Outline of the classification of flowering plants (Magnoliophyta)

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.

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The timescale of early land plant evolution

The pattern and ontogeny of leaf venation appear to guide or limit many aspects of leaf cell differentiation and function. Photosynthetic, supportive, stomatal, and other specialized cell types differentiate in positions showing a spatial relationship to the vascular system. These spatial relationships are of obvious importance to leaf function, which relies on venation for the servicing of cells engaged in photosynthesis, gas exchange, and other leaf processes. Although the need for coordinated organization of cell types around the vascular system is clear, the means by which this is achieved during development is not well understood. In the few systems in which it has been possible to follow the ontogeny of the venation along with the differentiation and function of surrounding cell types (e.g., in C4 grasses), observations suggest that the developing vascular system may have a role in providing positional landmarks that guide the differentiation of other cell types. Another possible explanation is that an underlying pattern guides the differentiation of both venation and surrounding cells. Whether the process of vascularization creates or reveals a pattern, studies to date are largely descriptive, and little is understood of the underlying mechanisms. These mechanisms must be highly regulated, as evidenced by the successful use of species-specific leaf vascular pattern as a taxonomic characteristic (e.g., Klucking, 1992) and by the predictable effect of certain mutations. In this review, we summarize the vascular patterns and their ontogenies in dicots and monocots, referring extensively but not exclusively to Arabidopsis and maize as examples. We also discuss a variety of models that seek to explain vascular pattern formation, and we provide a summary of molecular and genetic investigations of the process.

Leaf Vascular Pattern Formation.

• Plants have utterly transformed the planet, but testing hypotheses of causality requires a reliable time-scale for plant evolution. While clock methods have been extensively developed, less attention has been paid to the correct interpretation and appropriate implementation of fossil data. • We constructed 17 calibrations, consisting of minimum constraints and soft maximum constraints, for divergences between model representatives of the major land plant lineages. Using a data set of seven plastid genes, we performed a cross-validation analysis to determine the consistency of the calibrations. Six molecular clock analyses were then conducted, one with the original calibrations, and others exploring the impact on divergence estimates of changing maxima at basal nodes, and prior probability densities within calibrations. • Cross-validation highlighted Tracheophyta and Euphyllophyta calibrations as inconsistent, either because their soft maxima were overly conservative or because of undetected rate variation. Molecular clock analyses yielded estimates ranging from 568-815 million yr before present (Ma) for crown embryophytes and from 175-240 Ma for crown angiosperms. • We reject both a post-Jurassic origin of angiosperms and a post-Cambrian origin of land plants. Our analyses also suggest that the establishment of the major embryophyte lineages occurred at a much slower tempo than suggested in most previous studies. These conclusions are entirely compatible with current palaeobotanical data, although not necessarily with their interpretation by palaeobotanists.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.2011.03794.x

Establishing a time-scale for plant evolution

• Describe the role of grafting in human history. • Distinguish between the use of seedling and clonal rootstocks. • Describe how natural grafting can affect tree performance. • Describe how the rootstock and scion heal together during grafting. • Define how specific genetic, environmental, and management factors and polarity affect graft success. • Determine what kinds of plants can be grafted. • Define graft incompatibility— its symptoms, causes, and control. • Describe important ways the rootstock (root system) influences the scion (shoot system) and vice versa. 11 Principles of Grafting and Budding

A History of Grafting

The vascular anatomy of the flower with refutation of the theory of carpel polymorphism

The Morphology of Agathis australis1

The Ovule and ‘Seed’ of Araucaria Bidwillii with Discussion of the Taxonomy of the Genus

Destruction of phloem in young bean plants after treatment with 2,4-D.

Comparative effects of spray treatments with growth–regulating substances on the nut grass, cyperus rotundus l., and anatomical modifications following treatment with butyl 2,4–dichlorophenoxyacetate

Arthur J. Eames

Papers

The vascular anatomy of the flower with refutation of the theory of carpel polymorphism

The Morphology of Agathis australis1

The Ovule and ‘Seed’ of Araucaria Bidwillii with Discussion of the Taxonomy of the Genus

Destruction of phloem in young bean plants after treatment with 2,4-D.

Comparative effects of spray treatments with growth–regulating substances on the nut grass, cyperus rotundus l., and anatomical modifications following treatment with butyl 2,4–dichlorophenoxyacetate