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Silurian and Devonian Palaeobotany

! University of Aberdeen: The Rhynie Chert Flora. See also The Biota of Early Terrestrial Ecosystems: The Rhynie Chert. A learning resource website., Forestry: Archaeopteris - The First Modern Tree.

T. Alekseeva et al. (2016): Characteristics of Early Earth's Critical Zone Based on Middle—Late Devonian Paleosol Properties (Voronezh High, Russia). In PDF, Clays and Clay Minerals, 64: 677–694.
Note also here.
Note figure 13: Terrane map of western and central Laurussia (including the Laurentian sector) and adjacent areas in the late Devonian (Famennian).

M.F. Alexandru et al. (2010): Simulating fossilization to resolve the taxonomic affinities of thalloid fossils in Early Silurian (ca. 425 Ma) terrestrial assemblages. In PDF.

J.P. Allen and R.A. Gastaldo (2006): Sedimentology and taphonomy of the Early to Middle Devonian plant-bearing beds of the Trout Valley Formation, Maine. PDF file, in: DiMichele, W.A., and Greb, S., eds., Wetlands Through Time: Geological Society of America, Special Publication 399: 57-78.
See also here.

! J.M. Anderson et al. (1999): Patterns of Gondwana plant colonisation and diversification. In PDF, Journal of African Earth Sciences, 28: 145-167.
See also here.

Joseph E. Armstrong, Department of Biological Sciences, Illinois State University, Normal: Plant diversity. Lecture notes. Go to: The Invasion of Land.
Snapshots provided by the Internet Archive´s Wayback Machine.

Lorna Ash, Department of Biological Sciences, University of Alberta: Instructional Multimedia, Multimedia Topics, Botany. Go to: Hydrasperman Reproduction. Online and downloadable flash movie.

! Stanley M. Awramik, Department of Earth Science, University of California Santa Barbara:
The Record of Life on the Early Earth. Lecture notes, Powerpoint presentation.

! Nicholas H. Barton (Edinburgh University), Derek E.G. Briggs (Yale University), Jonathan A. Eisen (University of California, Davis), David B. Goldstein (Duke University Medical Center), and Nipam H. Patel (University of California, Berkeley): Evolution (by Cold Spring Harbor Laboratory Press). This textbook is designed to serve as the primary text for undergraduate courses in evolution. It differs from currently available alternatives in containing more molecular biology than is traditionally the case. Go to: Table of Contents: Some figures and tables free of charge! See: Evolution Figures: Chapter 4.

! 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. Provided by the Internet Archive´s Wayback Machine.

! H. Beraldi-Campesi (2013): Early life on land and the first terrestrial ecosystems. In PDF, Ecological Processes, 2. See also here.

L. Battison and M.D. Brasier (2009): Exceptional Preservation of Early Terrestrial Communities in Lacustrine Phosphate One Billion Years Ago. Abstract.
Now provided by the Internet Archive´s Wayback Machine.

J. Bek and J.V. Frojdová (2023): Spore Evidence for the Origin of Isoetalean Lycopsids? Open access, Life, 13.
Note figure 3: Phylogeny of isoetalean lycopsids.
Compare with figure 4: New scheme.
"... Spores with three apical papillae, reported as dispersed as well as in situ, were recorded continuously from the lower Silurian (Wenlockian) through the Devonian, Carboniferous, Permian, Mesozoic to Cenozoic era and form a phylogenetically independent lineage ..."

Ernst-Georg Beck, Merian-Schule Freiburg: Biokurs (in German). Go to: Präkambrium: Hadäan (4,6 Milliarden Jahre - 3,8 Milliarden Jahre).

S. Bengtson et al. (2017): Three-dimensional preservation of cellular and subcellular structures suggests 1.6 billion-year-old crown-group red algae. Open Access, PLoS Biol., 15: e2000735.

A. Bennici (2008): Origin and early evolution of land plants: Problems and considerations. Free access, Commun. Integr. Biol., 1: 212-218.

! 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.

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): The evolution of the first forests in the Devonian. In PDF. See also here.
Note figure 1: Schematic timeline of known Devonian forest types.
Figure 2: Reconstruction of stand of Calamophyton (cladoxylopsid) trees (2—3 m high) based on fossils from Lindlar, Germany (Mid Eifelian age).
Figure 3: Reconstruction of forest showing upright cladoxylopsid trees (up to at least 8 m) and recumbent aneurophytaleans, from Gilboa, New York.
Figure 4: Reconstruction of forest of lycopsids with cormose bases and attached rootlets, from Munindalen, Svalbard.

C.M. Berry (2019): Palaeobotany: The Rise of the Earth’s Early Forests. Free access, Current Biology, 29: R792-R794.
Note figure 1: The ecology and appearance of known early forests.

C.M. Berry and J.E.A. Marshall (2015): Lycopsid forests in the early Late Devonian paleoequatorial zone of Svalbard. In PDF, Geology, 43: 1043-1046.

! Debashish Bhattachatya et al. (2009): Eukaryotes (Eukaryota). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

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.

! A.M.C. Bowles et al. (2023): The origin and early evolution of plants. Open access, Trends in Plant Science, 28.
Note figure 2: Phylogeny of early plant evolution with a selection of available genomic resources.
Figure 3: Fossils of possible and probable early archaeplastids.
! Figure 4: Summary of molecular estimates for the timescale of archaeplastid evolution.
"... Molecular clock analyses estimate that Streptophyta and Viridiplantae emerged in the late Mesoproterozoic to late Neoproterozoic, whereas Archaeplastida emerged in the late-mid Palaeoproterozoic ..."

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

! C.K. Boyce (2008): How green was Cooksonia? The importance of size in understanding the early evolution of physiology in the vascular plant lineage. In PDF, Paleobiology, 34: 179–194.
See likewise here.

C. Kevin Boyce et al. (2007): Devonian landscape heterogeneity recorded by a giant fungus. PDF file, Geology, 35: 399-402.
This expired link is available through the Internet Archive´s Wayback Machine.


MSc Palaeobiology Students, Department of Earth Sciences, University of Bristol, (the author´s name appears on the title page for each section):
Fossil Lagerstätten. A catalogue of sites of exceptional fossil preservation. Go to: The Flora of the Rhynie Chert.
Diagrammatic reconstructions of Rhynia, Aglaophyton, Horneophyton.
Some reconstruction images here.
Websites still available via Internet Archive Wayback Machine.

25th New Phytologist/Colston Research Society Symposium, September 21-22, 2010. Clifton Hill House, University of Bristol, UK: Colonization of the terrestrial environment. PDF file, Abstracts.

! D.R. Broussard et al. (2018): Depositional setting, taphonomy and geochronology of new fossil sites in the Catskill Formation (Upper Devonian) of north-central Pennsylvania, USA, including a new early tetrapod fossil. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 511: 168-187. See also here (in PDF).

! Mark C. Brundrett (2002): Coevolution of roots and mycorrhizas of land plants. PDF file, New Phytologist, 154: 275-304.
Now recovered from the Internet Archive´s Wayback Machine.

Stephen Caine, UK: Links to Models of the Rhynie Chert Plants. Viewed on the Rhynie Chert Flora page, the Aberdeen University Geology Department web site, etc. Ventarura lyonii, from the Windyfied chert, Rhynie, Scotland (showing enlarge section of possible sporangial arrangement). See also (page hosted by the Rhynie chert Research Group, the University of Aberdeen): The Royal Society's Summer Science Exhibition in London 2004 (a Rhynie diorama). Some images taken of the exhibit, including a visit by H.R.H. The Prince of Wales (who takes notice of palaeobotany elsewhere?)

Alison Campbell, Penelope Cooke, Kathrin Cass and Kerry Earl, The "Evolution for Teaching" Website Project, University of Waikato, New Zealand: The Evolution of Life. Information about the evolution of life on Earth. Go to: Plant Evolution.

E. Capel et al. (2023): New insights into Silurian–Devonian palaeophytogeography. Free access, Palaeogeography, Palaeoclimatology, Palaeoecology. 613.

E. Capel et al. (2023): The effect of geological biases on our perception of early land plant radiation. Free access, Palaeontology, 66.
"... geological incompleteness remains a fundamental bias for describing early plant diversification. This indicates that, even when sampling is extensive, observed diversity patterns potentially reflect the heterogeneity of the rock record, which blurs our understanding of the early history of land vegetation ..."

! 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. (2022): Revisiting the Rebreuve plant assemblage from the Lower Devonian of Artois, northern France. In PDF, Botany Letters, 2022, .10.1080/23818107.2022.2101516.. .hal-03781273.
See also here.

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.

C. Cardona-Correa et al. (2016): Peat Moss–Like Vegetative Remains from Ordovician Carbonates. Free access, International Journal of Plant Sciences, 177: 523-538.

Sean Carrington, Department of Biological & Chemical Sciences, University of the West Indies (UWI), Barbados: BIODIVERSITY I, THE PLANT KINGDOM. An introduction to the world of plants from an evolutionary perspective. Go to: The Conquest of the Land.

! B. Cascales-Miñana et al. (2019): On the hydraulic conductance of three woody Devonian plants. In PDF, IAWA journal, 40: 446-465. See also here (abstract).

! Eric J. Chaisson, Wright Center for Science Education: Cosmic evolution: from big bang to humankind. Based on a course taught at Harvard University. This site offers background information and resources to understand the origins of matter and life in our universe, known as cosmic evolution. Questions from how the universe began to how humans evolved are addressed, using an interdisciplinary approach between life, Earth, space, and physical sciences.
Website now publicly accessible by the Internet Archive´s Wayback Machine.
Go to: Chemical Evolution.

W.G. Chaloner (1968): The cone of Cyclostigma kiltorkense Haughton, from the Upper Devonian of Ireland. In PDF.

A. Channing (2018): A review of active hot-spring analogues of Rhynie: environments, habitats and ecosystems. In PDF, Transactions of the Royal Society, B, 373: 20160490.
See also here.
"... Comparison with Yellowstone suggests the Rhynie plants were preadapted to their environment by life in more common and widespread environments with elevated salinity and pH such as coastal marshes, salt lakes, estuaries and saline seeps. ..."

A. Channing and D. Edwards (2009): Yellowstone hot spring environments and the palaeoecophysiology of Rhynie chert plants: towards a synthesis. In PDF, Plant Ecology & Diversity. See also here.

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

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.

Samuel J. Ciurca, Jr., Rochester, New York: Silurian Plants (under construction). Cooksonia.
Recovered from the Internet Archive´s Wayback Machine.

Catherine Clabby, The News & Observer Publishing Company: Rocks may tell tale of first land plants. Palaeobotanical research at UNC-Chapel Hill.
This expired link is available through the Internet Archive´s Wayback Machine.

! C.J. Cleal and B.A. Thomas (The Geological Conservation Review): Palaeozoic Palaeobotany of Great Britain (PDF file). GCR VOLUME No. 9. Introduction. History of research on British plant fossils. List of sites (Silurian, Devonian, Lower Carboniferous, Upper Carboniferous, Permian). Scroll down to figure 1.1.

C.J. Cleal & B. A. Thomas: A Provisional World List of Geosites for Palaeozoic Palaeobotany. Initiated by the IUGS to develop an inventory of globally important geological sites. GEOSITES provide a provisional list of candidate Palaeozoic palaeobotany sites. The results are summarized in 40 sites, which are intended to show the broad pattern of evolution in land floras from the middle Silurian to the end of the Permian.
Still available via Internet Archive Wayback Machine.
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.
See also here.

S. Conway Morris (1993): The fossil record and the early evolution of the Metazoa. PDF file, Nature, 361: 219–225.
See also here.

L. Cornet et al. (2012): A Devonian Callixylon (Archaeopteridales) from Ronquières, Belgium. In PDF, Review of Palaeobotany and Palynology, 183: 1-8.
See also here.

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.

M. D'Ario et al. (2023): Hidden functional complexity in the flora of an early land ecosystem. Free access, New Phytologist, doi: 10.1111/nph.19228.
"... Our approach highlights the impact of sporangia morphology on spore dispersal and adaptation
We discovered previously unidentified innovations among early land plants, discussing how different species might have opted for different spore dispersal strategies ..."

Marilyn Davis, Perspectives, Southern Illinois University, Carbondale: LUSH LIFE: What early land plants can tell us about earth’s family tree.
Now provided by the Internet Archive´s Wayback Machine.

N.S. Davies and M.R. Gibling (2010): Cambrian to Devonian evolution of alluvial systems: The sedimentological impact of the earliest land plants. Abstract, Earth-Science Reviews, 98: 171-200.

Anne-Laure Decombeix, Brigitte Meyer-Berthaud, Nick Rowe & Jean Galtier: Diversity of large woody lignophytes preceding the extinction of Archaeopteris: new data from the middle Tournaisian of Thuringia (Germany).

G.P. de Oliveira Martins et al. (2018): Are early plants significant as paleogeographic indicators of past coastlines? Insights from the taphonomy and sedimentology of a Devonian taphoflora of Paraná Basin, Brazil. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 505: 234-242. See also here.

Melanie DeVore, Georgia College and State University, Milledgeville, GA:
! The Evolution of Plants. Powerpoint presentation, slow download, 90 MB!
Still available via Internet Archive Wayback Machine.
Provided by D. Freile, New Jersey City University: Historical Geology.
Powerpoint presentations, still available via Internet Archive Wayback Machine.

! 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 surrounded by synoecosporal walls has led to algal interpretations ..."
! Note figure 4: Timescale of streptophyte phylogeny and the origin of land plant novelties.

! N.L. Dotzler (2009): Microbial life in the late Paleozoic: new discoveries from the Early Devonian and Carboniferous. In PDF, Thesis, Ludwig-Maximilians-Universität München.

E. Dowding et al. (2023): Survivorship dynamics of the flora of Devonian Angarida. Proceedings of the Royal Society of London, B, Biological Sciences, 290.
"... survivorship dynamics of early plant communities upon the palaeocontinent Angarida have demonstrated that transgression and volcanogenic nutrient influx were key to the survival of colonizing plants ..."

A. Duhin et al. (2022): Early land plants: Plentiful but neglected nutritional resources for herbivores? Open access, Ecology and Evolution, 12.

J.A. Dunlop and R.J. Garwood (2017): Terrestrial invertebrates in the Rhynie chert ecosystem. In PDF, Phil. Trans. R. Soc. B, 373: 20160493.

R.L. Dunstone and G.C. Young (2019): New Devonian plant fossil occurrences on the New South Wales South Coast: geological implications. In PDF, Australian Journal of Earth Sciences, 66. See also here.

! 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 et al. (2017): History and contemporary significance of the Rhynie cherts—our earliest preserved terrestrial ecosystem. Phil. Trans. R. Soc., B 373: 20160489. See also here (in PDF).
Note figure 1 and 2: Kidston and Lang’s original reconstructions of Rhynie gwynnevaughanii, Aglaophyton majus (Rhynia major), Asteroxylon mackiei and Horneophyton lignieri (Hornea lignieri).

! 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.

D. Edwards (2004): Embryophytic sporophytes in the Rhynie and Windyfield cherts. Abstract, Transactions of the Royal Society of Edinburgh: Earth Sciences, 94: 397–410. See also here (in PDF).

D. Edwards et al. (2002): Hepatic characters in the earliest land plants. Abstract, Nature, 374: 635-636.

D. Edwards (2000): The role of Mid-Palaeozoic mesofossils in the detection of early bryophytes. In PDF, Philosophical Transactions of the Royal Society B.

! D. Edwards, H. Kerp and H. Hass (1998): Stomata in early land plants: an anatomical and ecophysiological approach. Journal of Experimental Botany, Vol. 49, Special Issue, pp. 255–278.

! Mike Farabee, Estrella Mountain Community College Center, Avondale, Arizona: On-Line Biology Book. Introductory biology lecture notes. Go to: PALEOBIOLOGY: FOSSILS AND TIME, PALEOBIOLOGY: THE PRECAMBRIAN: LIFE'S GENESIS AND SPREAD, PALEOBIOLOGY: THE EARLY PALEOZOIC, and PALEOBIOLOGY: THE LATE PALEOZOIC.

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

Ben Fletcher, Department of Animal and Plant Sciences, University of Sheffield: The role of stomata in the early evolution of land plants, and How the atmosphere affects plants. See also: Modern-day representatives of early land plants.

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.

J.R. Flores et al. (2023): Dating the evolution of the complex thalloid liverworts (Marchantiopsida): total-evidence dating analysis supports a Late Silurian-Early Devonian origin and post-Mesozoic morphological stasis. In PDF, New Phytologist, doi: 10.1111/nph.19254.
See also here.
"... Phylogenetic analyses were performed on a combined dataset of 130 discrete characters and 11 molecular markers
[...] Total-evidence dating analyses support the radiation of Marchantiopsida during Late Silurian-Early Devonian (or Middle Ordovician when the outgroup is excluded) and that of Ricciaceae in the Middle Jurassic ..."

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.

X. Gao et al. (2022): Re-study of Guangdedendron micrum from the Late Devonian Xinhang forest. Free access, BMC Ecology and Evolution, 22.
Note the reconstruction in fig. 6.

! 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 et al. (2020): Back to the Beginnings: The Silurian-Devonian as a Time of Major Innovation in Plants and Their Communities PDF file, pp 367–398. In: Nature through Time: Virtual field trips through the Nature of the past. Springer, Textbooks in Earth Sciences, Geography and Environment. (eds Martinetto E., Tschopp E., Gastaldo R.A.), pp. 159–185. Springer International Publishing, Cham.
See likewise here.
! Note figure 15.20: Phylogenetic relationships between the major Paleozoic plant groups.

! 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. (2022): Earliest Evidence of Land Plants in Brazil. In PDF, In: Iannuzzi, R., Rößler, R., Kunzmann, L. (eds.): Brazilian Paleofloras. Springer.
See also here.
Note. fig. 3: Suggested life cycle of an early vascular plant from the early Devonian Rhynie Chert.
Fig. 4b: Suggested reconstruction of Cooksonia paranensis.
Fig. 5: Suggested life cycle of Cooksonia paranensis.

P. Gerrienne et al. (2019): Earliest Evidence of Land Plants in Brazil. Abstract, Brazilian Paleofloras. See also here (in PDF).

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.

R.W. Gess and C. Prestianni (2021): An early Devonian flora from the Baviaanskloof Formation (Table Mountain Group) of South Africa Open access, Scientific Reports, 11. See also:
Discovery of the oldest plant fossils on the African continent! EurekAlert, the American Association for the Advancement of Science (AAAS).

P. Giesen and C.M. Berry (2013): Reconstruction and growth of the early tree Calamophyton (Pseudosporochnales, Cladoxylopsida) based on exceptionally complete specimens from Lindlar, Germany (mid-Devonian): organic connection of Calamophyton branches and Duisbergia trunks. PDF file, Int. J. Plant Sci., 174: 665-686.

! I.J. Glasspool and R.A. Gastaldo (2022): Silurian wildfire proxies and atmospheric oxygen, Open avccess, Geology.
! Note figure 3: Silurian–Devonian charcoal plotted against three common models of Paleozoic pO2 and back-calculated measurements.
"... The frequency of charcoal data from Silurian sequences indicates that fires were not rare but an established part of the terrestrial biome from at least the Wenlock onward. ..." Also worth checking out:
International Spotlight Shines on Colby Geologists (by Bob Keyes, July 7, 2022, Colby News).

R. Gossmann et al. (2022): A stratigraphically significant new zosterophyllopsid from the Rhenish Lower Devonian (W Germany). Free access, Palaeobiodiversity and Palaeoenvironments, 102: 503–519.

! L.E. Graham et al. (2010): Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats. In PDF, American Journal of Botany, 97: 268-275. See also:
! T.N. Taylor et al. (2010): The enigmatic Devonian fossil Prototaxites is not a rolled-up liverwort mat: Comment on the paper by Graham et al.(AJB 97: 268-275). In PDF. See also:
! L.E. Graham et al. (2010): Rolled liverwort mats explain major Prototaxites features: Response to commentaries.

L.E. Graham et al. (2004): Resistant tissues of modern marchantioid liverworts resemble enigmatic Early Paleozoic microfossils. In PDF, PNAS, 101: 11025-11029.

! 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. 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.
Note figure 1: Evolution of wetland types in the Silurian and Devonian.
See also here.
Still available through the Internet Archive´s Wayback Machine.

J.D. Grierson and H.P. Banks (1983): A new genus of lycopods from the Devonian of New York State. In PDF, Botanical Journal of the Linnean Society, 86: 81-101. See also here.
Note figure 22: Diagrams illustrating fracture planes or weathering surfaces of compressed lycopod stems in a rock matrix.

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.

S. Hartenfels et al. (2022): The Rhenish Massif: More than 150 years of research in a Variscan mountain chain. Open access, Palaeobiodiversity and Palaeoenvironments, 102: 493–502.
Note figure 7: Reconstruction of the Mid-Devonian Lindlar forest.

Heckman, D.S., et al. 2001: Molecular evidence for the early colonization of land by fungi and plants. Science 293: 1129-1133.

Blair Hedges and Barbara K. Kennedy (Penn State), EurekAlert: First land plants and fungi changed earth's climate, paving the way for explosive evolution of land animals, new gene study suggests.

A.J. Hetherington et al. (2021): An evidence-based 3D reconstruction of Asteroxylon mackiei the most complex plant preserved from the Rhynie chert. Free access, eLife. See also here, and there. Worth checking out:
Zu den Wurzeln der pflanzlichen Evolution (, in German).
Forscher rekonstruieren, wie die ersten Wurzeln Fuß fassten (Der Standard, in German).

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).

L.A. Hoffman and A.M.F. Tomescu (2013): An early origin of secondary growth: Franhueberia gerriennei gen. et sp. nov. from the Lower Devonian of Gaspé (Quebec, Canada). OPen access, American Journal of Botany, 100: 754-763.

Patrick Honecker, University of Cologne: Ancestors of land plants revealed.
Still available through the Internet Archive´s Wayback Machine. Education 2001, Leaving the Water.

Ann Jelinek et al., Murrindindi Shire Council, Australia: Flora Fossil Site: YEA (PDF file). About Baragwanathia. See also here.

Uwe Kaulfuß, Technische Universität, Bergakademie Freiberg, Germany: Geologisches Oberseminar 2000/2001, Charakterisierung der Eroberung des Festlandes als Habitat. In German (PDF-file).

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: Chlorobionta: Land Plants.
Evolution of Land Plants.

! 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 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.

P. Kenrick (2001): Turning over a new leaf. PDF file, Nature, 410: 309-310. This expired link is available through the Internet Archive´s Wayback Machine.

Paul Kenrick, Department of Palaeontology, The Natural History Museum, London, UK: Palaeobotany: Fishing for the first plants. Nature 425, 248 - 249.

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

! P. Kenrick & P.R. Crane (1997): The origin and early evolution of plants on land. PDF file, Nature.
See also here.

H. Kerp and M. Krings (2023): The Early Devonian Rhynie chert–The world's oldest and most complete terrestrial ecosystem. PDF file, starting on PDF page 45. In: J. Reitner, M. Reich, J.-P. Duda (eds.): Abstracts, Fossillagerstätten and Taphonomy.


! H. Kerp (2017): Organs and tissues of Rhynie chert plants. Open access, Phil. Trans. R. Soc. B, 373: 20160495.

H. Kerp et al. (2013): Reproductive organs and in situ spores of Asteroxylon mackiei Kidston & Lang, the most complex plant from the lower Devonian Rhynie chert. In PDF, Int. J. Plant Sci., 174: 293-308.

! Hans Kerp, Palaeobotanical Research Group, Westfälische Wilhelms University, Münster. Click: "Rhynie Chert" (The Rhynie Chert and its Flora). A depiction of the silica permineralized fossil flora of Rhynie (Scotland), a 400 Million year old flora, which contains a wide diversity of taxa varying from unicellular fungi to the earliest anatomically preserved higher land plants and animal remains. Breathtaking thin section micro-photographs, e.g. in " V. The alternation of generations in early land plants": The male gametophyte with antheridia, the release of sperm from antheridium, etc. Including "The life cycle of Aglaophyton - Lyonophyton", modified after Taylor, Kerp & Hass, 2005, PNAS, v. 102, p. 5892-5897.

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.

Khudadad (2021): A Middle Devonian vernal pool ecosystem provides a snapshot of the earliest forests. Open access, PLoS ONE 16(9): e0255565.

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

M.E. Kotyk et al. 2002): Morphologically complex plant macrofossils from the Late Silurian of Arctic Canada. American Journal of Botany 89(6): 1004–1013. See also here.

! M. Krings and H. Kerp (2023): The fidelity of microbial preservation in the Lower Devonian Rhynie cherts of Scotland. PDF file, starting on PDF page 54. In: J. Reitner, M. Reich, J.-P. Duda (eds.): Abstracts, Symposium Fossillagerstätten and Taphonomy.

! M. Krings et al. (2017): Fungi and fungal interactions in the Rhynie chert: a review of the evidence, with the description of Perexiflasca tayloriana gen. et sp. nov. Free access, Phil. Trans. R. Soc. B, 373. See also here.

! M. Krings, LMU München: Mikroorganismen aus den Cherts von Esnost und Combres/Lay (Unterkarbon, Frankreich) sowie Rhynie (Unterdevon, Schottland). Scientific project report (in German).
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

Khudadad (2021): A Middle Devonian vernal pool ecosystem provides a snapshot of the earliest forests. Open access, PLoS ONE 16(9): e0255565.
Note figure 14: Representative fossils of roots systems belonging to three Middle Devonian tree clades.

U. Kutschera, Institut für Biologie, Universität Kassel, Germany, and K.J. Niklas, Department of Plant Biology, Cornell University, Ithaca, NY: The modern theory of biological evolution: an expanded synthesis. Worth to check out: Fig.2 Geological time scale with key events in the history of life, from the formation of the Earth to the present.

C.C. Labandeira et al. (2014): Middle Devonian liverwort herbivory and antiherbivore defence. In PDF, New Phytologist, 202: 247–258. 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. See also here.

M.A.K. Lalica and A.M.F. Tomescu (2023): Complex wound response mechanisms and phellogen evolution–insights from Early Devonian euphyllophytes. Abstract, New Phytologist, 239: 388-398.
"... The earliest occurrences of wound periderm pre-date the oldest known periderm produced systemically as a regular ontogenetic stage (canonical periderm), suggesting that periderm evolved initially as a wound-response mechanism. We hypothesize that canonical periderm evolved by exaptation of this wound sealing mechanism..."

! M.A.K. Lalica and A.M.F. Tomescu (2021): The early fossil record of glomeromycete fungi: New data on spores associated with early tracheophytes in the Lower Devonian (Emsian; c. 400 Ma) of Gaspé (Quebec, Canada). In PDF, Review of Palaeobotany and Palynology. See also here.
"... occurrence in fluvial-coastal environments and their putative mycorrhizal role suggest that glomeromycetes were relatively ubiquitous symbionts of tracheophytes, ..."

G. Le Hir et al. (2011): The climate change caused by the land plant invasion in the Devonian. In PDF, Earth and Planetary Science Letters, 310: 203-212.

! F. Leliaert et al. (2012): Phylogeny and Molecular Evolution of the Green Algae. PDF file, Critical Reviews in Plant Sciences, 31: 1-46.
See also here.

! F. Leliaert et al. (2011): Into the deep: new discoveries at the base of the green plant phylogeny. PDF file, BioEssays. 33: 683-692. See also here.
! Note figure 1: Phylogenetic relationships among the main lineages of green plants.
"... A schism early in their evolution gave rise to two major lineages, one of which diversified in the world’s oceans and gave rise to a large diversity of marine and freshwater green algae (Chlorophyta) while the other gave rise to a diverse array of freshwater green algae and the land plants (Streptophyta) ..."

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

! M. Libertín et al. (2022): The early land plant Cooksonia bohemica from the Pridoli, late Silurian, Barrandian area, the Czech Republic, Central Europe. In PDF, Historical Biology, DOI: 10.1080/08912963.2022.2144286.
See also here.
! Note figure 7: Reconstruction of Cooksonia bohemica.
! Figure 8: Reconstruction of Aberlemnia caledonica.

M. Libertín et al. (2018): Sporophytes of polysporangiate land plants from the early Silurian period may have been photosynthetically autonomous. Abstract, Nature Plants, 4: 269–271. See also here

L. Liu et al. (2022): A Late Devonian tree lycopsid with large strobili and isotomous roots. Open access, Communications Biology, 5.
Note figure 7: Reconstruction of a strobilus of Omprelostrobus gigas.

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

C.C. Loron et al. (2023): Molecular fingerprints resolve affinities of Rhynie chert organic fossils. Open access, Nature Communications, 4.
"... we demonstrate that the famously exquisite preservation of cells, tissues and organisms in the Rhynie chert accompanies similarly impressive preservation of molecular information. These results provide a compelling positive control that validates the use of infrared spectroscopy to investigate the affinity of organic fossils in chert. ..."

M. Lu et al. (2019): Geochemical Evidence of First Forestation in the Southernmost Euramerica from Upper Devonian (Famennian) Black Shales. Free access, Scientific Reports, 9.
"... Plant residues (microfossils, vitrinite and inertinite) and biomarkers derived from terrestrial plants and wildfire occur throughout the stratigraphic section, suggesting widespread forest in the southern Appalachian Basin, a region with no macro plant fossil record during the Famennian. Inorganic geochemical results, as shown by increasing values of SiO2/ Al2O3, Ti/Al, Zr/Al, and the Chemical Index of Alteration (CIA) upon time sequence, suggest enhanced continental weathering that may be attributed to the invasion of barren lands by rooted land plants. ..."

P. Maffre et al. (2022): The complex response of continental silicate rock weathering to the colonization of the continents by vascular plants in the Devonian. In PDF,
See also here.
"... The fossil record shows that, by the end of the Devonian, vascular plants and forests were common and widespread [...]
we build a mathematical description of the coupled response of the physical erosion and chemical weathering on the continents, to the colonization by vascular plants over the course of the Devonian.

Gustavo Prado de Oliveira Martins et al. (2018): Are early plants significant as paleogeographic indicators of past coastlines? Insights from the taphonomy and sedimentology of a Devonian taphoflora of Paraná Basin, Brazil. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 505: 234–242. See also here.

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.

K.K.S. Matsunaga and A.M.F. Tomescu (2017): An organismal concept for Sengelia radicans gen. et sp. nov. – morphology and natural history of an Early Devonian lycophyte. Free access, Annals of Botany, 19: 1097–1113.
Note figure 10: Whole-plant reconstruction of Sengelia radicans.
See also:
Whole-plant reconstruction of an early Devonian lycophyte (Botany One, 2017).

! K.K.S. Matsunaga and A.M.F. Tomescu (2016): Root evolution at the base of the lycophyte clade: insights from an Early Devonian lycophyte. Free access, Annals of botany, 117: 585–598.

! R.M. McCourt et al. (2023): Green land: Multiple perspectives on green algal evolution and the earliest land plants. In PDF, American Journal of Botany 110. See also here (Free to read).
! Note figure 1: Green plant diversification in the context of the fossil record.
"... Green plants, broadly defined as green algae and the land plants (together, Viridiplantae), constitute the primary eukaryotic lineage that successfully colonized Earth's emergent landscape.
[...] We present the process not as a step-by-step advancement from primitive green cells to an inevitable success of embryophytes, but rather as a process of adaptations and exaptations that allowed multiple clades of green plants ..."

Richard M. McCourt et al. (2004): Charophyte algae and land plant origins. PDF file, Trends in Ecology and Evolution, 19.

G. McLean (2017): A 'mystery fossil' is evidence for massive Devonian trees in Australia. In PDF, Records of the Australian Museum, 69: 101–118. See also here.
Note figure 10: A possible taphonomic process experienced by the Griffith specimen.

F.R. McSweeney et al. (2022): Lower Devonian Zosterophyllum-like plants from central Victoria, Australia, and their significance. In PDF, Memoirs of Museum Victoria, 81: 25–41.

M.-C. Meng et al. (2015): Vegetative Characters, Growth Habit and Microsporangiate Strobilus of Lycopsid Minostrobus chaohuensis. PLoS ONE, 10.

B. Meyer-Berthaud et al. (2016): The terrestrialization process: a palaeobotanical and palynological perspective. In PDF, Review of Palaeobotany and Palynology, 224: 1–3.

B. Meyer-Berthaud and A.L. Decombeix (2012): In the shade of the oldest forest. In PDF, Nature, 483.

Brigitte Meyer-Berthaud and Anne-Laure Decombeix (2007): Palaeobotany: A tree without leaves.

! 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.

R.L. Mitchell et al. (2021): Cryptogamic ground covers as analogues for early terrestrial biospheres: Initiation and evolution of biologically mediated proto-soils. Open access, Geobiology, 19: 292-306.
Note fig. 8: Illustrations summarising the key features in modern lichen, thalloid plant, moss and mixed proto-soils.

! 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.

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany: History of Palaeozoic Forests, SILURIAN PLANT FOSSILS. Link list page with rankings and brief explanations. Images of Silurian cryptospores and Parka decipiens. See also:
THE EARLIEST LIFE. Link list page with picture rankings. Images of precambrian microfossils and stromatolites. The links give the most direct connections to pictures available on the web; in many cases they are from sites that have additional palaeobotanical information. See also:
THE EARLIEST LAND PLANTS. Link list page with rankings and brief explanations. Images of Rhynia, Rhynia gwynne-vaughanii, Cooksonia, Cooksonia hemisphaerica, Baragwanathia, Cooksonia pertonii, Aglaophyton major, Lyonophyton rhynienensis, Horneophyton lignieri, Nothia aphylla, Crenaticaulis, Sawdonia, Sawdonia acanthotheca, Sawdonia ornata, Serrulacaulis furcatus, Rebuchia ovata, Zosterophyllum divaricatum, Zosterophyllum rhenanum, Psilophyton, Psilophyton crenulatum, Psilophyton dawsonii, Psilophyton dapsile, Psilophyton ornata, Pertica, Pertica quadrifaria, Asteroxylon, Asteroxylon mackiei. See also:
THE EARLY FORESTS AND THE PROGYMNOSPERMS. Images of Archaeopteris, Tetraxylopteris schmidtii, Callixylon, Archaeopteris gaspensis, Archaeopteris halliana, Archaeopteris hibernica. See also:
EARLIEST SEED PLANTS. Images of Moresnetia, Moresnetia zaleskyi, Elkinsia. Excellent!
These expired links are now available through the Internet Archive´s Wayback Machine.

Dennis C. Murphy: Devonian Times. Go to: Who's Who at Red Hill. A survey of Devonian plants (tracheophytes)and animals. Photographs and drawings of Barinophyton, Lepidodendropsis, Archaeopteris, Gillespiea randolphensis, Rhyacophyton ceratangium.

M.P. Nelsen and C.K. Boyce (2022): What to Do with Prototaxites? Abstract, International Journal of Plant Sciences.

D.L. Nickrent et al. (2000): Multigene phylogeny of land plants with special reference to bryophytes and the earliest land plants. PDF file, Molecular Biology and Evolution: 17: 1885-1895.

! K.J. Niklas (2023): Deciphering the hidden complexity of early land plant reproduction. Free 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.

! E.G. Nisbet and N.H. Sleep (2001): The habitat and nature of early life. PDF file, Nature, 409.
The link is to a version archived by the Internet Archive´s Wayback Machine.

Malcolm O'Neill and William York, Plant Cell Wall Research, The Complex Carbohydrate Research Center, The University of Georgia Athens: The Cell Walls of Lower Plants. Project announcement.

C.P. Osborne et al.(2004): Biophysical constraints on the origin of leaves inferred from the fossil record. PDF file, PNAS, 101: 10360-10362.
This expired link is available through the Internet Archive´s Wayback Machine.

£. 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.

Sid Perkins, Science now: ScienceShot: Ancient Forest Kept Good Company. Fossil tree stumps in a sandstone quarry near Gilboa, New York.

John Perlin, Eco-Links: The Tree That Changed the World. PDF file, very slow!

! J.M. Pettitt and C.B. Beck (1968): Archaeosperma arnoldii: a cupulate seed from the Upper Devonian of North America. In PDF, Contrib. Mus. Paleontol. Univ. Mich., 22: 139–154.

K.C. Pfeiler et al. (2018): An Early Devonian permineralized rhyniopsid from the Battery Point Formation of Gaspé (Canada). In PDF, Botanical Journal of the Linnean Society, 187: 292–302. See also here.

K.C. Pfeiler and A.M.F. Tomescu (2018): An Early Devonian permineralized rhyniopsid from the Battery Point Formation of Gaspé (Canada). Free access, Botanical Journal of the Linnean Society, 187: 292–302.

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.

! 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.

B.R. Pratt and J. van Heerde (2017): An arborescent lycopsid stem fragment from the Palliser Formation (Famennian) carbonate platform, southwestern Alberta, Canada, and its paleogeographic and paleoclimatic significance. In PDF, Canadian Journal of Earth Sciences, 54: 141-145. See also here (abstract).

C. Prestianni et al. (2013): Were all devonian seeds cupulate? A reinvestigation of Pseudosporogonites hallei, Xenotheca bertrandii, and Aglosperma spp. In PDF, Int. J. Plant Sci. 174, 832–851.

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.

W. Qie et al. (2023): Enhanced Continental Weathering as a Trigger for the End-Devonian Hangenberg Crisis. Open access, Geophysical Research Letters, 50: e2022GL102640.
Note figure 1A: Latest Devonian global paleogeographic reconstruction.
"... The colonization of land plants during the Devonian is believed to have played a key role in regulating Earth's climate. The initially rapid expansion of seed plants into unvegetated or sparsely vegetated uplands is considered to have caused enhanced rock dissolution relative to clay formation on end-Devonian continents ..."

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

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

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.

K.S. Renzaglia et al. (2017): Hornwort stomata: architecture and fate shared with 400 million year old fossil plants without leaves. Free access, Plant Physiology, 177: 788–797.

! 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.

G.J. Retallack and C. Huang (2021): Ecology and evolution of Devonian trees in New York, USA. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 299: 110-128. See also here.

G.J. Retallack (2021): Great moments in plant evolution. See also here (in PDF).
Please notice figure 1.

G.J. Retallack (2015): Silurian vegetation stature and density inferred from fossil soils and plants in Pennsylvania, USA. In PDF, Journal of the Geological Society.
Reconstructed Siluro-Devonian plants on PDF page 14.
See also here (abstract).

G.J. Retallack and C. Huang (2011): Ecology and evolution of Devonian trees in New York, USA. In PDF, Palaeogeography, Palaeoclimatology, Palaeoecology, 299: 110-128. 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).

Sue Rigby, Geology, Geophysics, Environmental Geoscience, Grant Institute, University of Edinburgh: COURSE MATERIALS. Go to: GEP COURSE MATERIALS,
Lecture 3: The early earth and the origin of life. PDF file.

S.M. Rimmer et al. (2015): The rise of fire: Fossil charcoal in late Devonian marine shales as an indicator of expanding terrestrial ecosystems, fire, and atmospheric change. In PDF, American Journal of Science, 315: 713-733.

Paul Rincon, BBC News Online: Fossils reveal oldest wildfire.

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.

! 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.

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

A. Salt (2018): Plants and Fungi: An ancient partnership. Botany One.

J.D. Schiffbauer et al. (2012): Thermally-induced structural and chemical alteration of organic-walled microfossils: an experimental approach to understanding fossil preservation in metasediments. In PDF, Geobiology, 10: 402-423.
This expired link is now available through the Internet Archive´s Wayback Machine.
See also here.

J.W. Schopf (1999), article starts on PDF page 105: Fossils and Pseudofossils: Lessons from the Hunt for Early Life on Earth. In PDF; In: Proceedings of the Workshop on Size Limits of Very Small Organisms, Space Studies Board, National Research Council, National Academies Press, Washington, DC. See also here. Geology, Evolution upset: Oxygen-making microbes came last, not first.

! A.C. Scott (1984): The early history of life on land. In PDF, Journal of Biological Education, 18. See also here.
Note figs. 5 and 6: Rconstructions of Silurian and Devonian plants.

! P.A. Selden (2016): Land Animals, Origins of. In PDF. In: Kliman, R. M. (ed.): Encyclopedia of evolutionary biology. Volume 2: 288-295. Oxford, Academic Press.
About the colonization of the land habitat from the sea by plants and animals.

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.

T. Servais et al. (2023): No (Cambrian) explosion and no (Ordovician) event: A single long-term radiation in the early Palaeozoic. Free access, Palaeogeography, Palaeoclimatology, Palaeoecology, 623.
Note figure 1: Overview of the major terminologies used in studies of early Palaeozoic (Cambrian, Ordovician, Silurian) biodiversity, including the Cambrian ‘Explosion’ and the Great Ordovician Biodiversification ‘Event,’ and stratigraphical position of some of the most significant Konservat-Lagerstätten.
Figure 3: Early Palaeozoic species richness curves of the three major biostratigraphical groups.

! 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.

! M.S. Smart et al. (2022): Enhanced terrestrial nutrient release during the Devonian emergence and expansion of forests: Evidence from lacustrine phosphorus and geochemical records. Free access, GSA Bulletin. Note also:
Können Wurzeln töten? By P. Heinemann, Frankfurter Allgemeine, March 07, 2023 (in German).

Society of American Foresters, Bethesda, Maryland: A Tree That Changed the World. About Archaeopteris, from the June 1999 issue of "The Forestry Source". See also (by NewsWise): Earliest Modern Tree Lived 360-345 Million Years Ago.

P. Steemans et al. (2023): A diverse Early Devonian palynoflora from the Waxweiler Lagerstätte (Klerf Formation, Rhenish Massif, Western Germany): palaeobotanical implications. In PDF, Palynology, 47.
See also here.

! P. Steemans et al. (2009): Origin and Radiation of the Earliest Vascular Land Plants. In PDF, Science, 324.

P. Steemans et al. (2007): Palaeophytogeographical and palaeoecological implications of a miospore assemblage of earliest Devonian (Lochkovian) age from Saudi Arabia. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology, 250: 237-254.
See also here.

! P. Steemans and E. Javaux (Editors), (Carnets de Géologie / Notebooks on Geology: Memoir 2005/02): Pre-Cambrian to Palaeozoic Palaeopalynology and Palaeobotany. Online articles from a meeting, organized by the NFSR Working Group: "Micropaléontologie végétale et Palynologie (MVP)" and supported by the NFSR, the University of Liège, and the French Community of Belgium (May 11, 2005). Excellent!

! W.E. Stein et al. (2019): Mid-Devonian Archaeopteris Roots Signal Revolutionary Change in Earliest Fossil Forests. Free access, Current Biology, See also here (in PDF).
Worth checking out:
Scientists have discovered the world’s oldest forest—and its radical impact on life (by Colin Barras, Science Magazine,

W.E. Stein et al. (2012): Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa. Abstract, Nature, 483. Numerous Eospermatopteris root systems in life position within a mixed-age stand of trees, large woody rhizomes with adventitious roots.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Fossil plant images from the oldest land plants. Go to: The evolution of plants. An introduction. See also:
The oldest land plants (1),
The oldest land plants (2).
Also worth to visit: The comprehensive site about
Cooksonia,a very old land plant: page 1,
and page 2.

P.K. Strother (2023): An evo-devo perspective on no Ordovician land plants. In PDF, Estonian Journal of Earth Sciences,
Note figure 1: Fossil record of microfossils related to land plant origins seen as character distributions.

! 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. (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).

P.K. Strother (2010): Thalloid carbonaceous incrustations and the asynchronous evolution of embryophyte characters during the Early Paleozoic. PDF file, International Journal of Coal Geology.
See also here.

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).
Still available through the Internet Archive´s Wayback Machine.

C. Strullu-Derrien et al. (2023): A fungal plant pathogen discovered in the Devonian Rhynie Chert. Open access, Nature Communications, 14.
"... The fungus forms a stroma-like structure with conidiophores arising in tufts outside the cuticle on aerial axes and leaf-like appendages of the lycopsid plant Asteroxylon mackiei. It causes a reaction in the plant that gives rise to dome-shaped surface projections. This suite of features in the fungus together with the plant reaction tissues provides evidence of it being a plant pathogenic fungus ..."

C. Strullu-Derrien et al. (2019): The Rhynie chert. Open access, Current Biology 29: R1211–R1223.

C. Strullu-Derrien et al. (2015): Fungal colonization of the rooting system of the early land plant Asteroxylon mackiei from the 407-Myr-old Rhynie Chert (Scotland, UK). In PDF, Botanical Journal of the Linnean Society, 179: 201–213. See also here.

Ralph E. Taggart, Department of Plant Biology, Department of Geological Sciences, Michigan State University: The First Vascular Land Plants.

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

! E.L. Taylor and T.N. Taylor (2012): Paleozoic mosses: Small, but no longer inconspicuous. In PDF, Geology, 40: 767-768.

! T.N. Taylor et al. (2004): Fungi from the Rhynie Chert: A view from the dark side. In PDF, Transactions of the Royal Society of Edinburgh, Earth Sciences, 94: 457-473.

! 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 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 and G.W. Rothwell (2006): Wetlands before tracheophytes: thalloid terrestrial communities of the Early Silurian Passage Creek biota (Virginia). PDF file, Wetlands Through Time. See also here (Google books).

Nigel H. Trewin, Stephen R. Fayers & Lyall I. Anderson, University of Aberdeen: The Biota of Early Terrestrial Ecosystems: The Rhynie Chert. The "Learning Resource" (updated 08/09/04) is primarily a resource site for students and teachers covering many aspects of the present knowledge of the unique Rhynie Chert deposit and its scientific significance (including a glossary and bibliography pages). The "Suggestions For Tutors" provides guidance for teachers (password protected). This part is primarily aimed at a university Honours degree level. The content is primarily of value in geology teaching, but has relevance to botany, zoology, ecology and history of science.

The link is to a version archived by the Internet Archive´s Wayback Machine.

Joshua Vance, University of Dayton: Rhynie Chert. Powerpoint presentation.

Y.P. Veenma et al. (2023): Biogeomorphology of Ireland's oldest fossil forest: Plant-sediment and plant-animal interactions recorded in the Late Devonian Harrylock Formation, Co. Wexford. Free access, Palaeogeography, Palaeoclimatology, Palaeoecology, 621.
Note figure 6, 7: Lignophyte root systems within the lower Sandeel Bay plant bed.
"... new evidence for early plant-sediment interactions from the Late Devonian (Famennian) Harrylock Formation (County Wexford, Ireland), which hosts standing trees that represent Ireland's earliest known fossil forest.
[...] Fossilized driftwood preserved in the lacustrine facies contains the earliest evidence for arthropod(?) borings in large vascular plant debris. Together these early examples show that plant-sediment and plant-animal interactions, frequently recorded in Carboniferous strata, were already in existence by the Devonian ..."

Matt von Konrat et al. (2010): A special issue of Phytotaxa dedicated to Bryophytes: The closest living relatives of early land plants. Editorial (PDF), Phytotaxa, 9: 5-10. Go to: Table of Contents (open access). See especially:
Matt von Konrat et al. (2010): Early Land Plants Today (ELPT): How many liverwort species are there? PDF file, Phytotaxa, 9: 22-40.

Ben Waggoner, Department of Biology, University of Central Arkansas, Conway, AR: Eukaryotes and Multicells: Origin. PDF file.

! D. Wang et al. (2019): The Most Extensive Devonian Fossil Forest with Small Lycopsid Trees Bearing the Earliest Stigmarian Roots. Current Biology, 29: 2604-2615. See also here (in PDF).
Note figure 6: Reconstruction of Guangdedendron.
! Note figure 7: Reconstruction of Xinhang Forest Landscape.
Also worth checking out: Ältester fossiler Wald Asiens entdeckt. Scinexx, in German.

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.

Wayne County Regional Educational Service Agency, (Wayne RESA): Origins of Life.
This expired link is available through the Internet Archive´s Wayback Machine.

C.H. Wellman et al. (2019): Filamentous green algae from the Early Devonian Rhynie chert. Free access, PalZ.

M. Wei-Haas (2019): Bizarre Fossils Reveal Asia's Oldest Known Forest. National Geographic Australia.

! 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 and A.C. Ball (2021): Early land plant phytodebris. Free access, Geological Society, London, Special Publications, 511: 309-320.

C.H. Wellman (2018): Palaeoecology and palaeophytogeography of the Rhynie chert plants: further evidence from integrated analysis of in situ and dispersed spores. Abstract, Phil. Trans. R. Soc. B, 373. See also here (in PDF).

! 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. (2014): Palaeophytogeography of Ordovician-Silurian land plants. In PDF.

C.H. Wellman et al. (2013): Palaeophytogeography of Ordovician-Silurian land plants. In PDF.

C.H. Wellman (2004): Palaeoecology and palaeophytogeography of the Rhynie chert plants: evidence from integrated analysis of in situ and dispersed spores. In PDF, Proc. R. Soc., B 271: 985-992.

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

! C.H. Wellman and Jane Gray (2000): The microfossil record of early land plants. PDF file, Phil. Trans. R. Soc. Lond. B, 355: 717-732.

Wikipedia, the free encyclopedia: Origin of life.

Wikipedia, the free encyclopedia: Rhynie chert.

Wikipedia, the free encyclopedia Wattieza.
See also Michelle Carr, Cosmos Online: Wattieza is world´s oldest tree. (with reconstruction of the crown portion).
This expired link is available through the Internet Archive´s Wayback Machine.

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

! 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.

Sabina Wodniok et al. (2011): Origin of land plants: Do conjugating green algae. PDF file, BMC Evolutionary Biology, 11: 104.

! 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.

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

H.-H. Xu et al. (2017): Unique growth strategy in the Earth’s first trees revealed in silicified fossil trunks from China. In PDF, PNAS, see also here

! 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).

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.

! J. Xue et al. (2016): Belowground rhizomes in paleosols: The hidden half of an Early Devonian vascular plant. In PDF, Proceedings of the National Academy of Sciences of the United States of America, 113. See also here (abstract).

W. Yuan et al. (2023): Mercury isotopes show vascular plants had colonized land extensively by the early Silurian. Free access, ScienceAdvances, 9; DOI: 10.1126/sciadv.ade9510.
Note figure 1: Conceptual model showing Hg cycling on Earth.
Figure 4: Critical records in Paleozoic sediments in stage/age level.
"... vascular plants were widely distributed on land during the Ordovician-Silurian transition (~444 million years), long before the earliest reported vascular plant fossil ..."

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Last updated January 04, 2024