SignificanceEstablishing the timescale of early land plant evolution is essential to testing hypotheses on the coevolution of land plants and Earth’s System. Here, we establish a timescale for early land plant evolution that integrates over competing hypotheses on bryophyte−tracheophyte relationships. We estimate land plants to have emerged in a middle Cambrian–Early Ordovocian interval, and vascular plants to have emerged in the Late Ordovician−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 a much earlier, middle Cambrian–Early Ordovician, origin.

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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50I.50II.52III.53IV.66V.71VI.72VII.7475References75 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.

A cored borehole through the Early Devonian Rhynie cherts at Rhynie, Aberdeenshire, NE Scotland, has revealed 53 chert beds in 35.41 m of core. The cherts originated as sinters deposited by hot-spring activity. Chert comprises 4.20 m of the cored succession, with the thickest bed, representing a single silicification event, being 0.31 m thick and the thickest composite chert (comprising six beds) 0.76 m thick. Average chert bed thickness is 80 mm. Forty-five plant-bearing chert beds are interbedded with sandstones, mudstones and shales. The sediments were deposited on an alluvial plain with local lakes, the area being periodically affected by hot-spring activity. Plants initially colonized both subaerial sand and sinter surfaces. Rhynia gwynne-vaughanii and Horneophyton lignieri commonly form the basal parts of the profiles with subsequent colonization by other genera. Rhynia is commonly found in life position above originally sandy substrates, and Horneophyton above sinter surfaces. The composition of the Rhynie vegetation is compared with coeval assemblages and, on the basis of current knowledge, it is concluded that there is no unequivocal evidence that the plants were adapted to life in the stressed environments in the immediate vicinity of hot springs.

Roots, as organs distinguishable developmentally and anatomically from shoots (other than by occurrence of stomata and sporangia on above-ground organs), evolved in the sporophytes of at least two distinct lineages of early vascular plants during their initial major radiation on land in Early Devonian times (c. 410-395 million years ago). This was some 15 million years after the appearance of tracheophytes and c. 50 million years after the earliest embryophytes of presumed bryophyte affinity. Both groups are known initially only from spores, but from comparative anatomy of extant bryophytes and later Lower Devonian fossils it is assumed that, during these times, below-ground structures (if any) other than true roots fulfilled the functions of anchorage and of water and nutrient acquisition, despite lacking an endodermis (as do the roots of extant Lycopodium spp.). By 375 million years ago root-like structures penetrated almost a metre into the substratum, greatly increasing the volume of mineral matter subject to weathering by the higher than atmospheric CO(2) levels generated by plant and microbial respiration in material with restricted diffusive contact with the atmosphere. Chemical weathering consumes CO(2) in converting silicates into bicarbonate and Si(OH)(4). The CO(2) consumed in weathering ultimately came from atmospheric CO(2) via photosynthesis and respiration; this use of CO(2) probably accounts for most of the postulated 10-fold decrease in atmospheric CO(2) from 400-350 million years ago, with significant effects on shoot evolution. Subsequent evolution of roots has yielded much-branched axes down to 40 microm diameter, a lower limit set by long-distance transport constraints. Finer structures involved in the uptake of nutrients of low diffusivity in soil evolved at least 400 million years ago as arbuscular mycorrhizas or as evaginations of "roots" ("root hairs").

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A new assemblage of arthropod cuticles from Upper Silurian rocks in Shropshire, England, includes at least two centipedes and a trigonotarbid arachnid. This unequivocal terrestrial fauna from the Silurian constitutes the earliest direct record of land animals. The presence of predatory arthropods suggests that complex terrestrial ecosystems were in place by the late Silurian (414 x 10(6) years before present) and that the animal invasion of the land occurred earlier than was previously thought.