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Abscission and Tissue Separation in Fossil and Extant Plants@
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Permineralized Plants and Petrified Forests@

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.

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.

Joseph E. Armstrong, Department of Biological Sciences, Illinois State University, Normal: Plant diversity. Lecture notes. Go to: The Invasion of Land.
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Lorna Ash, Department of Biological Sciences, University of Alberta: Instructional Multimedia, Multimedia Topics, Botany. Go to: Hydrasperman Reproduction. Online and downloadable flash movie.

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

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.

Andrea Bennici (2008): Origin and early evolution of land plants: Problems and considerations. PDF file, Commun Integr Biol., 1: 212-218. See also here, and there.

! 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): Palaeobotany: The Rise of the Earth's Early Forests. Open access, Current Biology, 29: R792-R794.

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.

Boston College: BC Scientist´s Fossil Discovery May Indicate Life on Land Evolved Earlier than Thought.

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

C. Kevin Boyce (2008): How green was Cooksonia? -- the importance of size in understanding the early evolution of physiology in the vascular plant lineage. PDF file, Paleobiology, 34: 179-194.
This expired link is available through the Internet Archive´s Wayback Machine.

C. Kevin Boyce et al. (2007): Devonian landscape heterogeneity recorded by a giant fungus. PDF file, Geology, 35: 399-402.
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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.
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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.

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.

! 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 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.

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.

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

Richard Cowen, Department of Geology, University of California, Davis: Comparing Plant and Animal Evolution.

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.

Marilyn Davis, Perspectives, Southern Illinois University, Carbondale: LUSH LIFE: What early land plants can tell us about earth’s family tree.
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Neil S. Davies and Martin R. Gibling (2010): Cambrian to Devonian evolution of alluvial systems: The sedimentological impact of the earliest land plants. PDF file, 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:
! Life of the PreCambrian: Archean & Proterozoic.
! The Evolution of Plants.
Powerpoint presentations. Provided by D. Freile, New Jersey City University: Historical Geology.

! 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.

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

! 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". In PDF, Phil. Trans. R. Soc. B, 370. See also here.

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.

William E. Friedman and Martha E. Cook (2000): The origin and early evolution of tracheids in vascular plants: integration of palaeobotanical and neobotanical data. Abstract and PDF file, Philos Trans R Soc Lond B Biol Sci, 355.

! Patricia G. Gensel (2008): The earliest land plants. The Annual Review of Ecology, Evolution, and Systematics, 39: 459-477.

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.

! 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.

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.

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 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). In PDF, American Journal of Botany, 100: 754-763.

Patrick Honecker, University of Cologne: Ancestors of land plants revealed. Education 2001, Leaving the Water.

C. Humphreys: Fossil bryophyte proxy contributes to palaeo-atmospheric CO2 predictions. In PDF. See also here.

C. Humphreys: Literature Review: Abiotic and Biotic Influences on the Productivity of Early Land Plants. In PDF. See also here.

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, Chlorobionta (Green Plants).

M. Alan Kazlev and Toby White: 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. Go to: Paleozoic Plants. Reconstructions of a Carboniderous swamp forest (from The Fossil Book - Fenton & Fenton, 1958, Doubleday & Co.), Early Devonian land plants (from Augusta & Burian), and the increasing terrestrial plant root depth penetration with time during the Devonian.

! 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.

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

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.

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, LMU München: Mikroorganismen aus den Cherts von Esnost und Combres/Lay (Unterkarbon, Frankreich) sowie Rhynie (Unterdevon, Schottland). Scientific project report (in German).

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.

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. (2011): Into the deep: new discoveries at the base of the green plant phylogeny. PDF file, BioEssays.

Frederik Leliaert et al.: Phylogeny and Molecular Evolution of the Green Algae. PDF file, Critical Reviews in Plant Sciences.

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

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

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

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. ..."

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 (2016): Root evolution at the base of the lycophyte clade: insights from an Early Devonian lycophyte. In PDF, Annals of botany.

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

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

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.

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

Mount Allison University, Canada ( Recap of Archean Proterozoic Life (Powerpoint presentation).

! 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!

! 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.

S.C. Morris (1993): The fossil record and the early evolution of the Metazoa. PDF file.

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.

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.

! Karl J. Niklas and Ulrich Kutschera (2010): The evolution of the land plant life cycle. PDF file, 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.

L. 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. See also here.
Note fig. 3: Landscape reconstruction showing aluvial plain in small river delta with stands of Pseudosporochnus, up to 4 m high.

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 (2017): An Early Devonian permineralized rhyniopsid from the Battery Point Formation of Gaspe (Canada). Abstract, bioRxiv. See also here (in PDF).

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.

! 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. In PDF, Plant Physiology, 177. See also here.

! 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 (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).

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. Am. J. Bot., 103: 652-662. See also here.

! 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.

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.

! 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.

! A.J. Shaw et al.(2011): Bryophyte diversity and evolution: Windows into the early evolution of land plants. In PDF, Am. J. Bot., 98: 352-369. See also here (abstract).

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. (2009): Origin and Radiation of the Earliest Vascular Land Plants. In PDF, Science, 324.

Philippe 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.

! 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 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 (2009): Thalloid carbonaceous incrustations and the asynchronous evolution of embryophyte characters during the Early Paleozoic. PDF file, International Journal of Coal Geology.

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. (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 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.

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. 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.

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

CHARLES H. WELLMAN, PETER L. OSTERLOFF, & UZMA MOHIUDDIN: Fragments of the earliest land plants. Nature Science 18/19.9.2003.

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.

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

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).

! 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).

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Last updated June 21, 2021