The extraordinary contemporary species richness and ecological predominance of flowering plants (angiosperms) are even more remarkable when considering the relatively recent onset of their evolutionary diversification. We examine the evolutionary diversification of angiosperms and the observed differential distribution of species in angiosperm clades by estimating the rate of diversification for angiosperms as a whole and for a large set of angiosperm clades. We also identify angiosperm clades with a standing diversity that is either much higher or lower than expected, given the estimated background diversification rate. Recognition of angiosperm clades, the phylogenetic relationships among them, and their taxonomic composition are based on an empirical compilation of primary phylogenetic studies. By making an integrative and critical use of the paleobotanical record, we obtain reasonably secure approximations for the age of a large set of angiosperm clades. Diversification was modeled as a stochastic, time-homogeneous birth-and-death process that depends on the diversification rate (r) and the relative extinction rate (epsilon). A statistical analysis of the birth and death process was then used to obtain 95% confidence intervals for the expected number of species through time in a clade that diversifies at a rate equal to that of angiosperms as a whole. Confidence intervals were obtained for stem group and for crown group ages in the absence of extinction (e = 0.0) and under a high relative extinction rate (epsilon = 0.9). The standing diversity of angiosperm clades was then compared to expected species diversity according to the background rate of diversification, and, depending on their placement with respect to the calculated confidence intervals, exceedingly species-rich or exceedingly species-poor clades were identified. The rate of diversification for angiosperms as a whole ranges from 0.077 (epsilon = 0.9) to 0.089 (epsilon = 0.0) net speciation events per million years. Ten clades fall above the confidence intervals of expected species diversity, and 13 clades were found to be unexpectedly species poor. The phylogenetic distribution of clades with an exceedingly high number of species suggests that traits that confer high rates of diversification evolved independently in different instances and do not characterize the angiosperms as a whole.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.0014-3820.2001.tb00826.x

Absolute diversification rates in angiosperm clades

Tertiary macrofossils of the flowering plant family Leguminosae (legumes) were used as time constraints to estimate ages of the earliest branching clades identified in separate plastid matK and rbcL gene phylogenies. Penalized likelihood rate smoothing was performed on sets of Bayesian likelihood trees generated with the AIC-selected GTR+ Gamma +I substitution model. Unequivocal legume fossils dating from the Recent continuously back to about 56 million years ago were used to fix the family stem clade at 60 million years (Ma), and at 1-Ma intervals back to 70 Ma. Specific fossils that showed distinctive combinations of apomorphic traits were used to constrain the minimum age of 12 specific internal nodes. These constraints were placed on stem rather than respective crown clades in order to bias for younger age estimates. Regardless, the mean age of the legume crown clade differs by only 1.0 to 2.5 Ma from the fixed age of the legume stem clade. Additionally, the oldest caesalpinioid, mimosoid, and papilionoid crown clades show approximately the same age range of 39 to 59 Ma. These findings all point to a rapid family-wide diversification, and predict few if any legume fossils prior to the Cenozoic. The range of the matK substitution rate, 2.1-24.6 x 10(-10) substitutions per site per year, is higher than that of rbcL, 1.6- 8.6 x 10(-10), and is accompanied by more uniform rate variation among codon positions. The matK and rbcL substitution rates are highly correlated across the legume family. For example, both loci have the slowest substitution rates among the mimosoids and the fastest rates among the millettioid legumes. This explains why groups such as the millettioids are amenable to species-level phylogenetic analysis with these loci, whereas other legume groups are not.

/pdf/evolutionary-rates-analysis-of-leguminosae-implicates-a-bf39lfvyg4.pdf

Evolutionary rates analysis of leguminosae implicates a rapid diversification of lineages during the tertiary

Fossil in situ spores and pollen grains: an annotated catalogue

Phylogeography posits that the sequence of speciation events within a clade should parallel the geographic migration and isolation of members of the clade through time. The primary historical features that govern migration and allopatry in land plants are changes in physical geography (e.g., oceans, mountains, and deserts) and in climate (e.g., moisture, temperature, and day length), features that are often interrelated. If we assume that living genera retain physiological stability through time, much as they retain the morphological features that allow their identification, then these environmental features of the geologic past may be used to test phylogeographic hypotheses of living genera based on phylogenetic analysis. The history of the climatic and geographic features of the Tertiary of the Northern Hemisphere agrees with many phylogenetically based phylogeographic hypotheses of living angiosperm genera but indicates that some hypotheses require reanalysis. While the parallel comparison of phylogene...

https://www.jstor.org/stable/10.1086/323880

The Use of Geological and Paleontological Evidence in Evaluating Plant Phylogeographic Hypotheses in the Northern Hemisphere Tertiary

The design and function of leaf venation are important to plant performance, with key implications for the distribution and productivity of ecosystems, and applications in paleobiology, agriculture and technology. We synthesize classical concepts and the recent literature on a wide range of aspects of leaf venation. We describe 10 major structural features that contribute to multiple key functions, and scale up to leaf and plant performance. We describe the development and plasticity of leaf venation and its adaptation across environments globally, and a new global data compilation indicating trends relating vein length per unit area to climate, growth form and habitat worldwide. We synthesize the evolution of vein traits in the major plant lineages throughout paleohistory, highlighting the multiple origins of individual traits. We summarize the strikingly diverse current applications of leaf vein research in multiple fields of science and industry. A unified core understanding will enable an increasing range of plant biologists to incorporate leaf venation into their research.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.12253

Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future

The recognition of numerous similarities between Isoetes L. and extinct arborescent lycopsids historically led to one of the best Known phylogenetic lineages in vascular plant evolution. The lycopsid reduction series proposed the derivation of the extant guillwort Isoetes from the Carboniferous lepidodendrids through a progressive reduction in both stems and stigmarian rooting systems. New information gained in the past decade demonstrates that the history of this group is more complex than the linear reduction sequence suggests (...)

Evolution of Isoetalean Lycopsids

The evolution of the isoetalean lycopsids spans much of the history of vascular plants, from Late, (or possibly Middle), Devonian to the current day genus Isoetes. The best known fossil members of this group are the arborescent lepidodendrids that dominated the Late Carboniferous coal swamps. Simpler unbranched isoetaleans with elongate stems also predated, coexisted with, and postdated the coal swamp trees, extending well into the Mesozoic. Whereas certain synapomorphies such as stigmarian rootlets, bipolar growth and secondary tissues unite the clade, other features characterize smaller subgroups of differing age, growth form and possibly, evolutionary lineage. Although some of these features are well known for plants of given time periods, particularly the Carboniferous, trends in character evolution have never been adequately documented for the group as a whole. A better understanding of such trends throughout the isoetalean fossil record could be valuable in distinguishing evolutionary lineages from convergence. It is interesting that several morphological characters of modern Isoetes are present as early as the Triassic: monolete microspores, sunken sporangia and elaborate ligules with glossopodia occur within elongate-stemmed Triassic forms. The dominant plant habit of modern Isoetes, a reduced cormose form that lacks appreciable stem elongation, originated at least by the Jurassic and typifies late Mesozoic and Cenozoic isoetaleans.

Isoetalean Lycopsid Evolution: from the Devonian to the Present

Several hundred permineralized fragments of a single type of plant occur in upper Pennsylvanian coal balls from the Appalachian Basin, and these provide evidence for the description and complete reconstruction of a new heterosporous lycophyte. Chaloneria cormosa gen. et sp. nov. is unbranched, ca. 2 m tall, and has a rounded rooting base. Secondary xylem is abundant at the base, diminishes distally, and is absent from the apical region. The stem has ligulate microphylls that are vegetative at proximal levels and grade upward into a fertile apex. Leaf cushions are not produced. In the fertile zone there are alternating regions of megasporophylls and microsporophylls. A second species, Chaloneria periodica sp. nov., has alternating vegetative and fertile zones and is based on Middle Pennsylvanian specimens that were previously referred to Polysporia mirabilis. Spores of both species are assignable to Valvisisporites and Endosporites. Plants with these sporomorphs comprise several genera rather than the sing...

Chaloneria Gen. Nov.; Heterosporous Lycophytes from the Pennsylvanian of North America

Many of the oldest definitive members of the Rosaceae are present in the Eocene upland floras of the Okanogan Highlands of northeastern Washington State and British Columbia, Canada. Over a dozen rosaceous taxa representing extant and extinct genera of all four traditionally recognized subfamilies are known from flowers, fruits, wood, pollen, and especially leaves. The complexity seen in Eocene Rosaceae suggests that hybridization and polyploidy may have played a pivotal role in the early evolution of the family. Increased species diversity and the first appearance of additional modern taxa occur during the Late Paleogene in North America and Europe. The Rosaceae become increasingly important components of fossil floras during the Neogene, with taxa adapted to many habitats.

A brief review of the fossil history of the family Rosaceae with a focus on the Eocene Okanogan Highlands of eastern Washington State, USA, and British Columbia, Canada

The evolution of plant ecosystems during the Cenophytic was complex and influenced by both abiotic and biotic factors. Among abiotic forces were tectonics, the distribution of continents and seas, climate, and fires; of biotic factors were herbivores, pests, and intra- and interspecific competition. The genus Quercus L. (Quercoideae, Fagaceae) evolved in this context to become an established member of the plant communities of the Northern Hemisphere, commencing in the Paleogene and spreading to a diverse range of environments in the later Cenozoic. Its palaeontological record, dominated by leaves and pollen, but also including wood, fruits and flowers, is widespread in Eurasia and North America. Consequently, a great number of species have been described, from the 19th century to the present day. Although Quercus is currently an ecologically and economically important component of the forests in many places of the Northern Hemisphere and Southeastern Asia, no comprehensive summary of its fossil record exists. The present work, written by an international team of palaeobotanists, provides the first synthesis of the fossil history of the oaks from their appearance in the early Paleogene to the Quaternary.

Kathleen B. Pigg

Papers

Evolution of Isoetalean Lycopsids

Isoetalean Lycopsid Evolution: from the Devonian to the Present

Chaloneria Gen. Nov.; Heterosporous Lycophytes from the Pennsylvanian of North America

A brief review of the fossil history of the family Rosaceae with a focus on the Eocene Okanogan Highlands of eastern Washington State, USA, and British Columbia, Canada

The Fossil History of Quercus