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What is Palaeobotany?
General Palaeobotany
Whole Plant Reconstructions
Overviews of Plant Fossil Lagerstätten and Their Palaeoenvironments
Tertiary Palaeobotany
Cretaceous Palaeobotany
Jurassic Palaeobotany
The Rhaetian@
Triassic Palaeobotany@
Early Triassic Floras@
Permian Palaeobotany
Carboniferous Palaeobotany
Silurian and Devonian Palaeobotany

! The Molecular Clock and/or/versus the Fossil Record
! Living Fossils@
! Focussed on the Fossil Record@
Chemotaxonomy and Chemometric Palaeobotany@
! Web Sites about Evolution@
! Teaching Documents about Evolution@
Teaching Documents about Palaeobotany@
Fossil Plant and Paleovegetation Reconstructions@
Progress in Palaeobotany and Palynology@
Classical Monographs and Textbooks in Palaeobotany@

Plant Evolution

Stephen T. Abedon, The Bacteriophage Ecology Group, Mansfield, Ohio State University, Columbus: Evolution of Plants. Brief lecture notes.
The link is to a version archived by the Internet Archive´s Wayback Machine.

! K.L. Adams and J.F. Wendel (2005): Polyploidy and genome evolution in plants. In PDF, Current opinion in plant biology.
Some references have been highlighted and annotated for recommended reading.

J.F. Allen and W.F.J. Vermaas (2010): Evolution of Photosynthesis. PDF file, In: Encyclopedia of Life Sciences (ELS), John Wiley & Sons.

! J. Anderson et al. (2007): Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology. In PDF, Strelitzia, 20, 279 p. See also here (abstract).

! J.M. Anderson et al. (1999): Patterns of Gondwana plant colonisation and diversification. Abstract, Journal of African Earth Sciences, 28: 145-l67. See also here (in PDF).

A. Antonelli et al. (2015): An engine for global plant diversity: highest evolutionary turnover and emigration in the American tropics. In PDF, American tropics. Front. Genet., 6. doi: 10.3389/fgene.2015.00130
See also here.

Hank Art et al., Williams College, Biology Dept., Williamstown MA: Field botany. Go to: Evolutionary Botany. Powerpoint download and links to aricles. See especially:
Early Land Plants.
Fossil Angiosperms.
Introduction to the Angiosperms.
Powerpoint presentations.

! J. Asar et al. (2022): Early diversifications of angiosperms and their insect pollinators: were they unlinked? Free access. Trends in Plant Science, 27: 858-869. See also here.
Note figure 1: Emergence of crown angiosperms and insect pollinators.
Figure 2. Phylogeny of seed plants, depicting pollination modes of both extinct and extant lineages.

L. Augusto et al. (2014): The enigma of the rise of angiosperms: can we untie the knot? In PDF, Ecology Letters.

F.J. Ayala, Walter M. Fitch, and Michael T. Clegg (eds.; 2000): Variation and Evolution in Plants and Microorganisms: Toward a New Synthesis 50 Years after Stebbins. Online book, National Academy of Sciences (2000).
This expired link is now available through the Internet Archive´s Wayback Machine.
See also here

S.-N. Bai (2017): Reconsideration of Plant Morphological Traits: From a Structure-Based Perspective to a Function-Based Evolutionary Perspective. Front. Plant Sci., 8: 345.

J. Barba-Montoya et al. (2018): Constraining uncertainty in the timescale of angiosperm evolution and the veracity of a Cretaceous Terrestrial Revolution. In PDF, New Phytologist, 218: 819-834.
See also here.
Note fig. 6: The time tree of tracheophytes encompassing uncertainty of calibration strategies.
"... We reject a post-Jurassic origin of angiosperms, supporting the notion of a cryptic early history of angiosperms ..."

E. Barley and K. Fitzpatrick, lecture presentation for Campbell Biology, ninth edition: Plant Diversity II: The Evolution of Seed Plants. Powepoint presentation.
Still available through the Internet Archive´s Wayback Machine.

J. Barros et al. (2015): The cell biology of lignification in higher plants. Free access, Annals of Botany, 115: 1053–1074.

R.M. Bateman (2020): Hunting the Snark: the flawed search for mythical Jurassic angiosperms. In PDF, Journal of Experimental Botany, 71: 22–35. See also here.

! R.M. Bateman et al. (1998): Early evolution of land plants: phylogeny, physiology, and ecology of the primary terrestrial radiation. PDF file, Annu. Rev. Ecol. Syst., 29: 263-292. Website saved by the Internet Archive´s Wayback Machine.

! R.M. Bateman and W.A. DiMichele (1994): Heterospory: the most iterative key innovation in the evolutionary history of the plant kingdom. In PDF, Biological Reviews.

BBC News, Friday, 3 May, 2002: "Oldest flower" found in China.

J.M. Beaulieu et al. (2015): Heterogeneous rates of molecular evolution and diversification could explain the Triassic age estimate for angiosperms. Abstract.

! B. Becker and B. Marin (2009): Streptophyte algae and the origin of embryophytes. In PDF, Annals of Botany, 103: 999–1004. See also here.
Note fig. 2: Diversification of green plants (Viridiplantae) and colonization of terrestrial habitats by streptophyte algae.

D.J. Beerling (2013): Atmospheric carbon dioxide: a driver of photosynthetic eukaryote evolution for over a billion years? In PDF, Philos. Trans. R. Soc. Lond. B, Biol. Sci., 367: 477-482.

! D. Beerling (2010): The Emerald Planet. How Plants Changed Earth´s History. In PDF.

D.J. Beerling and R.A. Berner (2005): Feedbacks and the coevolution of plants and atmospheric CO2. In PDF, PNAS, 102.

! C.D. Bell et al. (2010): The age and diversification of the angiosperms re-revisited. Free access, American Journal of Botany, 97: 1296-1303.

! M.J. Benton and F. Wu (2022): Triassic revolution. Free access, Frontiers in Earth Science, 10. See also here.
Note figure 9: Novel physiological and functional characteristics, new tetrapod, insect and plant groups in the Triassic on land.
"... On land, ongoing competition between synapsids and archosauromorphs through the Triassic was marked by a posture shift from sprawling to erect, and a shift in physiology to warm-bloodedness, with insulating skin coverings of hair and feathers. Dinosaurs, for example, originated in the Early or Middle Triassic, but did not diversify until after the CPE [Carnian Pluvial Episode]. ..."

! M.J. Benton et al. (2022): The Angiosperm Terrestrial Revolution and the origins of modern biodiversity. Free access, New Phytologist, 233: 2017–2035.
Note fig. 1: Evolution of hyperdiverse terrestrial life.
Fig. 3: Key stages in Earth history and angiosperm evolution through the Angiosperm Terrestrial Revolution.
Also worth checking out:
Flowering plants: an evolution revolution. (Univ. of Bristol, November 17, 2021).
How 'Flower Power' Quite Literally Transformed Earth Millions of Years Ago (by T. Koumoundouros, January 08,2022).

! H. Beraldi-Campesi (2013): Early life on land and the first terrestrial ecosystems. In PDF, Ecological Processes, 2. See also here.
Note figure 1: Suggested chronology of geological, atmospheric, and biological events during the Hadean, Archean, and Paleoproterozoic eons.

! Museum of Paleontology (UCMP), University of California at Berkeley, Plantae, Fossil Record: Chart of First Appearances of Major Plant Groups. Each of the taxonomic plant groups in pink boxes can be clicked upon to take you to an introduction.

Botanischer Garten und Botanisches Museum Berlin:
Von Nacktpflanzen und Schuppenbäumen. Ein Streifzug durch die Entwicklungsgeschichte der Pflanzen (in German). Easy to understand introduction.

! R.A. Berner et al. (2007): Oxygen and evolution. In PDF, Science, 316. Website saved by the Internet Archive´s Wayback Machine.

Robert A. Berner, Geology and Geophysics, Yale University, New Haven, Connecticut: The Rise of Plants and Their Effect on Weathering and Atmospheric CO2 (now via wayback archive). See also here.

Michael Bernstein, Washington and New Orleans, March 21-27, 2003: (American Chemical Society, EurekAlert): Scientists find evidence for crucial root in the history of plant evolution.

! C.M. Berry (2019): Palaeobotany: The Rise of the Earth’s Early Forests. Free access, Current Biology, 29: R792-R794. See also here (in PDF).

! J.A. Berry et al. (2010): Stomata: key players in the earth system, past and present. Abstract, Current opinion in plant biology, 13: 232–239. See also here (in PDF).

! M.I. Bidartondo et al. (2011): The dawn of symbiosis between plants and fungi. In PDF, Biology Letters. See also here.

E. Biffin et al. (2013): Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation. In PDF, Proc. R. Soc. B, 279: 341-348.

A.C. Bippus et al. (2022): The Role of Paleontological Data in Bryophyte Systematics. Abstract, Journal of Experimental Botany.
"... Paucity of the bryophyte fossil record, driven primarily by phenotypic (small plant size) and ecological constraints (patchy substrate-hugging populations), and incomplete exploration, results in many morphologically isolated, taxonomically ambiguous fossil taxa. Nevertheless, instances of exquisite preservation and pioneering studies demonstrate the feasibility of including bryophyte fossils in evolutionary inference. ..."

H.J.B. Birks (2020): Angiosperms versus gymnosperms in the Cretaceous. Open access, PNAS, 117: 30879-30881.

B. Blonder et al. (2011): Venation networks and the origin of the leaf economics spectrum. In PDF, Ecology Letters, 14: 91-100. See also here.

B. Bomfleur et al. (2018): Polar Regions of the Mesozoic-Paleogene Greenhouse World as Refugia for Relict Plant Groups. Chapter 24, in PDF, in: M. Krings, C.J. Harper, N.R. Cuneo and G.W. Rothwell (eds.): Transformative Paleobotany Papers to Commemorate the Life and Legacy of Thomas N. Taylor.
Note figure 24.2: Distribution of Dicroidium through space and time.

W.J. Bond and A.C. Scott (2010): Fire and the spread of flowering plants in the Cretaceous. In PDF, New Phytologist, 188: 1137-1150.

Boston College: BC Scientist´s Fossil Discovery May Indicate Life on Land Evolved Earlier than Thought.
The link is to a version archived by the Internet Archive´s Wayback Machine.

! J.L. Bowman et al. (2022): The renaissance and enlightenment of Marchantia as a model system. Open access, The Plant Cell, koac219. See also
Note fig. 1: Phylogenetic history of Marchantia plotted against the geologic timescale.
Figure 2. Life cycle of Marchantia polymorpha.

J.L. Bowman et al. (2017): Insights into Land Plant Evolution Garnered from the Marchantia polymorpha Genome. Abstract, Cell, 171: 287-304. See also here (in PDF).

C.K. Boyce and J.-E. Lee (2017): Plant Evolution and Climate over Geological Timescales. Abstract, Annual Review of Earth and Planetary Sciences, 45.

C.K. Boyce and M.A. Zwieniecki (2019): The prospects for constraining productivity through time with the whole-plant physiology of fossils Open access, New Phytologist, 223: 40–49.

C.K. Boyce and A.B. Leslie (2012): The Paleontological Context of Angiosperm Vegetative Evolution. In PDF, International Journal of Plant Sciences, 173: 561–568.
See also here.
"... a survey of the fossil record demonstrates that most anatomical traits that are now unique to the angiosperms were more broadly distributed among extinct lineages.
[...] Of all the various vegetative morphological traits that have been traditionally linked to angiosperm success, only the evolution of very high leaf vein densities appears to be truly unique to the angiosperms. ..."

C.K. Boyce and M.A. Zwieniecki (2012): Leaf fossil record suggests limited influence of atmospheric CO2 on terrestrial productivity prior to angiosperm evolution. Free access, PNAS, 109: 10403–10408.

! C.K. Boyce (2010): The evolution of plant development in a paleontological context. In PDF Current Opinion in Plant Biology, 13: 102-107.

C. Kevin Boyce et al. (2009). Angiosperm leaf vein evolution was physiologically and environmentally transformative. PDF file, Proceedings of the Royal Society B, 276: 1771-1776. See also here (abstract).

Snapshot provided by the Internet Archive´s Wayback Machine.

! Jamie Boyer, The New York Botanical Garden: The Paleoplant Website. An educational resource for students and teachers studying Earth's history, fossils, and evolution.
! Go to: Paleobotany Short-Course. Lecture notes, with downloadable Power Point presentations:
Paleobotany Overview; Life moves to land.
Diversification of land plants.
Rise and dominance of seed plants.
Origin and diversification of flowering plants.

J. Boyer (2008): Testing the Telome Concept: A Modeling Approach for Understanding the Growth of Early Vascular Plants. In PDF, Ph.D. Dissertation, Binghamton University State University of New York.

! M. Brasier et al. (2006): A fresh look at the fossil evidence for early Archaean cellular life. In PDF, Philos. Trans. R. Soc. Lond. B, Biol Sci., 361: 887–902. See also here.

J.C. Briggs (2014): Invasions, adaptive radiations, and the generation of biodiversity. In PDF, Environmental Skeptics and Critics, 3: 8-16.

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! Purported Triassic angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! The origin and evolution of angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

T.J. Brodribb and S.A.M. McAdam (2017): Evolution of the stomatal regulation of plant water content. Open access, Plant Physiology, 175: 639–649.

! Stephen P. Broker, Yale-New Haven Teachers Institute: The Evolution of Plants. Snapshot taken by the Internet Archive´s Wayback Machine.
The evolution of plants is briefly treated primarily in terms of a consideration of the concepts of time and change, and an appreciation of the great diversity of life on earth today (without images). Recommended for Biology, 9th and 10th grade level, and Botany, 11th and 12th grade level.

! N. Brocklehurst et al. (2020): The origin of tetrapod herbivory: effects on local plant diversity. Free access, Proc. R. Soc. B 287: 20200124.
"... findings suggest that plant richness was to some extent structured by vertebrate herbivory from its earliest origins more than 300 Mya. Studies of modern ecosystems suggest that this should be the case, ..."

J.W. Brown and S.A. Smith (2017): The Past Sure Is Tense: On Interpreting Phylogenetic Divergence Time Estimates. See also here (in PDF).

M.C. Brundrett (2002): Coevolution of roots and mycorrhizas of land plants. In PDF, New phytologist, 154: 275-304.
Provided by the Internet Archive´s Wayback Machine.

Neil Buckley, Department of Biological Sciences, SUNY Plattsburgh, Plattsburgh, NY: Bio 102 General Biology II Class Notes. Powerpoint presentations. Go to:
An overview of Plant Evolution. Key Moments in the life of Kingdom Plantae.
Websites still available via Internet Archive Wayback Machine.

G.E. Budd et al. (2021): Fossil data do not support a long pre-Cretaceous history of flowering plants. Free access, bioRxiv.

G.E. Budd et al. (2021): The use of geological and paleontological evidence in evaluating plant phylogeographic hypotheses in the Northern Hemisphere Tertiary. Free access, See also here (in PDF).

R.J.A. Buggs (2021): The origin of Darwin’s “abominable mystery”. Free access, American Journal of Botany, 108: 22–36.

R.J. Burnham (2008): Hide and Go Seek: What Does Presence Mean in the Fossil Record. Abstract, Annals of the Missouri Botanical Garden, 95: 51-71. See also here (in PDF).

! Plant Sciences, University of Cambridge (developed by Nicola Peart and Ben Roberts, with feedback from Katy Jordan, Howard Griffiths and Beverley Glover): Plant Evolution Timeline. This is a cut-down version of the full tool, designed primarily to aid plant scientists with their learning of plant evolution.
Click: "Timeline".
Including first occurence (or first appearance) of species and speciation and major groups of plants. See also:
Plant Evolution Timeline - Help.

Alison Campbell et al., Biology & Earth Science, The University of Waikato, New Zealand: Evolution for Teaching. This website has been developed to provide a web based resource for use by secondary teachers, especially in the science fields of evolution and geological time. Go to: Plant and Animal Evolution.

P.D. Cantino et al. (2007): Towards a phylogenetic nomenclature of Tracheophyta. PDF file, Taxon, 56: 822-846. See also here.

! E. Capel et al. (2022): The Silurian–Devonian terrestrial revolution: Diversity patterns and sampling bias of the vascular plant macrofossil record. In PDF, Earth-Science Reviews, 231.
See also here.
Note fig. 6: Silurian–Devonian diversity patterns of tracheophytes.
"... The sampling-corrected pattern of standing diversity suggests a clear increase of plant richness in the Pragian (Early Devonian) and Givetian (Middle Devonian) ..."

E. Capel et al. (2021): A factor analysis approach to modelling the early diversification of terrestrial vegetation. In PDF, Palaeogeography,Palaeoclimatology, Palaeoecology. See also here.

Sean Carrington, Department of Biological & Chemical Sciences, University of the West Indies (UWI), Barbados: The Plant Kingdom. An introduction to the world of plants from an evolutionary perspective.
Have a look for handouts and PDF files, or
navigate from here.

! M.R. Carvalho et al. (2021): Extinction at the end-Cretaceous and the origin of modern Neotropical rainforests Science, 372: 63–68. See also here (in PDF).
"... Plant diversity declined by 45% at the Cretaceous–Paleogene boundary and did not recover for ~6 million years. ..."
Please take notice: Wie der Asteroid den Regenwald prägte., in German.

! B. Cascales-Miñana et al. (2019): An alternative model for the earliest evolution of vascular plants. Abstract, Lethaia, 52: 445–453. See also here (in PDF).

! B. Cascales-Miñana and C.J. Cleal (2013): The plant fossil record reflects just two great extinction events. Abstract. See also here (in PDF).

B. Cascales-Miñana and J.B. Diez (2012): The effect of singletons and interval length on interpreting diversity trends from the palaeobotanical record. In PDF, Palaeontologia Electronica.

David D. Cass, Department of Biological Sciences, University of Alberta: Earliest Evidence of Flowering Plants. 32 slides.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

S.R.S. Cevallos-Ferriz et al. (2022): Paleobotany to understand evolution and biodiversity in Mexico. In PDF, Botanical Sciences, 100 (Special Issue): S34-S65.
See also here.

! A.S. Chanderbali et al. (2016): Evolving Ideas on the Origin and Evolution of Flowers: New Perspectives in the Genomic Era. In PDF, Genetics, 202: 1255–1265. See also here.

(?), University of Virginia, Charlottesville:
! Evolution of Land Plants.
Evolution of Seed Plants.
Powerpoint presentations.

! J. Chave et al. (2009): Towards a worldwide wood economics spectrum. In PDF, Ecology Letters, 12: 351–366.

S.M. Chaw et al. (1997): Molecular phylogeny of extant gymnosperms and seed plant evolution: analysis of nuclear 18S rRNA sequences. In PDF.

(?), University of Virginia, Charlottesville:
Evolution of Land Plants.
Evolution of Seed Plants.
Powerpoint presentations.

G. Chomicki et al. (2017): Evolution and ecology of plant architecture: integrating insights from the fossil record, extant morphology, developmental genetics and phylogenies. Annals of Botany 120: 855–891. See also here (in PDF).
Note Fig. 9: A timeline of plant architectures and branching mechanisms through time.

! M.J.M. Christenhusz et al. (2021): Biogeography and genome size evolution of the oldest extant vascular plant genus, Equisetum (Equisetaceae). Free access, Annals of Botany, 127: 681–695.
Note figure 2: Tree of Equisetum.
Figure 3: Spatiotemporal evolution of Equisetum.
"... With a calculated age of 342 Mya, we place the origin of the ancestor of the extant members of Equisetaceae in the Early Carboniferous, with Equisetum evolving at some point after that, probably soon, but definitely before 175 Mya, making it possibly the oldest extant genus of vascular plants. ..."

! T.Y.S. Choo and I.H. Escapa (2018): Assessing the evolutionary history of the fern family Dipteridaceae (Gleicheniales) by incorporating both extant and extinct members in a combined phylogenetic study. Abstract, American Journal of Botany 105: 1–14. See also here (in PDF).

Paul F. Ciesielski, Dept. Geological Sciences, University of Florida: Evolution of Earth and Life. Go to: Transition of plants to land.
Snapshot provided by the Internet Archive´s Wayback Machine.

Citable reviews in the life sciences (Wiley). Go to:
Plant Evolution.

J.W. Clark and P.C.J. Donoghue (2018): Whole-Genome Duplication and Plant Macroevolution. In PDF, Trends in Plant Science, 23: 933-945. See also here.

! J.T. Clarke et al. (2011): Establishing a time-scale for plant evolution. PDF file, New Phytologist. See also here.
! Note figure 2: A representative tree of relationships between model representatives of the major land plant lineages whose plastid or nuclear genomes have been fully sequenced.
Figure 7: Chronogram for land plant evolution.
Figure 8: Chronograms for the six molecular clock analyses conducted.
"... 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. ..."

Regine Claßen-Bockhoff (2001): Plant Morphology: The Historic Concepts of Wilhelm Troll, Walter Zimmermann and Agnes Arber. Free PDF file, Annals of Botany, 88: 1153-1172.

! C.J. Cleal and B. Cascales-Miñana (2021, start on PDF-page 39): Evolutionary floras - revealing large-scale patterns in Palaeozoic vegetation history. Journal of Palaeosciences, 70: 31-42.

C.J. Cleal et al. (2021): Palaeobotanical experiences of plant diversity in deep time. 1: How well can we identify past plant diversity in the fossil record? Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology.

C.J. Cleal and B. Cascales-Miñana (2014): Composition and dynamics of the great Phanerozoic Evolutionary Floras. Abstract.

J.L. Cloudsley-Thompson (2005): Ecology and Behaviour of Mesozoic Reptiles, The Mesozoic Environment. In PDF. See also here,

J.C. Coates et al. (2011): Plants and the Earth system - past events and future challenges. In PDF, New Phytologist, 89: 370-373.

C. Coiffard et al. (2012): Rise to dominance of angiosperm pioneers in European Cretaceous environments . Abstract. See also here ( and there (

! M. Coiro et al. (2022): Cutting the long branches: Consilience as a path to unearth the evolutionary history of Gnetales. Open access, Front. Ecol. Evol. 10:1082639. doi: 10.3389/fevo.2022.1082639.
See also here.
Note figure 2: Phylograms showing the presence of long branches in the Gnetales. Figure 6: Top: examples of macrofossils with gnetalean affinity.

! M. Coiro et al. (2019): How deep is the conflict between molecular and fossil evidence on the age of angiosperms? Free access, New Phytologist, doi: 10.1111/nph.15708.
"... Critical scrutiny shows that supposed pre-Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms. ..."

F.L. Condamine et al. (2020): The rise of angiosperms pushed conifers to decline during global cooling. Free access, Proceedings of the National Academy of Sciences, 117: 28867–28875.
Note figure 1: An overview of hypothetical determinants of conifer diversification over time.
Figure 2: Global diversification of conifers inferred from a molecular phylogeny and the fossil record.
Figure 3: Drivers of conifer diversification dynamics.

! F.L. Condamine et al. (2015): Origin and diversification of living cycads: a cautionary tale on the impact of the branching process prior in Bayesian molecular dating. In PDF, BMC Evolutionary Biology.
Note figure 1: The node calibration procedure used for dating the cycads.
! Figure 2: Time-calibrated phylogeny of Cycadales.
! "... We also provide new insights into the history of cycad diversification because we found (i) periods of extinction along the long branches of the genera consistent with fossil data, and (ii) high diversification rates within the Miocene genus radiations. ..."

Richard Cowen, Department of Geology, University of California, Davis: Comparing Plant and Animal Evolution.
Still available via Internet Archive Wayback Machine.

C.J. Cox et al. (2014): Conflicting Phylogenies for Early Land Plants are Caused by Composition Biases among Synonymous Substitutions. Syst. Biol., 63: 272-279.

! P.R. Crane and A.B. Leslie (2013): Major Events in the Evolution of Land Plants. In PDF. The Princeton Guide to Evolution.
1. Phylogenetic framework.
2. Origin and diversification of land plants.
3. Origin and diversification of vascular plants.
4. Origin and diversification of seed plants.
5. Origin and diversification of flowering plants.
6. Innovation in the land plant body.
7. Innovation in land plant reproduction.
8. Co-evolution with animals.
9. Patterns of extinction.
See also here, and there (Google books).

P.R. Crane et al. (2010): Darwin and the Evolution of Flowers. PDF file, Phil. Trans. R. Soc. B, 365: 347-350.
The link is to a version archived by the Internet Archive´s Wayback Machine.

Peter R. Crane et al. (2004): Fossils and plant phylogeny. Free access, American Journal of Botany, 91: 1683-1699.

! W.L. Crepet and K.J. Niklas (2009): Darwin´s second "abominable mystery": Why are there so many angiosperm species? Open access, American Journal of Botany, 96: 366-381.

! W.L. Crepet (2008): The Fossil Record of Angiosperms: Requiem or Renaissance? Abstract, Annals of the Missouri Botanical Garden, 95: 3-33.
"... a reasonably good fossil record of angiosperms is emerging
[...] One of its most striking aspects is the rapid radiation of angiosperm taxa that are now unusually diverse around two particular times in geological history: the Turonian and Early Tertiary. Possible reasons for these intervals of rapid radiation among angiosperms will be discussed.

W.L. Crepet et al. (2004): Fossil evidence and phylogeny: the age of major angiosperm clades based on mesofossil and macrofossil evidence from Cretaceous deposits. Free access, American Journal of Botany, 91: 1666-1682.
! Beautifully preserved charcoalified flowers!

Cyberinfrastructure for Phylogenetic Research (CIPRES). Building the Tree of Life: A National Resource for Phyloinformatics and Computational Phylogenetics. CIPRES is a collaboration of many american museums and institutions. Go to:
! Getting to the Roots of Plant Evolution (Powerpoint presentation). See also the Exercise Handout (PDF file).
Snapshots provided by the Internet Archive´s Wayback Machine.

R. Daber (2012), Academic Universal-Lexikon:
Evolution: Pflanzen erobern das Festland (in German).
These expired links are now available through the Internet Archive´s Wayback Machine.

! T.W. Dahl and S.K.M. Arens (2020): The impacts of land plant evolution on Earth's climate and oxygenation state – An interdisciplinary review. Open access, Chemical Geology, 547.

D. J. Daniels, Glendale High School, Glendale, Arizona: Advanced Placement Biology 2000, D. J. Daniels´ Glendale High Biology Page, Land Plants, Evolution and Diversity.

C.C. Davis and S. Matthews (2019); Davis Lab, Harvard University Herbaria, Cambridge, MA:
! Evolution of Land Plants. In PDF.
Evolution of land plants in a nutshell, including a selected annotated bibliography.
Issued in Oxford Bibliographies, selected bibliographies for a variety of academic topics.

C.C. Davis and H. Schaefer (2011): Plant Evolution: Pulses of Extinction and Speciation in Gymnosperm Diversity. Open access, Current Biology.

H.J. de Boer et al. (2012): A critical transition in leaf evolution facilitated the Cretaceous angiosperm revolution. In PDF, Nature Communications, 3.

O. De Clerck et al. (2012): Diversity and Evolution of Algae: Primary Endosymbiosis. In PDF, Advances in Botanical Research, 64.
This expired link is available through the Internet Archive´s Wayback Machine.

! A.L. Decombeix et al. (2019): Plant hydraulic architecture through time: lessons and questions on the evolution of vascular systems. In PDF, IAWA Journal, 40: 387-420. See also here and there.

P.M. Delaux and S. Schornack (2021): Plant evolution driven by interactions with symbiotic and pathogenic microbes. In PDF, Science, American Association for the Advancement of Science (AAAS), 371 (6531), pp.eaba6605. ff10.1126/science.aba6605ff. ffhal-03327916ff.
See also here.
"... Delaux and Schornack review how insights from a range of plant and algal genomes reveal sustained use through evolution of ancient gene modules as well as emergence of lineage-specific specializations. Mosses, liverworts, and hornworts have layered innovation onto existing pathways to build new microbial interactions. ..."

P.M. Delaux et al. (2012): Molecular and biochemical aspects of plant terrestrialization. In PDF, Perspectives in Plant Ecology, Evolution and Systematics, 14: 49-59.

! L.E.V. Del-Bem (2018): Xyloglucan evolution and the terrestrialization of green plants. Free access, New Phytologist, 219: 1150–1153.

C.F. Delwiche et al. (2017): Land Plant Model Systems Branch Out. In PDF, Cell, 171.
"Liverworts may be the sister taxon to all other land plants, and the genome shows features that illuminate the ancestor of all land plants and give insights into how plant systems function and evolved".

! C.F. Delwiche and E.D. Cooper (2015): The Evolutionary Origin of a Terrestrial Flora. Abstract, Current Biology. Please take notice:
! From algae to land plants (and vice versa). Did some freshwater algae descend from a terrestrial ancestor? In PDF.

B. De Rybel et al. (2016): Plant vascular development: from early specification to differentiation. Abstract, Nat. Rev. Mol. Cell Biol., 2016: 30-40. see also here and there (in PDF).

Melanie DeVore, Department of Biological and Environmental Science, Georgia College and State University: Plant Origin and Evolution. PowerPoint presentation (87.4 MB!).

! J. De Vries and J.M. Archibald (2018): Plant evolution: landmarks on the path to terrestrial life. Free access, New Phytologist, 217: 1428-1434.

J. de Vries et al. (2018): Embryophyte stress signaling evolved in the algal progenitors of land plants. In PDF, PNAS, 115. See also here (abstract), and there (in German).

! D.L. Dilcher (2001): Paleobotany: some aspects of non-flowering and flowering plant evolution. In PDF, Taxon.
See also here.
For early angiospermous fossil floras see figure 1 (on PDF page 5).

David Dilcher (2000): Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. PDF file, Proc Natl Acad Sci U S A., 97: 7030-7036. See also here.

Adam Dimech, Department of Economic Development, Jobs, Transport & Resources (DEDJTR), Australia. How did the world´s plants evolve? A guide to the history of the plant kingdom.

! W.A. DiMichele et al. (2008): The so-called "Paleophytic-Mesophytic" transition in equatorial Pangea. Multiple biomes and vegetational tracking of climate change through geological time. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 268: 152-163. See also here (abstract), and there (still available via Internet Archive Wayback Machine).
! "... the evidence for a global “Paleophytic” vs. “Mesophytic” “vegetation” is simply unsubstantiated by the fossil record.
[...] The vegetational changes occurring in the late Paleozoic thus can be understood best when examined as spatial–temporal changes in biome-scale species pools responding to major global climate changes, locally and regionally manifested. ..."

! W.A. DiMichele et al. (2006): From wetlands to wet spots: Environmental tracking and the fate of Carboniferous elements in Early Permian tropical floras. PDF file. In Greb, S.F., and DiMichele, W.A., Wetlands through time: Geological Society of America Special Paper 399, p. 223–248. See also here and there (Google books).

! W.A. DiMichele and R.M. Bateman (2005): Evolution of Land Plant Diversity: Major Innovations and Lineages through Time. In PDF, In: Krupnick, G.A. & Kress, W.J. (eds): Plant Conservation. A Natural History Approach. Chicago University Press, Chicago, IL, 3–14.

W.A. DiMichele et al. (2004): Long-term stasis in ecological assemblages: evidence from the fossil record. PDF file, Annu. Rev. Ecol. Evol. Syst., 35: 285-322. Provided by the Internet Archive´s Wayback Machine.

! W.A. DiMichele (1999): EVOLUTIONARY AND PALEOECOLOGICAL IMPLICATIONS OF TERRESTRIAL FLORAL CHANGES IN THE LATE PALEOZOIC TROPICS. Abstract, 1999 GSA Annual Meeting, Denver, Colorado; The Geological Society of America (GSA).
This expired link is now available through the Internet Archive´s Wayback Machine.

Adam Dimech, Burnley College, University of Melbourne, Australia: Plant Evolution. This website is designed to serve as an introduction to the theory behind the evolution of the world's flora, with some emphasis placed on the Australian flora.

D.S. Domozych et al. (2013): The Cell Walls of Green Algae: A Journey through Evolution and Diversity. In PDF.

! P.C.J. Donoghue et al. (2021): The evolutionary emergence of land plants. In PDF, Current Biology, 31: R1281-R1298.
See also here.
"... The oldest possible fossil evidence for land plants occurs as late Cambrian cryptospores, but their irregular arrangements and occurrence in ‘packets’ of multiple spore-like bodies sur- rounded by synoecosporal walls has led to algal interpretations ..."
! Note figure 4: Timescale of streptophyte phylogeny and the origin of land plant novelties.

P. Donoghue (2019): Evolution: The Flowering of Land Plant Evolution. Abstract, Current Biology. See also:
Evolution: The flowering of land plant evolution - whence and whither? (in PDF).

! M.J. Donoghue et al. (2015): Confluence, synnovation, and depauperons in plant diversification. Open access, New Phytologist, 207: 260–274.

M.J. Donoghue and E.J. Edwards (2014): Biome shifts and niche evolution in plants. In PDF, Annu. Rev. Ecol. Evol. Syst., 45: 547-572.

M.J. Donoghue (2005): Key innovations, convergence, and success: macroevolutionary lessons from plant phylogeny. In PDF, Paleobiology, 31: 77-93.
See also here.

M.J. Donoghue (1989): Phylogenies and the analysis of evolutionary sequences, with examples from seed plants. Open access, Evolution, 43.

! M.J. Donoghue (1989): Phylogenies and the analysis of evolutionary sequences, with examples from seed plants. Open access, Evolution, 43.

J.A. Doyle and P.K. Endress (2014): Integrating Early Cretaceous Fossils into the Phylogeny of Living Angiosperms: ANITA Lines and Relatives of Chloranthaceae Int. J. Plant Sci., 175: 555–600.

! J.A. Doyle (2013): Phylogenetic analyses and morphological innovations in land plants. Free access, Annual Plant Reviews book series, Volume 45: The Evolution of Plant Form. See also here (in PDF).

! J.A. Doyle (2012): Molecular and fossil evidence on the origin of angiosperms. In PDF, Annual Review of Earth and Planetary Sciences, 40: 301-26.

James A. Doyle, Section of Evolution and Ecology, University of California, Davis: PHYLOGENY OF VASCULAR PLANTS. Abstract, Annu. Rev. Ecol. Syst. 1998. 29:567-599. See also here.

X.Y. Du et al. (2021): Simultaneous diversification of Polypodiales and angiosperms in the Mesozoic. In PDF, Cladistics, 37.
See also here.
Note fig. 2: Summary chronograms of Polypodiales.
Fig. 5. Comparison of lineage through time plots for Polypodiales and angiosperms.
"... The estimated divergence patterns of Polypodiales and angiosperms converge to a scenario in which their main lineages were established simultaneously shortly before the onset of the Cretaceous Terrestrial Revolution ..."

! D. Edwards et al. 2022a): Piecing together the eophytes–a new group of ancient plants containing cryptospores. Free access, New Phytologist, 233: 1440–1455.

! D. Edwards et al. 2022b): Earliest record of transfer cells in Lower Devonian plants. Free access, New Phytologist, 233: 1456–1465.

! D. Edwards and P. Kenrick (2015): The early evolution of land plants, from fossils to genomics: a commentary on Lang (1937) "On the plant-remains from the Downtonian of England and Wales". Open access, Phil. Trans. R. Soc. B 370.
Note figure 4: Relationships among major groups of land plants showing the hypothesized broad range of clades to which cryptophytes (extinct cryptospore-producing plants) might belong.

! A. Elgorriaga et al. (2019): Relictual Lepidopteris (Peltaspermales) from the Early Jurassic Cañadón Asfalto Formation, Patagonia, Argentina. Abstract, Int. J. Plant Sci., 180. See also here (in PDF), and there.
"... and its youngest species, Lepidopteris ottonis, has been used as a Rhaetian marker for several European, Greenlandic, and American localities ..."
"... Lepidopteris scassoi represents the youngest occurrence of the genus by more than 20 Myr. Lepidopteris and Dicroidium lineages, dominant in Southern Hemisphere Triassic ecosystems, show a similar overall pattern of origination (Late Permian), diversification (late Early-Middle Triassic), and decline (Late Triassic), with relict occurrences during the Early Jurassic. ..."

C. Elliott-Kingston et al. (2021): Creating a university evolution garden: An integrated learning approach for teaching land plant evolution. Open access, Plants People Planet, 3: 761-774.

P.K. Endress (2011): Angiosperm ovules: diversity, development, evolution. Free access, Annals of Botany, 107: 1465-1489.

M.J. Farabee, Estrella Mountain Community College, Phoenix, Arizona:
! On-Line Biology Book. Table of Contents. Introductory biology lecture notes. Go to:
! The modern view of evolution.

J.A. Fawcett and Y. Van de Peer (2010): Angiosperm polyploids and their road to evolutionary success. Trends in Evolutionary Biology.
See also here.

T.S. Feild et al. (2011): Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution. In PDF, PNAS, 108: 8363-8366.

F.A.A. Feijen et al. (2018): Evolutionary dynamics of mycorrhizal symbiosis in land plant diversification. In PDF, Scientific reports.

C. Ferrándiz et al. (2010): Carpel development. In PDF, Advances in Botanical Research, 55: 1-73.
See also here.

! K.J. Field and S. Pressel (2018): Unity in diversity: structural and functional insights into the ancient partnerships between plants and fungi. In PDF, New Phytologist. See also here

K.J. Field et al. (2015): Symbiotic options for the conquest of land. In PDF, Trends in Ecology and Evolution, 30: 477-486. See also here.

S. Fields (2021): Diversification of Angiosperms During the Cretaceous Period. In PDF, Undergraduate Student Theses, Environmental Studies Program at DigitalCommons@University of Nebraska,Lincoln. See also here.

J. Folsom, The Huntington Library, Art Collections, and Botanical Gardens, San Marino, CA: Plant Trivia Timeline. This expired link is available through the Internet Archive´s Wayback Machine. See also:
here (PDF file). The Timeline gives world history from the viewpoint of a botanist. It is the story of plant discovery and use, and addresses the roles of plants in human civilization.

David Ford, Canopy Dynamics Lab, School of Environmental and Forest Resources, University of Washington, Seattle, WA:
! Biol220 TAs. Botany lecture notes (Powerpoint presentations). See especially:
The Importance of Plants, their origins and ways of life.
Plant evolution timeline on Powerpoint slide 11, 18 and 22!

! F. Forrest (2009): Calibrating the Tree of Life: fossils, molecules and evolutionary timescales. Free access, Annals of Botany, 104: 789–794.
"... New methods have now been proposed to resolve potential sources of error associated with the calibration of phylogenetic trees, particularly those involving use of the fossil record.
[...] ! "...the fossil record remains the most reliable source of information for the calibration of phylogenetic trees, although associated assumptions and potential bias must be taken into account. ..."

! C.S.P. Foster (2016): The evolutionary history of flowering plants. In PDF, Journal & Proceedings of the Royal Society of New South Wales, 149: 65-82.

P.J. Franks et al. (2012): Megacycles of atmospheric carbon dioxide concentration correlate with fossil plant genome size. Abstract, Phil. Trans. R. Soc. B, 367: 556-564.

W.A. Friedman (2020): Darwin in the garden: Engaging the public about evolution with museum collections of living objects. Open access, Plants, People, Planet, 2: 294–301.
"... Polls continue to show distressingly high percentages of people around the world do not accept that evolution has occurred.
[...] It is time for botanical gardens and arboreta around the world to commit to leveraging their living collections of museum objects to explain and demonstrate the roles of mutation, variation, and selection in the evolutionary process. In doing so, much could be accomplished to increase scientific literacy at a societal level.

W.E. Friedman (2017): Insights into how the world turned green. In PDF, New Phytologist.

W.E. Friedman et al. (2004): The evolution of plant development. Free access, American Journal of Botany 91: 1726-1741.

W.E. Friedman and M.E. Cook (2000): The origin and early evolution of tracheids in vascular plants: integration of palaeobotanical and neobotanical data. In PDF, Phil.Trans. R. Soc. Lond. B, 355: 857-868.
See also here.
! Note figure 2. The three major types of early tracheids.

! E.M. Friis et al. (2011): Early Flowers and Angiosperm Evolution. Abstract, Cambridge University Press.
See also here (in PDF, long download time) and there (Google books).
Also worth checking out: Book Review, by P.J. Rudall, Botanical Journal of the Linnean Society, 170. In PDF.
"... This long-awaited book represents not only a remarkable tour de force of palaeobotanical literature, but also a potentially enduring biological textbook. ..."

Else Marie Friis, Kaj Raunsgaard Pedersen and Peter R. Crane (2010): Diversity in obscurity: fossil flowers and the early history of angiosperms. PDF file, Phil. Trans. R. Soc. B, 365: 369-382. Some of the specimens are charcoalified and have retained their original three-dimensional shape. See also here.

! M.W. Frohlich & M.W. Chase (2007): After a dozen years of progress the origin of angiosperms is still a great mystery. In PDF, Nature, 450: 1184-1189.
See also here.

! Q. Fu et al. (2023): Micro-CT results exhibit ovules enclosed in the ovaries of Nanjinganthus. Open access, Scientific Reports, 13.
Note figure 4: Micro-CT results exhibit ovules enclosed in the ovaries of Nanjinganthus.

J.M.R. Fürst-Jansen et al. (2020): Evo-physio: on stress responses and the earliest land plants. Free access, Journal of Experimental Botany, 71: 3254–3269.

! D.J. Futuyma and A.A. Agrawal (2009): Macroevolution and the biological diversity of plants and herbivores. In PDF.

! Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine: Plant Associations of the Mesophytic. Lecture Notes.

J.F. Genise et al. (2020): 100 Ma sweat bee nests: Early and rapid co-diversification of crown bees and flowering plants. Open access, PLoS ONE 15: e0227789.

! P.G. Gensel (2021): When did terrestrial plants arise? Abstract, Science, 373: : 736-737.
"... There has been a discrepancy in the time of land plant origination between molecular clock estimations (based on genes and RNA) and fossil record estimates (based on morphology). On page 792 of this issue, Strother and Foster (6) describe fossilized spores whose characteristics raise the possibility that land plants arose by co-opting algal genes, along with acquiring de novo genes, and that the former would account for the molecular clock predating the fossil record. ..."

! P.G. Gensel (2008): The earliest land plants. In PDF, The Annual Review of Ecology, Evolution, and Systematics, 39: 459-477.
See also here.

P. Gerrienne et al. (2016): Plant evolution and terrestrialization during Palaeozoic times - the phylogenetic context. Abstract, Review of Palaeobotany and Palynology.

Philippe Gerrienne et al. (2011): A Simple Type of Wood in Two Early Devonian Plants. Abstract, Science, 333. See also here (E. Brown, The Sacramento Bee), and there.

Philippe Gerrienne and Paul Gonez (2010): Early evolution of life cycles in embryophytes: A focus on the fossil evidence of gametophyte/sporophyte size and morphological complexity. Journal of Systematics and Evolution, 49: 1-16.

! M.R. Gibling et al. (2014): Palaeozoic co-evolution of rivers and vegetation: a synthesis of current knowledge. In PDF, Proceedings of the Geologists´ Association, 125: 524-533.

! M.R. Gibling and N.S. Davies (2012): Palaeozoic landscapes shaped by plant evolution. In PDF, Nature Geoscience, 5. See also here (abstract).

B. Gieren (2006): Die Landpflanzenevolution im Phanerozoikum aus petrographischer und geochemischer Sicht. PDF file, in German. Thesis, Georg-August-Universität, Gõttingen.

E.M. Gifford and A.S. Foster (1988): Morphology and Evolution of Vascular Plants. In PDF, 3rd edition, (New York: Freeman). See also here.

M.A. Gitzendanner et al. (2018): Methods for exploring the plant tree of life. In PDF, Applications in Plant Sciences, 6. See also here. Introduction for the special issue: Methods for Exploring the Plant Tree of Life.

R. Gorelick and K. Olson (2011): Is lack of cycad (Cycadales) diversity a result of a lack of polyploidy? Abstract, Botanical Journal of the Linnean Society, 165: 156-167.

R. Gorelick (2001): Did insect pollination cause increased seed plant diversity? PDF file, Biological Journal of the Linnean Society, 74: 407-427.

S.R. Gradstein and H. Kerp (2012): A Brief History of Plants on Earth. Google books, The Geologic Time Scale 2012. See also here (Table of contents, Elsevier).

Alan Graham (1993): 3. History of the Vegetation: Cretaceous (Maastrichian) - Tertiary. PDF file, Vol. 1. Flora of North America north of Mexico. See also here.

! Linda E. Graham et al. (2000): The origin of plants: Body plan changes contributing to a major evolutionary radiation. Abstracts, Proceedings of the National Academy of Sciences, 97: 4535-4540.
! See also at here. (in PDF).

J. Gravendyck et al. (2022): Early Angiosperms - How far can we reliably go back in the pollen record. Abstract, 11th European Palaeobotany and Palynology Conference Abstracts, Program and Proceedings, Swedish Museum of Natural History, Stockholm.

J. Gray and W. Shear (1992): Early life on land. In PDF, American Scientist.

S.F. Greb et al. (2006): Evolution and Importance of Wetlands in Earth History. PDF file, In: DiMichele, W.A., and Greb, S., eds., Wetlands Through Time: Geological Society of America, Special Publication, 399: 1-40. Rhacophyton and Archaeopteris in a Devonian wetland as well as Pennsylvanian, Permian, Triassic and Cretaceous wetland plant reconstructions.

K. Gurung et al. (2022): Climate windows of opportunity for plant expansion during the Phanerozoic. Open access, Nature Communications, 13.
Note figure 1: Approximate estimations of plant evolution and Phanerozoic time periods.
Figure 7: Potential biomass of plant functional types across the Phanerozoic.
"... we identify two key ‘windows of opportunity’ during the Ordovician and Jurassic-Paleogene capable of supporting dramatic expansions of potential plant biomass. These conditions are driven by continental dispersion, paleolatitude of continental area and a lack of glaciation, ..."

S.G. Hao and J.Z. Xue (2013): Earliest record of megaphylls and leafy structures, and their initial diversification. In PDF, Chin. Sci. Bull., 58: 2784-2793.

! J. Harholt et al. (2015): Why Plants Were Terrestrial from the Beginning. Abstract, Trends in Plant Science, 21: 96-101.

B.J. Harris et al. (2021): Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants. bioRxiv, see also here.
Note figure 3: The timescale of land plant evolution.
"... Here we investigate the evolution of the land plants (embryophytes) and their two main lineages, the tracheophytes (vascular plants) and bryophytes (non-vascular plants).
[...] extant tracheophytes and bryophytes are both highly derived; as a result, understanding the origin of land plants requires tracing character evolution across the diversity of modern lineages.

! C.J. Harrison and J.L. Morris (2017): The origin and early evolution of vascular plant shoots and leaves. Free access, Phil. Trans. R. Soc. B, 373: 20160496.

C.J. Harrison (2017): Development and genetics in the evolution of land plant body plans. In PDF, Phil. Trans. R. Soc., B 372. See also here (abstract).

M. Haworth et al. (2011): Stomatal control as a driver of plant evolution. In PDF, J. Exp. Bot., 62: 2419-2423.

Alan Haywood, Leeds: Plants and Earth History. Powerpoint presentation.

T. He et al. (2019): Fire as a key driver of Earth's biodiversity. In PDF, Biological Reviews. See also here.

T. He and B.B. Lamont (2017): Baptism by fire: the pivotal role of ancient conflagrations in evolution of the Earth´s flora. National Science Review. See also here (in PDF).

! T.A. Heath et al. (2014): The fossilized birth–death process for coherent calibration of divergence-time estimates. Open access, Proceedings of the National Academy of Sciences, USA, 111: E2957–E2966.

Scott A. Heckathorn, Department of Environmental Sciences, University of Toledo, Ohio, USA:
Biodiversity lecture notes, Powerpoint presentations. See especially:
Plant Diversity I: How Plants Colonized Land.
Plant Diversity II:&xnbsp; The Evolution of Seed Plants.
These expired links are available through the Internet Archive´s Wayback Machine.

J.B. Hedges (2004): A molecular timescale of eukaryote evolution and the rise of complex multicellular life. BMC evolutionary biology.

University of Heidelberg, Germany:
Aktuelle Themen in der pflanzlichen Biodiversitätsforschung (in PDF). Lecture notes, in German.
J. Griller, PDF page 1-18: "Die verwandtschaftliche Stellung der Moose".
A. Olbrich, PDF page 19-55: "Verwandtschaftliche Beziehungen der Farnpflanzen".
(?), PDF page 56-92: "Die verwandtschaftliche Stellung der Gymnospermen".
F. Haßfeld, PDF page 93-102: "Die verwandtschaftliche Stellung der Angiospermen".

! P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

F. Herrera et al. (2015): A New Voltzian Seed Cone from the Early Cretaceous of Mongolia and Its Implications for the Evolution of Ancient Conifers. In PDF, Int. J. Plant Sci., 176: 791-809.

J. Herting and T. Stützel (2021): Evolution of the coniferous seed scale. Free access, Annals of Botany.

A.J. Hetherington and L. Dolan (2019): Rhynie chert fossils demonstrate the independent origin and gradual evolution of lycophyte roots. Abstract, Current opinion in plant biology, 47: 119-126. See also here and there (in PDF).

A.J. Hetherington and L. Dolan (2018): Stepwise and independent origins of roots among land plants. Free access, Nature, 561: 235–238. Author manuscript; available in PMC 2019.

T.E. Higham et al. (2022): The Evolution of Mechanical Properties of Conifer and Angiosperm Woods. In PDF, Integrative and Comparative Biology, 62: 668–682.
See also here.

! M.F. Hohmann-Marriott and R.E. Blankenship (2011): Evolution of Photosynthesis. In PDF, Annual Review of Plant Biology, 62: 515-548.
See also here.
Note figure 2: Evolution of life and photosynthesis in geological context, highlighting the emergence of groups of photosynthetic organisms.

Kent E. Holsinger, Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT: Reproductive systems and evolution in vascular plants (PDF file).

M. Hrabovský (2020): LEAF EVOLUTION AND CLASSIFICATION. 1. LYCOPODIOPSIDA. In PDF, Acta Botanica Universitatis Comenianae, 55.

M. Hübers and H. Kerp (2012): Oldest known mosses discovered in Mississippian (late Visean) strata of Germany. In PDF, Geology, 40: 755–758.
See also here.

I.B. Huegele and S.R. Manchester (2020): An Early Paleocene Carpoflora from the Denver Basin of Colorado, USA, and Its Implications for Plant-Animal Interactions and Fruit Size Evolution. Free access, Int. J. Plant Sci., 181: 646–665.

! C.E. Hughes et al. (2015): Evolutionary plant radiations: where, when, why and how? In PDF, New Phytologist, 207: 249-253. See also here.

! Norman F. Hughes (1994): The Enigma of Angiosperm Origins. 405 pages. Provided by Cambridge University Press through the Google Print Publisher Program.

Norman F. Hughes (1982): Palaeobiology of Angiosperm Origins: Problems of Mesozoic Seed-Plant Evolution. Provided by Google books.

A. Ielpi et al. (2022): The impact of vegetation on meandering rivers. In PDF, Nature Reviews Earth & Environment, 3: 165–178.
See also here.
! Note fig. 2: Graphical timeline summary of the main evolutionary and fluvial-geomorphic events that accompanied the Palaeozoic rise of land plants, with select plant types and their approximate first appearance.
! Fig. 4: Meandering rivers in barren and vegetated landscapes.

K. Ishizaki (2017): Evolution of land plants: insights from molecular studies on basal lineages. Abstract, Bioscience, Biotechnology, and Biochemistry, 81.

K. Jordan et al.(2008): An Interactive Timeline of Plant Evolution. See also here (in PDF). Note fig. 1.

N.A. Jud (2015): Fossil evidence for a herbaceous diversification of early eudicot angiosperms during the Early Cretaceous. In PDF, Proc. R. Soc., B, 282. See also here.

Kenneth G. Karol, Richard M. McCourt, Matthew T. Cimino, and Charles F. Delwiche, Science Magazine: The Closest Living Relatives of Land Plants. This analysis supports the hypothesis that the land plants are placed phylogenetically within the Charophyta, identifies the Charales (stoneworts) as the closest living relatives of plants.

! O. Katz (2018): Extending the scope of Darwin’s ‘abominable mystery’: integrative approaches to understanding angiosperm origins and species richness. Open access, Annals of Botany, 121: 1–8.
See also here (Botany One).

M. Alan Kazlev and Toby White, Australia: Palaeos: The trace of Life on Earth. The Palaeos Site is dedicated to providing a detailed and - at least in parts - comprehensive overview of the history of life on Earth. Use the menu bars at the top and (in longer pages) bottom of the page to navigate. Go to: The Evolutionary History of Green Plants. All in a nutshell about green plant evolution-related topics, including textbook illustrations and the coverage of major phyla and kingdoms.

J.E. Keeley et al. (2011): Fire as an evolutionary pressure shaping plant traits. PDF file, Trends in Plant Science, 16.

Kelber, K.-P. (2003): Sterben und Neubeginn im Spiegel der Paläofloren. PDF file (17 MB!), in German. Plant evolution, the fossil record of plants and the aftermath of mass extinction events. pp. 38-59, 212-215; In: Hansch, W. (ed.): Katastrophen in der Erdgeschichte - Wendezeiten des Lebens.- museo 19, Heilbronn.

! P. Kenrick (2017): Changing expressions: a hypothesis for the origin of the vascular plant life cycle. Free access, Phil. Trans. R. Soc. B, 373: 20170149.
Note reconstructions of early land plants in fig. 4 and 5: Aglaophyton majus, Horneophyton lignieri, Remyophyton delicatum, Lyonophyton rhyniense, Lycopodium annotinum.

P. Kenrick and C. Strullu-Derrien (2014): The Origin and Early Evolution of Roots. In PDF, Plant Physiology, 166: 570-580. See also here (abstract).

! P. Kenrick et al. (2012): A timeline for terrestrialization: consequences for the carbon cycle in the Palaeozoic. In PDF, Philosophical Transactions of the Royal Society B, 367: 519-536.
Website saved by the Internet Archive´s Wayback Machine.

! Paul Kenrick (2011): Timescales and timetrees. PDF file, New Phytologist, 192. See also here.

P. Kenrick (2000): The relationships of vascular plants. PDF file.

! Paul Kenrick & Peter R. Crane: The origin and early evolution of plants on land. PDF file, Nature 1997.
The link is to a version archived by the Internet Archive´s Wayback Machine.

! H. Kerp et al. (2020): Plants, spores, and pollen. Pdf file, in: F.M. Gradstein et al. (eds.): The Geological Time Scale 2020.

Hans Kerp, Palaeobotanical Research Group, Westfälische Wilhelms University, Münster: A History of Palaeozoic Forests. An introductory text with many helpful links directly related to the history of Palaeozoic forests. 7 chapters provide information about: The earliest land plants; Towards a tree-like growth habit; The earliest forests; The Carboniferous coal swamp forests; The floral change at the end of the Westphalian; Stefanian and Rotliegend floras; Is there a floral break in the Permian?
Now provided by the Internet Archive´s Wayback Machine.

Michael Knee, Department of Horticulture and Crop Science, Ohio State University, Columbus: General Plant Biology Online Resources. Lecture notes.
Available through the Internet Archive´s Wayback Machine.

! A.H. Knoll and M.A. Nowak (2017): The timetable of evolution. Free access, Science Advances, 3.
Note fig. 1: The evolutionary timetable, showing the course of evolution as inferred from fossils, environmental proxies, and high-resolution geochronology.

A.H. Knoll (2014): Paleobiological Perspectives on Early Eukaryotic Evolution. In PDF, see also here.

M. Koltzenburg and G. Weitbrecht, Reutlingen, Germany: Floren- und Vegetationsgeschichte Plant evolution in a nutshell (DOC file, in German).
Now recovered from the Internet Archive´s Wayback Machine.

J. Kowal et al. (2018): From rhizoids to roots? Experimental evidence of mutualism between liverworts and ascomycete fungi. In PDF, Annals Of Botany, 121: 221-227. See also here.

Valentin A. Krassilov (1987): Palaeobotany of the mesophyticum: state of the art. In PDF, Review of Palaeobotany and Palynology, 50: 231-254. Provided by the Internet Archive´s Wayback Machine.

! M. Krings et al. (2012): Fungal Endophytes as a Driving Force in Land Plant Evolution: Evidence from the Fossil Record. In PDF; D. Southworth (ed.): Biocomplexity of Plant-Fungal Interactions (John Wiley & Sons).

U. Kutschera and K.J. Niklas (2004): The modern theory of biological evolution: an expanded synthesis. PDF file, Naturwissenschaften, 91: 255-276.

! C.C. Labandeira et al. (2016): The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies. See also here (in PDF). Proc. R. Soc., B 283.
Heritagedaily: Paleobotanist plays role in discovery of "Jurassic butterflies". An artist´s rendering of the butterfly Oregramma illecebrosa, consuming pollen drops from Triassic bennettitales.

Conrad C. Labandeira (2010): The Pollination of Mid Mesozoic Seed Plants and the Early History of Long-proboscid Insects. PDF file, Annals of the Missouri Botanical Garden, 97: 469-513. See also here.

C.C. Labandeira (2005): Invasion of the continents: cyanobacterial crusts to tree-inhabiting arthropods. In PDF, Trends in Ecology and Evolution, 20.

B. Laenen et al. (2014): Extant diversity of bryophytes emerged from successive post-Mesozoic diversification bursts. In PDF. See also here (abstract).

A.D.B. Leakey and J.A. Lau (2012): Evolutionary context for understanding and manipulating plant responses to past, present and future atmospheric [CO2]. Phil. Trans. R. Soc. B, 367: 613-629. See als here (in PDF).

A.R. Leitch and I.J. Leitch (2012): Ecological and genetic factors linked to contrasting genome dynamics in seed plants. In PDF, New Phytologist, 194: 629-646.

S. Lehtonen et al. (2020): Exploring the phylogeny of the marattialean ferns. Open access, Cladistics.
Note fig. 4: Parsimony-dated phylogeny and Bayesian historical biogeography of the marattialean ferns.
"... We resolved the fossil genera Marattiopsis, Danaeopsis and Qasimia as members of the monophyletic family Marattiaceae, and the Carboniferous genera Sydneia and Radstockia as the monophyletic sister of all other marattialean ferns. ..."

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T.M. Lenton et al. (2016): Earliest land plants created modern levels of atmospheric oxygen. Free access, PNAS, 113.

! T.M. Lenton (2001): The role of land plants, phosphorus weathering and fire in the rise and regulation of atmospheric oxygen. In PDF, Global Change Biology, 7: 613-629.

! K. Lepot (2020): Signatures of early microbial life from the Archean (4 to 2.5 Ga) eon. Free access, Earth-Science Reviews, 209. See also here.

A.B. Leslie and J.M. Losada (2019): Reproductive Ontogeny and the Evolution of Morphological Diversity in Conifers and Other Plants. Free access, Integrative and Comparative Biology, 59: 548–558.
Note figure 4: Depictions of ovuliferous complex and ovule/seed ontogeny in different conifer groups through time.

A.B. Leslie et al. (2018): An overview of extant conifer evolution from the perspective of the fossil record. Abstract, American Journal of Botany, 105: 1–14. See also here (in PDF).

! A.B. Leslie et al. (2015): Integration and macroevolutionary patterns in the pollination biology of conifers. In PDF, Evolution, 69: 1573-1583.

! A.B. Leslie et al. (2012): Hemisphere-scale differences in conifer evolutionary dynamics. In PDF, PNAS, 109: 16217-16221. See also here.

Gerhard Leubner, The Seed Biology Place, University Freiburg, Germany: Seed evolution. Origin and evolution of the seed habit.

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"... With a well-resolved plastid tree and 62?fossil calibrations, we dated the origin of the crown angiosperms to the Upper Triassic, with major angiosperm radiations occurring in the Jurassic and Lower Cretaceous. This estimated crown age is substantially earlier than that of unequivocal angiosperm fossils, and the difference is here termed the ‘Jurassic angiosperm gap’. ..."

F.-W. Li et al. (2018): Fern genomes elucidate land plant evolution and cyanobacterial symbioses. Open access, Nature Plants, 4: 460–472.

! R. Ligrone et al. (2012): Major transitions in the evolution of early land plants: a bryological perspective. Free access, Annals of botany, 109: 851–871.

! A. Linkies et al. (2010). The evolution of seeds. In PDF, New Phytologist, 186: 817-831. See also here (abstract).

B.H. Lomax et al. (2014): Reconstructing relative genome size of vascular plants through geological time. Free access, New Phytologist, 201: 636–644.

! C.V. Looy et al. (2014): The late Paleozoic ecological-evolutionary laboratory, a land-plant fossil record perspective. In PDF, The Sedimentary Record, 12: 4-18. See also here.

! G.Q. Liu et al. (2022): The Molecular Phylogeny of Land Plants: Progress and Future Prospects. Open access, Diversity, 14 (from the Special Issue Ecology, Evolution and Diversity of Plants).
Note figure 1: Summary of phylogenetic relationships among major clades of land plants.

! Y. Liu et al. (2022): The Cycas genome and the early evolution of seed plants. Open access, Nature Plants, 8: 389–401.
"... Although the major cycad lineages are ancient, modern cycad species emerged from several relatively recent diversifications ..."

Y. Lu et al. (2014): Phylogeny and divergence times of gymnosperms inferred from single-copy nuclear genes. PloS one.

! W.J. Lucas et al. (2013): The Plant Vascular System: Evolution, Development and Functions. In PDF, Journal of Integrative Plant Biology, 55: 294-388. See also here.

S. Magallón et al. (2015): A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity. In PDF, New Phytologist. See also here.

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! S. Magallón (2009): Flowering plants (Magnoliophyta). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

S. Magallón and A. Castillo (2009): Angiosperm diversification through time. Free access, American Journal of Botany, 96: 349-365.

! S. Magallón et al. (2013): Land plant evolutionary timeline: Gene effects are secondary to fossil constraints in relaxed clock estimation of age and substitution rates. Open access, American Journal of Botany, 100: 556-573.

! Susana Magallón and Khidir W. Hilu (2009): Land plants (Embryophyta). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).
These expired links are now available through the Internet Archive´s Wayback Machine.

K. Mao et al. (2012): Distribution of living Cupressaceae reflects the breakup of Pangea. In PDF, Proc. Natl. Acad. Sci., 109: 7793-7798.

P.D. Mannion et al. (2014): The latitudinal biodiversity gradient through deep time. In PDF, Trends in Ecology & Evolution, 29: 42–50. See also here.

A.O. Marron et al. (2016): The Evolution of Silicon Transport in Eukaryotes. In PDF, Mol. Biol. Evol. See also here.

! W.F. Martin and J.F. Allen (2018): An algal greening of land. Free access, Cell, 174: 256-258. See also here.
Note figure 1: Streptophyte Algae and the Rise of Atmospheric Oxygen.

! F.M. Martin et al. (2017): Ancestral alliances: Plant mutualistic symbioses with fungi and bacteria. In PDF, Science, 356. See also here.

! C. Martín-Closas (2003): The fossil record and evolution of freshwater plants: a review. PDF file, Geologica Acta, 1: 315-338.

Patrick T. Martone et al. (2009): Discovery of Lignin in Seaweed Reveals Convergent Evolution of Cell-Wall Architecture. Abstract, Current Biology, Volume 19, Issue 2, 169-175. See also here.

N.P. Maslova et al. (2021): Recent Studies of Co-Evolutionary Relationships of Fossil Plants and Fungi: Success, Problems, Prospects. In PDF, Paleontological Journal, 55: 1–17. See also here.

! S. Mathews (2009): Phylogenetic relationships among seed plants: persistent questions and the limits of molecular data. Free access, American Journal of Botany, 96: 228-236.

W.J. Matthaeus et al. (2022): Stems matter: Xylem physiological limits are an accessible and critical improvement to models of plant gas exchange in deep time. In PDF, Front. Ecol. Evol., 10:955066. doi: 10.3389/fevo.2022.955066.
See also here.
Note figure 1: Vascular plant hydraulic pathway conducting element features.

M.R. May et al. (2021): Inferring the Total-Evidence Timescale of Marattialean Fern Evolution in the Face of Model Sensitivity. Free access, Systematic Biology, syab020, See also here.

S.A.M. McAdam et al. (2021): Stomata: the holey grail of plant evolution. In PDF, Am. J. Bot., 108: 366–371. See also here.

J.C. McElwain et al. (2016): Assessing the role of atmospheric oxygen in plant evolution. Abstract, starting on PDF page 44.
Abstracts, XIV International Palynological Congress, X International Organisation of Palaeobotany Conference, Salvador, Brazil.

! J.C. McElwain and S.W. Punyasena (2007): Mass extinction events and the plant fossil record. PDF file, Trends in Ecology and Evolution, 22: 548-557. See also here (abstract).

! S. McLoughlin (2021): Gymnosperms: History of Life: Plants: Gymnosperms. In PDF, p. 476-500; In: Elias, S. & Alderton, D. (eds.), Encyclopedia of Geology, Amsterdam, Elsevier. See also here.
! Note fig. 8: One model for the evolution of seed plants showing the stratigraphic ranges and relative abundance of the major groups.

S. McLoughlin (2017): Antarctica’s Glossopteris forests. In PDF, In: 52 More Things You Should Know About Palaeontology, eds. A. Cullum, A.W. Martinius. Nova Scotia: Agile Libre, p. 22-23. See also here.

S. McLoughlin and B.P. Kear (2014): Gondwanan Mesozoic biotas and bioevents. Abstract.

! Stephen McLoughlin (2001): The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. In PDF, Australian Journal of Botany, 49: 271-300. See also here (abstract).

! B.J.W. Mills et al. (2017): Nutrient acquisition by symbiotic fungi governs Palaeozoic climate transition. Open access, Phil. Trans. R. Soc. B, 373.

R.L. Mitchell et al. (2023): Terrestrial surface stabilisation by modern analogues of the earliest land plants: A multi-dimensional imaging study. Open access, Geobiology.
Note figure 1: Summary chart highlighting the evolution of different CGC elements [cryptogamic ground covers] from contrasting molecular, phylogenetic and fossil dating methods, and schematic land plant phylogeny of modern terrestrial organisms, focussing on the bryophytes and specific liverwort genera.

Sebastian Molnar, Department of Zoology, University of British Columbia, Vancouver: Evolution and the Origins of Life. A directory of introductions concerning evolution, with a bias to Plant Biology and Evolution. Excellent examples about how evolution works can be seen from the plant world. Go to: Angiosperm Origins and Evolution, or The Evolution of Polyploidy, and Summary: Polyploid Evolution.

! J.L. Morris et al. (2018): The timescale of early land plant evolution. In PDF, PNAS, 115. See also here.

! J.L. Morris et al. (2015): Investigating Devonian trees as geo-engineers of past climates: linking palaeosols to palaeobotany and experimental geobiology. In PDF, Palaeontology, 58: 787-801. See also here.

! Terence M. Murphy, Thomas L. Rost and Michael G. Barbour (2015), University of California, Davis, CA:
Plant Biology. Lecture notes, in PDF. Please take notice: Book announcement.
See for instance: Bryophytes.
The Early Tracheophytes.

N.S. Nagalingum et al. (2011): Recent Synchronous Radiation of a Living Fossil. Abstract.
"Using fossil-calibrated molecular phylogenies, we show that cycads underwent a near synchronous global rediversification beginning in the late Miocene, followed by a slowdown toward the Recent. Although the cycad lineage is ancient, our timetrees indicate that living cycad species are not much older than ~12 million years". See also here. In PDF, Science 334.

Y. Nie et al. (2020): Accounting for uncertainty in the evolutionary timescale of green plants through clock-partitioning and fossil calibration strategies. In PDF, Syst. Biol., 69: 1–16. See also here.

Nature Science Update (December 8, 1999): One for the Vine. "A prickly climbing vine", Vasovinea tianii (Gigantopteridales), that lived more than 250 million years ago could shed light on the origin of flowering plants.
This expired link is available through the Internet Archive´s Wayback Machine.

M.P. Nelsen et al. (2020): No support for the emergence of lichens prior to the evolution of vascular plants. In PDF, Gebiology, 18: 3-13. See also here.

A.B. Nicotra et al. (2011): The evolution and functional significance of leaf shape in the angiosperms. In PDF, Functional Plant Biology, 38: 535-552. See also here.

! Y. Nie et al. (2020): Accounting for uncertainty in the evolutionary timescale of green plants through clock-partitioning and fossil calibration strategies. In PDF, Syst. Biol., 69: 1–16. See also here.
! Note figure 5: Time-tree of green plants.
! "... By taking into account various sources of uncertainty, we estimate that crown-group green plants originated in the Paleoproterozoic–Mesoproterozoic (1679.7–1025.6 Ma), crown-group Chlorophyta and Streptophyta originated in the Mesoproterozoic–Neoproterozoic (1480.0–902.9 Ma and 1571.8–940.9 Ma), and crown-group land plants originated in the Ediacaran to middle Ordovician (559.3– 459.9 Ma). ..."

S. Nigris et al. (2021): Fleshy Structures Associated with Ovule Protection and Seed Dispersal in Gymnosperms: A Systematic and Evolutionary Overview. Open access, Critical Reviews in Plant Sciences, 40.

K.J. Niklas and B.H. Tiffney (2022): Viridiplantae Body Plans Viewed Through the Lens of the Fossil Record and Molecular Biology. Open access, Integrative and Comparative Biology,
"... A review of the fossil record coupled with insights gained from molecular and developmental biology reveal a series of body plan transformations that gave rise to the first land plants. Across diverse algal clades, including the green algae and their descendants, the plant body plan underwent a unicellular -- colonial -- simple multicellular -- complex multicellular transformation series. ..."
Note figure 4: Scenarios for the evolution of the first land plant sporophyte resulting from delayed zygotic meiosis.

K.J. Niklas et al. (2017): The evolution of hydrophobic cell wall biopolymers: from algae to angiosperms. Abstract, J. Exp. Bot.

Karl J. Niklas (2016): Plant Evolution: An Introduction to the History of Life. Book announcement.
Worth checking out: ! Introduction.
See also here (Google books).

! K.J. Niklas (2015): Measuring the tempo of plant death and birth. Open access, New Phytologist.

! K.J. Niklas and U. Kutschera (2010): The evolution of the land plant life cycle. Free access, New Phytologist, 185: 27-41.

K.J. Niklas (2004): Computer models of early land plant evolution. In PDF, Annu. Rev. Earth Planet. Sci., 32: 47-66.

Karl Niklas, Plant Biology, Cornell University (page hosted by Access Excellence): Plant Evolution: Adaptation or Historical Accident?. See also here, and there.

K.J. Niklas and T. Speck (2001): Evolutionary trends in safety factors against wind-induced stem failure. Open access, American Journal of Botany, 88: 1266-1278.

! K.J. Niklas (2000): The Evolution of Plant Body Plans - A Biomechanical Perspective. In PDF, Annals of Botany, 85: 411-438.
See also here.

! K.J. Niklas (1994): Morphological Evolution Through Complex Domains of Fitness. In: Fitch, W.M. And Ayala, F.J. (eds.):
! Tempo And Mode In Evolution: Genetics And Paleontology 50 Years After Simpson. Open access! National Academies Press (US); Washington (DC).

Karl Niklas, (Section of Plant Biology, Cornell University), Access Excellance, BioForum 4, "Theoretical Issues in Plant Biology". Now available by the Internet Archive´s Wayback Machine.
BioForum is a series of lectures, presented by California Academy of Sciences, in which scientists share their research results with high school biology teachers. Plant Evolution: Adaptation or Historical Accident? Is plant evolution largely adaptive or is it contingent on historical accidents? Dr. Niklas discuss in some detail a computer generated model dealing with the early evolution of land plants that can be used to predict how plant architecture must change to assure evolutionary success as the environment changes.

H. Nowak et al. (2020): Palaeophytogeographical Patterns Across the Permian–Triassic Boundary. Open access, Front. Earth Sci.

T. Nyman et al. (2012): Climate-driven diversity dynamics in plants and plant-feeding insects. Free access, Ecology Letters, 14: 1-10.

! S.L. Olson et al. (2018): Earth: Atmospheric Evolution of a Habitable Planet. PDF file, In: Deeg H., Belmonte J. (eds.) Handbook of Exoplanets. Springer. See also here.
Worth checking out: Figure 2, co-evolution of life and surface environments on Earth.

A.A. Óladóttir, Iceland GeoSurvey, Reykjavik, Iceland: An Introduction to the Mesozoic Palaeobotany. In PDF.

J. Ollerton and E. Coulthard (2009): Evolution of Animal Pollination. In PDF, Science, 326.

Mark E. Olson (2012): Linear Trends in Botanical Systematics and the Major Trends of Xylem Evolution. In PDF.

! R. Omlor and J.W. Kadereit (2005), Institut für Spezielle Botanik und Botanischer Garten, Mainz: Evolution der Landpflanzen (in German).

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See also here.

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Barry A. Palevitz, "Discovering Relatives in the Flowering Plant Family Tree". The Scientist, Volume 13, 1999: Search for: "flowering plant". Registration procedure required.

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! J.G. Pausas et al. (2018): Unearthing belowground bud banks in fire-prone ecosystems. Free access, New Phytologist, 217: 1387-1778.
Note figure 1: Stylized diagrams of 16 belowground bud bank (BBB) structures that enable plants to resprout following fire.
Figure 3: Oldest time of origin for different belowground bud bank (BBB) organs in selected angiosperm families.
"... Recognizing the diversity of BBBs provides a basis for understanding the many evolutionary pathways available to plants for responding to severe recurrent disturbances. ..."

J.G. Pausas et al. (2017): Flammability as an ecological and evolutionary driver. In PDF, Journal of Ecology, 105: 289–297.

J.G. Pausas and D. Schwilk (2012): Fire and plant evolution. Free access, New Phytologist, 193: 301-303.

£. Pawlik et al. 2020): Impact of trees and forests on the Devonian landscape and weathering processes with implications to the global Earth's system properties – A critical review. In PDF, Earth-Science Reviews, 205: doi 10.1016/j.earscirev.2020.103200.
See also here.
Note fig. 2. Spatial configuration of continents in the Devonian.
Note fig. 3: Landscape reconstruction with stands of Pseudosporochnus, up to 4 m high, with Protopteridium in shruby layer and herbaceous Drepanophycus and Protolepidodendron in understorey.
Note fig. 6: A close look at trees diversification and selected accompanying events in the Devonian.

Pearson Education, Inc.: Overview of Land Plant Evolution. Powerpoint presentation.
Snapshot taken by the Internet Archive´s Wayback Machine.

! D. Peris et al. (2017): False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance. In PDF, Current Biology, 27. See also here.

! Rémy J. Petit and Arndt Hampe (2006): Some Evolutionary Consequences of Being a Tree. PDF file, Annu. Rev. Ecol. Evol. Syst., 37: 187-214.

D. Peyrot et al. (2019): The greening of Western Australian landscapes: the Phanerozoic plant record. Journal of the Royal Society of Western Australia, 102: 52-82. See also here. Worth checking out:
! Figure 9: Major plant-evolutionary events and vegetation changes in Western Australia.

H.W. Pfefferkorn et al. (2017): Impact of an icehouse climate interval on tropical vegetation and plant evolution. In PDF, Stratigraphy, 14: 365-376. See also here. (in German):
Der Farnsamer aus dem Perm (2023).
Landeroberung früher als gedacht. Bereits die Vorfahren heutiger Landpflanzen haben an Land gelebt (2016).
Die Entstehung der Pflanzenwelt (2013).
The link is to a version archived by the Internet Archive´s Wayback Machine.

! A. Piombino (2016): The Heavy Links between Geological Events and Vascular Plants Evolution: A Brief Outline. In PDF, International journal of evolutionary biology.

N.D. Pires and L. Dolan (2012): Morphological evolution in land plants: new designs with old genes. In PDF, Philosophical Transactions of the Royal Society B, 367: 508-518.

! J. Pittermann et al. (2015): The structure and function of xylem in seed-free vascular plants: an evolutionary perspective. In PDF. See also here.

J. Pittermann (2010): The evolution of water transport in plants: an integrated approach. In PDF, Geobiology.
See also here.

! A.R.G. Plackett and J.C. Coates (2016): Life’s a beach – the colonization of the terrestrial environment. In PDF, New Phytologist, 212: 831–835. See also here.

Tõnu Ploompuu, Biology, Tallinn Pedagogical University, Tallinn, Estonia: Resting and active evolution. Possible preadaptations in the early evolution of Angiosperms. See also here.

George Poinar and Greg Poinar (2018): The antiquity of floral secretory tissues that provide today’s fragrances. Abstract, Historical Biology. See also:
Schnupperten schon Dinos Blumenduft? Kreidezeitliche Blütenpflanzen könnten bereits Düfte produziert haben. In German,

Z.A. Popper et al. (2011): Evolution and Diversity of Plant Cell Walls: From Algae to Flowering Plants. In PDF, Annu. Rev. Plant Biol., 62: 567-590.

Z.A. Popper and M.G. Tuohy (2010): Beyond the Green: Understanding the Evolutionary Puzzle of Plant and Algal Cell Walls. PDF file, Plant Physiology, 53: 373-383.

! J.C. Preston et al. 2022): Plant structure and function: Evolutionary origins and underlying mechanisms. Free access, Plant Physiology.

Ernst Probst, fossilien-news, Mainz-Kostheim, Germany: Rekorde der Urzeit aus der Pflanzenwelt (in German).
The link is to a version archived by the Internet Archive´s Wayback Machine.

! K.M. Pryer et al. (2004): Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. Open access, American Journal of Botany, 91: 1582-1598.

J. Pšenicka et al. (2021): Dynamics of Silurian plants as response to climate changes. Open access, Life, 11.
Note figure 1: Silurian time scale showing conodont and graptolite biozones, stage slices and generalized 13Ccarb curve.
Figure 2: Silurian palaeocontinental reconstructions.

C. Puginier et al. (2021): Plant–microbe interactions that have impacted plant terrestrializations. Free access, Plant Physiology.
Note figure 1: 1 Phylogenetic tree of the Viridiplantae. showing the evolution of the AMS [arbuscular mycorrhizal symbiosis], the putative evolutions of lichens and clades that contain LFA [lichen forming algae] and terrestrial species.
Figure 3: Lichens and their tolerance against terrestrial-related constraints.

! M.N. Puttick et al. (2018): The Interrelationships of Land Plants and the Nature of the Ancestral Embryophyte. Free access, Current Biology, 28: 733–745.

! Y.-L. Qiu et al. (2006): The deepest divergences in land plants inferred from phylogenomic evidence. In PDF, PNAS, 103: 15511-15516

Yin-Long Qiu and Jeffrey D. Palmer (1999): Phylogeny of early land plants: insights from genes and genomes. In PDF.
Now recovered from the Internet Archive´s Wayback Machine.

J. Quirk et al. (2015): Constraining the role of early land plants in Palaeozoic weathering and global cooling. Proc. R. Soc., B 282.

! L. Ragnia and T. Greb (2018): Secondary growth as a determinant of plant shape and form. Open access, Seminars in Cell & Developmental Biology, 79: 58-67.

! J.-H. Ran et al. (2018): Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms. Abstract.

S. Ratti et al. (2011): Did Sulfate Availability Facilitate the Evolutionary Expansion of Chlorophyll a+c Phytoplankton in the Oceans? In PDF, Geobiology 9, no. 4: 301–312. See also here (abstract).

J.A. Raven (2017): Evolution and palaeophysiology of the vascular system and other means of long-distance transport. In PDF, Phil. Trans. R. Soc. B, 373: 20160497.

J.A. Raven and M. Andrews (2010): Evolution of tree nutrition. In PDF, Tree Physiology, 30: 1050-1071. See also here.

J.A. Raven (2002): Selection pressures on stomatal evolution. PDF file, New Phytologist.

J.A. Raven and D. Edwards (2001): Roots: evolutionary origins and biogeochemical significance. PDF file, J. Exp. Bot., 52: 381-401.

R.R. Reisz and J. Müller (2004): Molecular timescales and the fossil record: a paleontological perspective. In PDF, Trends in Genetics.

S.S. Renner (2022): Plant Evolution and Systematics 1982–2022: Changing Questions and Methods as Seen by a Participant. In PDF, Progress in Botany.
See also here.
"... With DNA data came lab work, bioinformatics, and both the need and the possibility to collaborate, which brought systematists out of their niche, gave comparative biology a huge push, and resulted in a better integration of biodiversity studies within biology. ..."

K.S. Renzaglia et al. (2000): Vegetative and reproductive innovations of early land plants: implications for a unified phylogeny. Abstract, Phil. Trans. R. Soc. Lond., B 355: 769-793.

H. Renault et al. (2017): A phenol-enriched cuticle is ancestral to lignin evolution in land plants. Nat. Commun. 8.

G.J. Retallack (2021): Great moments in plant evolution. In PDF, Proceedings of the National Academy of Sciences of the United States of America (PNAS), 118. See also here.

Authored by the The Rhynie Chert Research Group, University of Aberdeen, with contributions and support by the Palaeobotanical Research Group, University of Münster, Germany, the Centre for Palynology, University of Sheffield, The Natural History Museum, London, and The Royal Museum, National Museums of Scotland: The Biota of Early Terrestrial Ecosystems, The Rhynie Chert. A resource site for students and teachers covering many aspects of the present knowledge of this unique geological deposit (including a glossary and bibliography pages). The website´s second part provides guidance for teachers in this subject area and as such will require a password to enter (obtainable from the authors).

J.D. Richey et al. (2021): Modeled physiological mechanisms for observed changes in the late Paleozoic plant fossil record. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 562.
"... (1) The existence of pCO2 and precipitation thresholds for loss of physiological viability that provide a mechanism for replacement of wet-adapted lycopsids and medullosans by marattialean tree ferns, which were tolerant of periodic drought, and the subsequent dominance of seasonally dry-adapted cordaitaleans and conifers. ...
(2) Under drier conditions, the combination of higher drought tolerance and primary productivity for marattialean tree ferns, conifers, and cordaitaleans provided an ecophysiological advantage over lycopsids and medullosans. ...
although the shift to more drought-tolerant plants in the Late Pennsylvanian and early Permian could have led to increased biomass and surface runoff, their ability to affect climate was likely limited by aridity and changes in vegetation density. ..."

J.P. Rose et al. (2016): Shape analysis of moss (Bryophyta) sporophytes: Insights into land plant evolution. Free access, Am. J. Bot., 103: 652-662.

Anita Roth-Nebelsick et al. (2001): Evolution and Function of Leaf Venation Architecture: A Review. PDF file, Annals of Botany 87: 553-566. See also here.

G.W. Rothwell et al. (2018): Tree of death: The role of fossils in resolving the overall pattern of plant phylogeny. Free access, American Journal of Botany, 105: 1–4. See also here and there.

! G.W. Rothwell et al. (2014): Plant evolution at the interface of paleontology and developmental biology: An organism-centered paradigm. Open access, Am. J. Bot., 101: 899-913.

G.W. Rothwell et al. (2012): The seed cone Eathiestrobus gen. nov.: Fossil evidence for a Jurassic origin of Pinaceae. In PDF, American Journal of Botany, 99: 708–720.

G.W. Rothwell et al. (2009): Is the anthophyte hypothesis alive and well? New evidence from the reproductive structures of Bennettitales. Free access, American Journal of Botany, 96: 296-322. Note fig. 1: Cycadeoidea spp. Characteristic features of Cycadeoidea plants.
! Table 2. Contrasting characters of Bennettitales and Cycadales.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens, OH: Vascular Plant Morphology. Archived by Internet Archive Wayback Machine. This course covers the structure, development, reproductive biology and relationships of vascular plants. The course is structured to emphasize the evolutionary changes that led to the diversity of modern tracheophytes. Go to Cordaitales and Coniferales (PDF file).

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens: Angiophytes: Using Whole Plant Concepts to Interpret Angiosperm Origins.
The link is to a version archived by the Internet Archive´s Wayback Machine.

Nick Rowe and Thomas Speck (2005): Plant growth forms: an ecological and evolutionary perspective. PDF file, New Phytologist, 166: 61-72. See also here.

Dmitry A. Ruban (2012): Mesozoic mass extinctions and angiosperm radiation: does the molecular clock tell something new? In PDF.

! C.V. Rubinstein and V. Vajda (2019): Baltica cradle of early land plants? Oldest record of trilete spores and diverse cryptospore assemblages; evidence from Ordovician successions of Sweden. Free access, GFF, DOI: 10.1080/11035897.2019.1636860.

! P.J. Rudall (2021): Evolution and patterning of the ovule in seed plants. Free access, Biological Reviews. See also here.

Paula J. Rudall and Richard M. Bateman (2010): Defining the limits of flowers: the challenge of distinguishing between the evolutionary products of simple versus compound strobili. In PDF, Philos. Trans. R. Soc. London, B Biol. Sci., 365: 397-409. See also here (abstract).

! B.R. Ruhfel et al. (2014): From algae to angiosperms - inferring the phylogeny of green plants (Viridiplantae) from 360 plastid genomes. In PDF, BMC Evolutionary Biology, 14. See also here.

Tyson Sacco, Cornell University: Trends in Green Plant Evolution. Powerpoint presentation.

! L. Sack and C. Scoffoni (2013): Leaf venation: structure, function, development, evolution, ecology and applications in the past, present and future. Free access, New Phytologist, 198: 983–1000.
Note figure 6: Evolution of terrestrial plants and their traits, including leaf vein traits against geological periods and time.

R.F. Sage (2004): The evolution of C4 photosynthesis. Free access, New Phytologist, 161: 341–370.
"... C4 photosynthesis in the dicots originated in arid regions of low latitude, implicating combined effects of heat, drought and/or salinity as important conditions promoting C4 evolution. Low atmospheric CO2 is a significant contributing factor ..."

! M.A. Salamon et al. (2018): Putative Late Ordovician land plants. Free Access, New Phytologist, 218: 1305–1309.

A. Salt (2019): When did the first flowers open? Botany One.

I. Sanmartín and F. Ronquist (2004): Southern Hemisphere Biogeography Inferred by Event-Based Models: Plant versus Animal Patterns. PDF file, Syst. Biol., 53: 216-243.

! P. Sarkar et al. (2009): Plant cell walls throughout evolution: towards a molecular understanding of their design principles. In PDF, Journal of Experimental Botany, 60: 3615–3635. See also here.

H. Sauquet et al. (2022): What is the age of flowering plants? In PDF, Journal of Experimental Botany,
See also here.
Note fig. 1: Hypothetical time tree of the angiosperms.
Note fig. 2. Crown-group angiosperm age estimates obtained in fossil-calibrated molecular dating and palaeontological macroevolutionary modelling studies published over the last 6 years.

! H. Sauquet et al. (2017): The ancestral flower of angiosperms and its early diversification. Nature Communications, 8.

C. Schirarend and R. Vogt, Botanischer Garten und Botanisches Museum Berlin-Dahlem, Freie Universität Berlin: Von Nacktpflanzen und Schuppenbäumen. Plant evolution in a nutshell (in German). Part of the description of the exhibition "Stammesgeschichte der Pflanzen" (in German).
These expired links are now available through the Internet Archive´s Wayback Machine.

B.E. Schirrmeister et al. (2011): The origin of multicellularity in cyanobacteria. Open access, BMC Evolutionary Biology, 11.

H. Schneider, (2006), starting on PDF page 65: Plant morphology as the cornerstone to the integration of fossil and extant taxa in phylogenetic systematics. PDF file, in German. Species, Phylogeny and Evolution, 1. Themenheft Phylogenetisches Symposium Göttingen: Der Stellenwert der Morphologie in der heutigen Phylogenetische Systematik.
This expired link is available through the Internet Archive´s Wayback Machine.

H. Schneider et al. (2004): Ferns diversified in the shadow of angiosperms. In PDF, see also here (abstract).

! J.W. Schopf et al. (2007): Evidence of Archean life: Stromatolites and microfossils. In PDF, Precambrian Research, 158: 141-155.
See also here.

! J.W. Schopf (2006): Fossil evidence of Archaean life. In PDF, Transactions of the Royal Society, B 361: 869–885.

! M. Schreiber et al. (2022): The greening ashore. Free access, Trends in Plant Science.
"... Two decisive endosymbiotic events, the emergence of eukaryotes followed by the further incorporation of a photosynthesizing cyanobacterium, laid the foundation for the development of plant life. ..."

! E. Schuettpelz and K.M. Pryer (2008): Fern phylogeny. In PDF.

! Viridiplantae Geology, Evolution upset: Oxygen-making microbes came last, not first.

D.H. Scott (1909), President of the Linnean Society: Darwin and Modern Science: The Palaeontological Record: Plants. Website hosted by "The Unofficial Stephen Jay Gould Archive".

M. Seale (2020): The Fat of the Land: Cuticle Formation in Terrestrial Plants. Free access, Plant Physiology, 184: 1622–1624.

! M.A. Selosse and C. Strullu-Derrien (2015): Origins of the terrestrial flora: A symbiosis with fungi? In PDF, BIO Web of Conferences, 4.

M.-A. Selosse et al. (2015): Plants, fungi and oomycetes: a 400-million year affair that shapes the biosphere. New Phytologist. 10th New Phytologist Workshop on the "Origin and evolution of plants and their interactions with fungi", London, UK, September 2014.

M.A. Selosse and C. Strullu-Derrien (2015): Origins of the terrestrial flora: A symbiosis with fungi? In PDF, BIO Web of Conferences, 4.

S. Sgorbati et al. (2018): Was Charles Darwin right in his explanation of the ‘abominable mystery’? Free access, Italian Botanist, 5: 25–30.

! A.J. Shaw et al.(2011): Bryophyte diversity and evolution: Windows into the early evolution of land plants. Free access, Am. J. Bot., 98: 352-369.

! D. Silvestro et al. (2020): Fossil data support a pre-Cretaceous origin of flowering plants. In PDF, Nature Ecology & Evolution. See also here.
"... Yet, our results indicate that an early, pre-Cretaceous origin of angiosperms is supported not only by molecular phylogenetic hypotheses but also by an analysis of the fossil record ..."

D. Silvestro et al. (2016): Fossil biogeography: a new model to infer dispersal, extinction and sampling from palaeontological data. In PDF, Phil. Trans. R. Soc., B, 371. See also here.

! D. Silvestro et al. (2015): Revisiting the origin and diversification of vascular plants through a comprehensive Bayesian analysis of the fossil record. In PDF, New Phytologist, 207: 425-436.

! M.G. Simpson (2010): Evolution and diversity of green and land plants. PDF file, p. 55–72. In: Simpson MG, (ed.): Plant systematics. 2nd ed., Cambridge (MA): Academic Press.

Stephen A. Smith et al. (2010): An uncorrelated relaxed-clock analysis suggests an earlier origin for flowering plants. PDF file, PNAS, 107: 5897-5902. See also here, and there.

Stephen A. Smith and Jeremy M. Beaulieu (2009): Life history influences rates of climatic niche evolution in flowering plants. In PDF, Proc. R. Soc. B, 276: 4345-4352. See also here.

! S.A. Smith and M.J. Donoghue (2008): History in Flowering Plants Rates of Molecular Evolution Are Linked to Life. In PDF, Science, 322. See also here (abstract).

P.S. Soltis et al. (2019): Darwin review: angiosperm phylogeny and evolutionary radiations. In PDF, Proc. R. Soc. B, 286: 20190099. See also here.

D.E. Soltis et al. (2018): Using and navigating the plant tree of life. In PDF, American Journal of Botany, 105: 287–290. See also here.

! D.E. Soltis et al. (2009): Polyploidy and angiosperm diversification. Free access, Am. J. Bot., 96: 336-348.

Pamela Soltis (website by American Institute of Biological Sciences): Flowering Plants: Keys to Earth´s Evolution and Human Well-Being.

P.S. Soltis and D.E. Soltis (2004): The origin and diversification of angiosperms. Free access, American Journal of Botany, 91: 1614-1626.

! D. Soltis et al., Florida Museum of Natural History (FLMNH), University of Florida: Deep Time. A comprehensive phylogenetic tree of living and fossil angiosperms.
Deep Time explore the ways in which angiosperm fossils can be appropriately integrated into the phylogenetic framework for extant taxa, with the ultimate goal of forming a comprehensive phylogenetic tree of living and fossil angiosperms. This includes the evaluation and prioritization of the fossil record, the critical appraisal of the age of fossils, the construction of a morphological data matrix for fossils and extant angiosperms, the integration of fossils into the angiosperm tree and the calibration of divergence times.

I. Sørensen et al. (2010): How Have Plant Cell Walls Evolved? In PDF, Plant Physiology, 153: 366–372. See also here.

T. Speck and O. Speck (2019): Quo vadis plant biomechanics: Old wine in new bottles or an up-and-coming field of modern plant science? Open access, American Journal of Botany, 106: 1399-1403.

V. Spencer et al. (2020): What can lycophytes teach us about plant evolution and development? Modern perspectives on an ancient lineage. Open access, Evolution & Development, 23: 174-196 (Special Issue: Plant Evolution & Development).

! J.S. Sperry (2003): Evolution of water transport and xylem structure. PDF file, International Journal of Plant Sciences. See also here (abstract).

! R.A. Spicer (1989): Physiological characteristics of land plants in relation to environment through time. In PDF, Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 80.
See also here.

! R. Spicer and A. Groover (2010): Evolution of development of vascular cambia and secondary growth. Open access, New Phytologist, 186: 577-592.
Note figure 1: Orientation of cells and tissues within a woody stem.
Figure 2: A phylogeny of vascular plants illustrating multiple origins of secondary growth via a vascular cambium.

A.K. Srivastava, Birbal Sahni Institute of Palaeobotany, Lucknow, India: Taxonomy, palaeobotany and biodiversity About the angiosperm origin (PDF file, page 2). CURRENT SCIENCE, VOL. 81, NO. 10.

! P. Steemans et al. (2009): Origin and Radiation of the Earliest Vascular Land Plants. In PDF, Science, 324.
Now provided by the Internet Archive´s Wayback Machine.

! W.E. Stein and J.S. Boyer (2006): Evolution of land plant architecture: beyond the telome theory. In PDF, Paleobiology 32: 450-482.
See also here.

W.E. Stein (1987): Phylogenetic analysis and fossil plants. PDF file, Review of palaeobotany and palynology.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Plant life from the Silurian to the Cretaceous. Go to:
The evolution of plants. A concise report of the development of the flora.

! P.F. Stevens and Hilary Davis, Missouri Botanical Garden, St. Louis: Angiosperm Phylogeny Website. The focus of this site is on angiosperms and emphasis is placed on plant families. You can also navigate from the Orders- or the Families-website. Go to:

R.A. Stockey et al. (2009): Introduction to the Darwin special issue: The abominable mystery. Free access, American Journal of Botany, 96: 3-4.

R.A. Stockey (1981): Some comments on the origin and evolution of conifers Canadian Journal of Botany, 59: 1932-1940. See also here.

Ruth A. Stockey, Department of Biological Sciences, University of Alberta, Edmonton: Paleobotany of Angiosperm Origins. Go to: Course Outline. Chiefly bibliographies and weblinks.

! P.K. Strother and C. Foster (2021): A fossil record of land plant origins from charophyte algae. Abstract, Science, 373: 792-796.
"... 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). ..."

P.K. Strother and C.H. Wellman (2021): The Nonesuch Formation Lagerstätte: a rare window into freshwater life one billion years ago. Open access, Journal of the Geological Society, 178.
"... Nonesuch microbiota, when viewed as a Lagerstätte, opens up a window onto the early evolution of unicellular eukaryotes, presenting an essential baseline of both eukaryotic diversity and cell structure well in advance of eukaryotic diversification documented in marine deposits from the later Neoproterozoic. ..."

P.K. Strother et al. (2021): A possible billion-year-old holozoan with differentiated multicellularity. Open access, Current Biology, 31: 2658-2665.e2

P.K. Strother et al. (2011): Earth´s earliest non-marine eukaryotes. In PDF, Nature 473: 505-509. See also here (abstract). See also the supplementary information (PDF, 5 MB).

Paul K. Strother, Palaeobotany Laboratory, Weston Observatory, Department of Geology & Geophysics, Boston College, Weston, Massachusetts: Links to Resources in Paleobotany, go to: Lectures, "Cryptospores and the Origin of Land Plants" (Powerpoint presentation). Attention, 132 MB!

! C. Strullu-Derrien et al. (2018): The origin and evolution of mycorrhizal symbioses: from palaeomycology to phylogenomics. Free access, New Phytologist, 220: 1012–1030.
! Note figure 1: Geological timescale with oldest known fossils. Left: Antiquity of genomic traits related to mycorrhizal evolution based on molecular clock estimates. Right: Oldest known fossils.
Figure 5: Simplified phylogenetic tree showing the minimum stratigraphic ranges of selected groups based on fossils (thick bars) and their minimum implied range extensions (thin lines).

C. Strullu-Derrien et al. (2016): Origins of the mycorrhizal symbioses. PDF file, In: F Martin (ed.): Molecular Mycorrhizal Symbiosis, John Wiley & Sons.

C. Strullu-Derrien (2014): The earliest wood and its hydraulic properties documented in c. 407-million-year-old fossils using synchrotron microtomography. Abstract, Botanical Journal of the Linnean Society, 175: 423-437.

D. Su et al. (2022): Large-scale phylogenomic analyses reveal the monophyly of bryophytes and neoproterozoic origin of land plants Open access, Molecular Biology and Evolution, 38: 3332–3344.
! Note figure 1: The concatenation species tree of land plants and their algal relatives.
! Figure 2: The coalescent species tree of land plants and their algal relatives.
"... We found that studies favoring a Neoproterozoic origin of land plants (980–682 Ma) are informed more by molecular data whereas those favoring a Phanerozoic origin (518–500 Ma) are informed more by fossil constraints. Our divergence time analyses highlighted the important contribution of the molecular data (time-dependent molecular change) when faced with contentious fossil evidence.
[..] A careful integration of fossil and molecular evidence will revolutionize our understanding of how land plants evolved.

D. Su et al. (2021): Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants. Free access, Molecular Biology and Evolution, 38: 3332–3344.
"... 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. ..."

T. Su et al. (2022): Tracing the Evolution of Plant Diversity in Southwestern China. Open access, Diversity, 14. See also here.

M. Sundaram et al. (2019): Accumulation over evolutionary time as a major cause of biodiversity hotspots in conifers. In PDF, Proc. R. Soc. B, 286: 20191887. See also here.

Ken Sytsma, Department of Botany , UW-Madison, Madison, WI:
Plant Geography. This course will examine the distributions of plants (and other organisms) from the perspectives of both ecology (biomes, climate, vegetation) and history (floristics, phylogenetics, paleobotany).
! Go to the PowerPoint Lectures (PDF files).
See especially:
Origin and Biogeography of Ancient Floras: Paleozoic (in black and white).
The same in color.
Origin and Biogeography of Ancient Floras: Mesozoic (in black and white).
The same in color.

M. Tanrattana et al. (2019): A new approach for modelling water transport in fossil plants. In PDF, IAWA Journal 40: 466–487.

D.W. Taylor and H. Li (2018): Paleobotany: Did flowering plants exist in the Jurassic period? eLife, 7: e43421.
"... we infer that Nanjinganthus shows substantial similarity to predicted models of ancestral characters and Early Cretaceous angiosperms, so the evidence suggests that it is a Jurassic flowering plant. ..."

! David W. Taylor and Leo J. Hickey (1996): Flowering Plant Origin, Evolution & Phylogeny. Google books (some pages omitted); American Institute of Biological Sciences (Springer), 404 pages.

! E.L. Taylor and T.N. Taylor (2009): Seed ferns from the late Paleozoic and Mesozoic: Any angiosperm ancestors lurking there? Open access, American Journal of Botany, 96: 237-251.
! "... In our opinion, it will be more productive to attempt to solve Darwin’s mystery if there were greater attention directed at mining the rock record in the hope of discovering more informative and new specimens, than to continue to construct new phylogenies using the same, often ambiguous characters. ..."
Worth checking out: Glossopterid vegetative and reproductive organs:
Note fig. 2: Suggested reconstruction of Ottokaria zeilleri.
Fig. 10: Suggested reconstruction of a Glossopteris megasporophyll with seeds attached to adaxial surface.
12: Diagrammatic reconstruction of Denkania indica.
Reproductive organs of Caytoniales and Corystospermales:
15. Suggested reconstruction of Caytonia cupule showing attachment of seeds and “stigmatic lip”.
16. Reconstruction of Caytonanthus arberi.
19. Suggested reconstruction of Umkomasia asiatica.
21. Diagrammatic reconstruction of Umkomasia uniramia.
Reproductive organs of Corystosperms and Petriellales:
25. Suggested reconstruction of Pilophorosperma geminatum.
28. Suggested reconstruction of Pteruchus fremouwensis.
30. Suggested reconstruction of Petriellaea triangulata.
32. Diagrammatic cutaway of Petriellaea triangulata cupule.
Reproductive organs of peltasperms:
34. Suggested reconstruction of Autunia conferta ovuliferous organ.
36. Suggested reconstruction of two Autunia conferta megasporophylls.
37. Suggested reconstruction of Peltaspermum rotula megasporophyll showing several ovules.
39. Suggested reconstruction of Peltaspermum thomasii axis bearing numerous megasporophylls.
40. Suggested reconstruction of Peltaspermopsis polyspermis.
41. Suggested reconstruction of Lepidopteris frond with pollen organs of the Antevsia-type at the tip.
42. Suggested reconstruction of Antevsia zeilleri pollen organ showing pinnate axis bearing clusters of pollen sacs.

! W. Testo and M. Sundue (2016): A 4000-species dataset provides new insight into the evolution of ferns. Abstract, Molecular Phylogenetics and Evolution, 105: 200–211. See also here (in PDF).

Greg Thorn, Department of Biology, University of Western Ontario, Canada:
! Evolution of Plants . Lecture notes, e.g.:
Evolution of the Angiosperms. Powerpoint presentation.

Greg Thorn, Department of Biology, University of Western Ontario: Evolution of Plants (Powerpoint presentations). Navigate from here with information from the Syllabus. See e.g. Lecture 16: Evolution of Plants. The evolution of early angiosperms.

A.S. Thorpe et al. (2011): Interactions among plants and evolution. In PDF, Journal of Ecology, 99: 729-740. Now provided by the Internet Archive´s Wayback Machine.

Bruce H. Tiffney (University of California, Santa Barbara), Access Excellance, BioForum 4, "Theoretical Issues in Plant Biology". BioForum is a series of lectures, presented by California Academy of Sciences, in which scientists share their research results with high school biology teachers. The Influence of Plants on the Evolution of Terrestrial Communities. "The Influence of Plants on the Evolution of Terrestrial Communities" is a tour de force through some 450 million years of plant evolution, giving you a feeling for the life forms based on sequential evidence of the fossil records and a consideration of the climates and major physical events that prevail.

! A.M.F. Tomescu (2021): Mysteries of the bryophyte–tracheophyte transition revealed: enter the eophytes. Free access, New Phytologist, Note fig. 1: Timeline and evolutionary hypothesis for early land plants. Worth checking out:
! D. Edwards et al. 2022a): Piecing together the eophytes–a new group of ancient plants containing cryptospores. Free access, New Phytologist, 233: 1440–1455.
! D. Edwards et al. 2022b): Earliest record of transfer cells in Lower Devonian plants. Free access, New Phytologist, 233: 1456–1465.

! A.M.F. Tomescu and G.W. Rothwell (2022): Fossils and Plant Evolution: Structural Fingerprints and Modularity in the Evo-Devo Paradigm. Free access, Evodevo, 13.
See also here.
Note fig. 8: The realization that a reproductive program can be activated in the intercalary meristem of individual equisetacean internodes.

! A.M.F. Tomescu et al. (2009): Carbon isotopes support the presence of extensive land floras pre-dating the origin of vascular plants. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 283: 46-59.

A.M.F. Tomescu (2016): Development: Paleobotany at the High Table of Evo-Devo. Free access, Current Biology, 26: R505-R508.

E. Trembath-Reichert et al. (2015): Four hundred million years of silica biomineralization in land plants. Free Access, Proc. National Academy of Sciences USA, 112: 5449–5454.

! C.J. van der Kooi and J. Ollerton (2020): The origins of flowering plants and pollinators. Free access, Science, 368: 1306-1308.
See also here (in PDF).

! Alain Vanderpoorten and Bernard Goffinet (2009): Introduction to Bryophytes, Evolutionary significance of bryophytes. In PDF, Cambridge University Press. See also here.

A. Vasco et al. (2016): Challenging the paradigms of leaf evolution: Class III HD-Zips in ferns and lycophytes. In PDF, New Phytologist, 212: 745–758. See also here.

! A. Vasco et al. (2013): The evolution, morphology, and development of fern leaves. In PDF, Frontiers in plant science. See also here (abstract).

! A. Vatén and D.C. Bergmann (2012): Mechanisms of stomatal development: an evolutionary view. In PDF, EvoDevo, 3.

B. Vento et al. (2023): Phylogenetic relationships in Nothofagus: The role of Antarctic fossil leaves. In PDF, Acta Palaeontologica Polonica, 68.

G.J. Vermeij (2017): How the land became the locus of major evolutionary innovations. In PDF, Current Biology; 27: 3178–3182. See also here.

G.J. Vermeij and L. Dudley (2000): Why are there so few evolutionary transitions between aquatic and terrestrial ecosystems? In PDF, Biological Journal of the Linnean Society, 70: 541-554.

! Elizabeth Anne Viau, Charter College of Education, California State University, Los Angeles: World Builders, Session Eight, Terrestrial Botany, Plants on Land. Go to: Important Landmarks in the Evolution of Land Plants.

M. Vicent et al. (2014): Insight into fern evolution: a mechanistic approach to main concepts and study techniques. In PDF, Botanica Complutensis, 38: 7-24. See also here.

R. Vidal-Russell and D.L. Nickrent (2008): The first mistletoes: Origins of aerial parasitism in Santalales. In PDF, Molecular Phylogenetics and Evolution, 47: 523-537.
Provided by the Internet Archive´s Wayback Machine.

A. Villarreal et al. (2015): Divergence times and the evolution of morphological complexity in an early land plant lineage (Marchantiopsida) with a slow molecular rate. Abstract, New Phytologist. See also here (in PDF).

School of Science and Engineering, University of Waikato, New Zealand: Evolution for Teaching. Earth's History and Evolution. Teaching resources. Go to: Plant Evolution, Prokaryotes, Algae and Plants.

! M.W. Wallace et al. (2017): Oxygenation history of the Neoproterozoic to early Phanerozoic and the rise of land plants. In PDF, Earth and Planetary Science Letters, 466: 12–19. See also here.

! S. Wallace et al. (2011): Evolutionary development of the plant spore and pollen wall. Open access, AoB PLANTS, 2011, plr027.

D.-M. Wang et al. (2015): Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China. Annals of Botany.

Q. Wang and K.-S. Mao (2015): Puzzling rocks and complicated clocks: how to optimize molecular dating approaches in historical phytogeography. In PDF, New Phytologist. See also here. (abstract).

X. Wang et al. (2018): Editorial: Evolution of Reproductive Organs in Land Plants. Open access, Front. Plant Sci., 9.

! X. Wang (2017): A Biased, Misleading Review on Early Angiosperms. In PDF, Natural Science, 9: 399-405.
Please note: P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

X.Q. Wang and J.H. Ran (2014): Evolution and biogeography of gymnosperms. In PDF, Molecular phylogenetics and evolution. See also here.

! L.M. Ward et al. (2016): Timescales of Oxygenation Following the Evolution of Oxygenic Photosynthesis. In PDF, Orig. Life Evol. Biosph.,46: 51-65.

! C.H. Wellmann et al. (2023). Terrestrialization in the Ordovician. Open access, Geological Society, London, Special Publications, 532.
"... This contribution reviews the evidence for terrestrial organisms during the Ordovician (microbial, land plant, fungal, animal)
[...] We conclude that the Ordovician was a critical period during the terrestrialization of planet Earth that witnessed the transition from a microbial terrestrial biota to one dominated by a vegetation of the most basal land plants. ..."

! C.H. Wellman (2014): The nature and evolutionary relationships of the earliest land plants. Abstract, New Phytologist, 202: 1–3. See also here (in PDF).

C.H. Wellman et al. (2003): Fragments of the earliest land plants. In PDF, Nature, 425: 282–285.
See also here.

G.D.A. Werner et al. (2014): A single evolutionary innovation drives the deep evolution of symbiotic N2-fixation in angiosperms. Open access, Nature Communications, 5: 4087.

Biology Department, Western Washington University, Bellingham, Washington: Plant Evolution. Powerpoint presentation. See also here, or there.

C.D. Whitewoods and E. Coen (2017): Growth and development of three-dimensional plant form. Abstract, Current Biology.

! N.J. Wickett et al. (2014): Phylotranscriptomic analysis of the origin and early diversification of land plants. In PDF, PNAS 111, see also here.

Niklas Wikström et al. (2001): Evolution of the angiosperms: calibrating the family tree. PDF file, Proc. R. Soc. Lond., B, 268: 2211-2220.

! Wikipedia, the free encyclopedia:
Plant evolution,
Evolutionary history of plants,
Timeline of plant evolution,
Plant evolutionary developmental biology.

Wikispaces, Tangient LLC, San Francisco, CA:
CDS Biology Website:
The Colonization of Land by Plants and Fungi. Powerpoint presentation.

N. Wikström et al. (2022): No phylogenomic support for a Cenozoic origin of the “living fossil” Isoetes. OPen access, American Journal of Botany.

! P. Wilf and I.H. Escapa (2015): Reply to Wang & Mao (2015): Molecular dates must be independently testable. In PDF.

! P. Wilf and I.H. Escapa (2015): Green Web or megabiased clock? Plant fossils from Gondwanan Patagonia speak on evolutionary radiations. In PDF, New Phytologist, 207: 283-290.

D.M. Wilkinson and T.N. Sherratt (2016): Why is the world green? The interactions of top-down and bottom-up processes in terrestrial vegetation ecology. In PDF, Plant Ecology & Diversity, 9: 127-140. See also here.

K.J. Willis and K.J. Niklas (2004): The role of Quaternary environmental change in plant macroevolution: the exception or the rule? In PDF, Phil. Trans. R. Soc. Lond., B 359: 159-172.

Kathy Willis and Jennifer McElwain (2014): The Evolution of Plants, Second Edition. Go to:
! Sample Material (in PDF). About the evolutionary record and methods of reconstruction.

Kathy Willis, School of Geography and the Environment, University of Oxford, & Jenny McElwain, Field Museum of Natural History, Chicago (Oxford University Press): The Evolution of Plants. Book announcement. Snapshot taken by the Internet Archive´s Wayback Machine. Go to: PowerPoint illustrations. Illustrations from the book in PowerPoint format. See also:
! Biome maps. Downloadable full-color images from the book.

Kathy Willis, School of Geography and the Environment, University of Oxford, & Jenny McElwain, Field Museum of Natural History, Chicago (Oxford University Press): The Evolution of Plants. Book announcement. Go to: Chapter 06, Flowering plant origins (PDF file).
Snapshots provided by the Internet Archive´s Wayback Machine.

R. Williams (2021): Discovered: Fossilized Spores Suggestive of Early Land Plants. The Scientist.

S. Williams (2017): The Weird Growth Strategy of Earth´s First Trees. The Scientist » News & Opinion » Daily News.
"Ancient fossils reveal how woodless trees got so big: by continuously ripping apart their xylem and knitting it back together".

Kathy Willis and Jennifer McElwain: The Evolution of Plants. Oxford University Press, Second Edition. Don't miss the
Companion Website
and some samples in Google books.
Note chapter 1: The evolutionary record and methods of reconstruction (in PDF).

! J.P. Wilson et al. (2023): Physiological selectivity and plant-environment feedbacks during Middle and Late Pennsylvanian plant community transitions. Free access, Geological Society, London, Special Publication, 535.

! J.P. Wilson et al. (2017): Dynamic Carboniferous tropical forests: new views of plant function and potential for physiological forcing of climate. In PDF, New Phytologist, 215: 1333–1353. See also here.
! Figure 2 shows the fungal evolution and abundance of coal basin sediments over the Phanerozoic.

! S.L. Wing and L.D. Boucher (1998): Ecological aspects of the Cretaceous flowering plant radiation. In PDF, Annu. Rev. Earth Planet. Sci. 1998 26: 379-421.

! S.L. Wing et al. (1992): Mesozoic and early Cenozoic terrestrial ecosystems. In PDF. In: Behrensmeyer, A.K., Damuth, J.D., DiMichele, W.A., Potts, R., Sues, H., Wing, S.L. (eds): Terrestrial Ecosystems Through Time : Evolutionary Paleoecology of Terrestrial Plants and Animals. University of Chicago Press, Chicago, pp.327–416.

! S. Woudenberg et al. (2022): Deep origin and gradual evolution of transporting tissues: Perspectives from across the land plants. In PDF, Plant Physiology.
See also here. Note figure 4: Summary of the early fossil record of transporting tissues.

! Q. Wu et al. (2021): High-precision U-Pb age constraints on the Permian floral turnovers, paleoclimate change, and tectonics of the North China block. Free access, Geology. See also here.
"... The great loss of highly diverse and abundant Cathaysian floras and the widespread invasion of the Angaran floras under arid climate conditions in the North China block happened during the late Cisuralian to Guadalupian, but its exact timing is uncertain due to the long hiatus. ..."

X. Zhang et al. (2017): How the ovules get enclosed in magnoliaceous carpels. PLoS ONE, 12: e0174955.

S. Xiao and Q. Tang (2018): After the boring billion and before the freezing millions: evolutionary patterns and innovations in the Tonian Period. In PDF, Emerging Topics in Life Sciences, 2: 161–171. See also here,

C. Xiong et al. (2013): Diversity Dynamics of Silurian-Early Carboniferous Land Plants in South China. PLoS ONE, 8.

H. Xu et al. (2022): The earliest vascular land plants from the Upper Ordovician of China. In PDF, ResearchSquare, DOI:
! Note fig. 4: Phylogeny and evolutionary timescale of early plant groups, with stratigraphic ranges of several key land-dwelling characters.

! H.-H. Xu et al. (2017): Unique growth strategy in the Earth´s first trees revealed in silicified fossil trunks from China. Abstract, Proceedings of the National Academy of Sciences of the United States of America, 114: 12009–12014. See also:
! D. Yuhas (2018): Ancient Tree Structure Is Like a Forest unto Itself. Arboreal fossils reveal an unusual and complex structure. Scientific American. See further:
Paläobotaniker lüften das Geheimnis der Urbäume (, in German).

S. Yashina et al. (2012): Regeneration of whole fertile plants from 30,000-y-old fruit tissue buried in Siberian permafrost. In PDF, PNAS, 109: 4008-4013.
See also here.
"... This natural cryopreservation of plant tissue over many thousands of years demonstrates a role for permafrost as a depository for an ancient gene pool, ..."
Also worth checking out: Scientists revive a 30,000 year old Pleistocene-era plant (by Gareth Branwyn, October 7, 2022).

! H.S. Yoon et al. (2004): A molecular timeline for the origin of photosynthetic eukaryotes. PDF file, Mol. Biol. Evol., 21: 809-818. See also here.

! A.E. Zanne et al. (2014): Three keys to the radiation of angiosperms into freezing environments. In PDF, Nature. Provided by the Internet Archive´s Wayback Machine.

N.E. Zavialova (2015): Evolutionary Transformations of Sporoderm Ultrastructure in Certain Monophyletic Lineages of Higher Plants. In PDF, Botanica Pacifica, 4.

! Z. Zhan et al. (2022): Origin and evolution of green plants in the light of key evolutionary events. Free access, Journal of Integrative Plant Biology, 64: 516–535.

L. Zhang et al. (2020): The water lily genome and the early evolution of flowering plants. Open access, Nature, 577: 79–84. Worth checking out:
Fig. 1d: Summary phylogeny and timescale of 115 plant species. Blue bars at nodes represent 95% credibility intervals of the estimated dates.

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This index is compiled and maintained by Klaus-Peter Kelber, Würzburg,
Last updated March 23, 2023

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