical cells, presumably comprising a developmental sequence. By their stratified organization the nematophytes differ from extant and extinct algae and bryophytes and the enigmatic Spongiophyton. A complex anatomy and septate tubes suggest affinity with lichenized fungi. Limited data support a fungal rather than embryophyte chemistry, but a photobiont is missing. Nematophytes, globally widespread in cryptogamic covers from mid-Ordovician times, added to the biodiversity in early terrestrial ecosystems and enhanced chemical weathering.

Six monophyletic groups of charophycean green algae are recognized within the Streptophyta. Although incongruent with earlier studies based on genes from three cellular compartments, chloroplast and nuclear phylogenomic analyses have resolved identical relationships among these groups, placing the Zygnematales or the Zygnematales + Coleochaetales as sister to land plants. The present investigation aimed at determining whether this consensus view is supported by the mitochondrial genome and at gaining insight into mitochondrial DNA (mtDNA) evolution within and across streptophyte algal lineages and during the transition toward the first land plants. We present here the newly sequenced mtDNAs of representatives of the Klebsormidiales (Entransia fimbriata and Klebsormidium spec.) and Zygnematales (Closterium baillyanum and Roya obtusa) and compare them with their homologs in other charophycean lineages as well as in selected embryophyte and chlorophyte lineages. Our results indicate that important changes occurred at the levels of genome size, gene order, and intron content within the Zygnematales. Although the representatives of the Klebsormidiales display more similarity in genome size and intron content, gene order seems more fluid and gene losses more frequent than in other charophycean lineages. In contrast, the two members of the Charales display an extremely conservative pattern of mtDNA evolution. Collectively, our analyses of gene order and gene content and the phylogenies we inferred from 40 mtDNA-encoded proteins failed to resolve the relationships among the Zygnematales, Coleochaetales, and Charales; however, they are consistent with previous phylogenomic studies in favoring that the morphologically complex Charales are not sister to land plants.

in Undetermined Sporangia, with their in situ cryptospores, spore masses, and a possible axis were identified in marginal marine Ludlovian deposits from Gotland. Sweden by means of light- and scanning electron microscopy. The sporangia identified are elongate, with fragments of the sporangia covering preserved, and contain well-preserved in situ embryophyte cryptospores identified as Laevolancis divellomedia (monad) and Dyadospora murusdensa (dyad). A single detached axis was identified possessing superficial longitudinal striations resembling epidermal cell patterning. The axis ends in a cup-shaped structure that possibly represents the basal part of a sporangium. masses of cryptospore monads shrouded in amorphous organic material, most probably representing coprolites of terrestrial arthropods, were found in the same beds. The spores in these masses are degraded in sharp contrast to the excellent preservation of the dispersed and sporangia-hosted spores. The monad, L divellomedia, a common local constituent of the dispersed microflora, was identified in the coprolites. This is the first report of undisputed Silurian land plant remains in Baltica and the study reveals one of the earliest examples of arthropod-plant interactions. (Less)

As part of a study to explore diversity and disparity in Early Devonian terrestrial vegetation, several hundreds of sporangia with in situ spores have been isolated from a Lochkovian locality in Shropshire. These include a small number (seven) of sporangia showing dehiscence into four valves and containing permanent sculptured dyads, belonging to the Cymbohilates horridus complex and C. cymosus, which are recorded in coeval dispersed spore assemblages. A further, previously described, mesofossil comprises an incomplete sporangium containing C. horridus that terminates a naked isotomously branching stem with stomata. The valvate sporangia are placed in a new genus, Partitatheca, containing four species, P. splendida (type), P. horrida, P. densa and P. cymosa, their names reflecting the names of the dispersed spore species and varieties. Complex ultrastructure in the walls of the dyads is similar to that in earlier dyads in the Dyadospora complex where it provides evidence for a hepatic affinity of the earliest embryophytes, but the new taxa present a combination of bryophyte and tracheophyte characters and are considered to represent a new embryophyte lineage. General discussion includes the development of dyads, more particularly their relevance to understanding the diversity in meiotic processes, and the disappearance of dyads from the dispersed spore record prior to the Middle Devonian.

Assimilatory nitrate reductase (NR; EC 1.7.1.1-3) catalyzes the reduction of nitrate to nitrite. This enzyme has a conserved structure common to fungi, algae and plants. However, some differences in the amino acid sequence between plant and algal NR suggest that the activity regulation mechanisms have changed during plant evolution. Since only NRs from angiosperms have been studied, the search and analysis of NR genes and proteins from the moss Physcomitrella patens, a basal land plant, was performed to widen the knowledge of land plant NR structure. A family of three nr genes, named ppnia1;1, ppnia1;2 and ppnia2, was localized in the P. patens genome. The predicted proteins are canonical NRs with the conserved domains Molybdene-Cytochorme b –Cytochrome b reductase and possess 20 amino acid residues important for the enzymatic function conserved in plant and algal NRs. Interestingly, moss NRs lack a consensus sequence, common to angiosperm NRs, that is a target for posttranslational regulation. A phylogenetic tree with embryophyte and green algae NR sequences was constructed and P. patens NRs localized at the base of embryophyte NR evolution. The data presented here suggest that bryophytes and vascular plants have different systems to regulate NR activity.