Links for Palaeobotanists

Home / Introductions to both Fossil and Recent Plant Taxa / Filicales

Cyanobacteria and Stromatolites
Seed Plants in General
! Triassic Palaeobotany, Palynology and Stratigraphy@
! Herbaria@
! Taxonomy and Plant Classification Databases@
! Living Fossils@
Plant Photographs@
! Paleovegetation Reconstructions@
Picture Search@


! Nan Crystal Arens, C. Strömberg and A. Thompson, Department of Integrative Biology, and Paleobotany Section, Museum of Paleontology (UCMP), University of California at Berkeley: Virtual Paleobotany. The Virtual Paleobotanical Laboratory, a comprehensive treatment of the fossil record of land plants, is divided into 12 chapters, lab I through XII. Each lab has a title page, a page with questions around the group or subject of study, a list of literature and links for further reading and exploration, and a virtual gallery of images from the lab. Go to: Sphenopsids and Ferns.

S.R. Ash (2001): The fossil ferns of Petrified Forest National Park, Arizona, and their paleoclimatological implications. Proceedings of the 6th Fossil Resource Conference.
Provided by the Internet Archive´s Wayback Machine.

! Lorna Ash & Heather Kroening, Department of Biological Sciences, University of Alberta: Instructional Multimedia, Multimedia Topics, Botany. Go to: Equisetum life cycle, Fern Life Cycle. See also here. Online and downloadable flash 4 movies. Excellent!

! Australian National Herbarium: The Fern Pages. This resource is intended to include information about the taxonomy and distribution of Australian and regional ferns and their allies as well at general information about pteridophyts. See also: A classification of the ferns and their allies - a work in progress, and
Pteridophytes: The Ferns and their Allies.

! B. Axsmith et al. (2018): A Triassic Mystery Solved: Fertile Pekinopteris From the Triassic of North Carolina, United States. Chapter 10, in PDF, in: M. Krings, C.J. Harper, N.R. Cuneo and G.W. Rothwell (eds.): Transformative Paleobotany Papers to Commemorate the Life and Legacy of Thomas N. Taylor. See also here.
Note fig. 10.1: A suggested reconstruction of Pekinopteris auriculata.

Brian Axsmith (2007): A new species of the fern Cynepteris from the Late Triassic of Arizona: Implications for the early diversification of the Schizaeales. Abstract, Botany & Plant Biology 2007, Botanical Society of America, Chicago.

M. Barbacka et al. (2019): Ferns of the Lower Jurassic from the Mecsek Mountains (Hungary): taxonomy and palaeoecology. Free access, PalZ, 93: 151–185. See also here (in PDF), and there.

M. Barbacka et al. (2016): New data about Matonia braunii (Göppert) Harris from the Early Jurassic of Poland and its ecology. In PDF, Geological Quarterly, 60: 857–868. See also here.

! The Museum of Paleontology (UCMP), University of California, Berkeley: Introduction to the Pteridopsida, The Ferns.

A.C. Bippus et al. (2019): Fossil fern rhizomes as a model system for exploring epiphyte community structure across geologic time: evidence from Patagonia. Open access, PeerJ., 7: e8244.
Note figure 2E: Coprolite-filled gallery in osmundaceous leaf base.

C. Blanco-Moreno et al. (2019): A novel approach for the metric analysis of fern fronds: Growth and architecture of the Mesozoic fern Weichselia reticulata in the light of modern ferns. Open access, PLoS ONE, 14: e0219192.

J. Bodnar et al. (2018): Middle Triassic dipterid ferns from west-central Argentina and their relationship to palaeoclimatic changes. In PDF, Acta Palaeontologica Polonica, 63: 397–416.

! B. Bomfleur et al. (2017): The fossil Osmundales (Royal Ferns)—a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes Open access, PeerJ, DOI 10.7717/peerj.3433.

B. Bomfleur et al. (2015): Osmunda pulchella sp. nov. from the Jurassic of Sweden - reconciling molecular and fossil evidence in the phylogeny of modern royal ferns (Osmundaceae). In PDF, BMC Evolutionary Biology, 5. See also here.

! B. Bomfleur et al. (2014): Fossilized Nuclei and Chromosomes Reveal 180 Million Years of Genomic Stasis in Royal Ferns. Science, 343, abstract.

! C. Kevin Boyce 2005): Patterns of segregation and convergence in the evolution of fern and seed plant leaf morphologies. PDF file, Paleobiology, 31: 117-140.

Charles Kevin Boyce, Organismic and Evolutionary Biology, Harvard Univ.: PATTERNS OF MORPHOLOGICAL EVOLUTION IN THE LEAVES OF FERNS AND SEED PLANTS. Abstract, GSA Annual Meeting, November 5-8, 2001.

The British Pteridiological Society.
The Objects of the Society are to promote all aspects of pteridology by encouraging the appreciation, conservation, cultivation and scientific study of ferns, horsetails, clubmosses and quillworts through publications, meetings, the provision of grants and other appropriate means. Go to: An Introduction to Ferns.
Snapshot taken by the Internet Archive´s Wayback Machine. This introduction (to ferns and other pteridophytes) is based on a chapter from the book "A World of Ferns", by Josephine M. Camus, A. Clive Jermy & Barry A. Thomas, Natural History Museum Publications, London.

F.M. Cardillo & T.S. Samuels, Department of Biology, Manhattan College and the College of Mt. St. Vincent: WHITTAKER FIVE KINGDOM SYSTEM (1978) Plant Classification, KINGDOM IV - Plantae, Division Pterophyta, Order Filicales.
Snapshots taken by the Internet Archive´s Wayback Machine.

Sean Carrington, Department of Biological" Chemical Sciences, University of the West Indies (UWI), Barbados: Ferns.

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

T. Choo et al. (2016): Monotypic colonies of Clathropteris meniscioides (Dipteridaceae) from the Early Jurassic of central Patagonia, Argentina: implications for taxonomy and palaeoecologyand palaeoecology. In PDF, Palaeontographica B, 294: 85-109. See also here.

! M.J.M. Christenhusz et al. (2011): A linear sequence of extant families and genera of lycophytes and ferns. PDF file, Phytotaxa, 19: 7-54.

! Michael Clayton, Department of Botany, University of Wisconsin, Madison: Instructional Technology (BotIT). Some image collections. Excellent! Go to:

! M.E. Collinson (2002): The ecology of Cainozoic ferns. Abstract, Review of Palaeobotany and Palynology.

E.P. Coturel et al. (2018): New species of osmundaceous fertile leaves from the upper Triassic of Argentina. Open access, Acta Palaeobotanica, 58: 107–119.

Jim Croft, The Australian National Herbarium: A classification of the ferns and their allies - a work in progress. This classification of the genera of ferns and their allies is a loose, perhaps tenuous, concensus of a number of published systems, some of which are available on the web. It tries to reflect contemporary views on phylogenetic relationships and as such will change from time to time.
The link is to a version archived by the Internet Archive´s Wayback Machine.

S.H. Deng and P. Shang (2000): A Brief Review of the Mesozoic Filicopsida in China. PDF file.

Dagmar Dietrich et al. (2000): Analytical X-Ray Microscopy on Psaronius sp.: A Contribution to Permineralization Process Studies. Abstract, Mikrochim. Acta, 133: 279-283.

! W.A. DiMichele and T.L. Phillips (2002): The ecology of Paleozoic ferns. In PDF, Review of Palaeobotany and Palynology.

! dmoz, the Open Directory Project:
Science: Biology: Flora and Fauna: Plantae:
Polypodiophyta. See also:
Earth Sciences: Paleontology: Paleobotany: Taxa.

J.Y. Dubuisson et al. (2009): Epiphytism in ferns: diversity and history. In PDF, Comptes rendus biologies. See also here (abstract).

Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey, UK: Research activities,
Fern palaeobiology.
Now recovered from the Internet Archive´s Wayback Machine.

! Encyclopedia of Earth (supported by the Environmental Information Coalition and the National Council for Science and the Environment). Expert-reviewed information about the Earth. For everyone, please take notice. Provided by the Internet Archive´s Wayback Machine. The scope of the Encyclopedia of Earth is the environment of the Earth broadly defined, with particular emphasis on the interaction between society and the natural spheres of the Earth. Excellent! Go to:
! Fern.

H.J. Falcon-Lang (2005): Adpressed tree-fern trunks from the Early Pennsylvanian Joggins Formation of Nova Scotia. In PDF, Atlantic Geology, 41: 169–172.

M. Farahimanesh et al. (2014): The fern Stauropteris oldhamia Binney: New data on branch development and adaptive significance of the hypodermal aerenchyma. In PDF, C. R. Palevol., 13: 473–481.

G. Grimm et al. (2015): Using more than the oldest fossils: Dating Osmundaceae with three Bayesian clock approaches. Open access, Systematic Biology, 64: 396-405. See also here (in PDF).

G. Guignard et al. (2009): A dipteridaceous fern with in situ spores from the Lower Jurassic in Hubei, China. In PDF, Review of Palaeobotany and Palynology, 156: 104–115.

! C.H. Haufler (2014): Ever since Klekowski: Testing a set of radical hypotheses revives the genetics of ferns and lycophytes. In PDF, American Journal of Botany, 101: 2036-2042.

Xiaoyuan He et al. (2010): Anatomically Preserved Marattialean Plants from the Upper Permian of Southwestern China: The Trunk of Psaronius laowujiensis sp. nov. PDF file, Int. J. Plant Sci 171: 662-678.

F. Herrera et al. (2017): An exquisitely preserved filmy fern (Hymenophyllaceae) from the Early Cretaceous of Mongolia. Free access, American Journal of Botany, 104: 1370-1381. See also here (in PDF).

Monte Hieb and Harrison Hieb, Plant Fossils of West Virginia: Ferns and Seed Ferns. Fossil Plants of the Middle Pennsylvanian Period. Go to: How to tell Neuropteris from Pecopteris or Alethopteris?
Snapshots taken by the Internet Archive´s Wayback Machine.

Josef Hlasek: Photo Gallery wildlife pictures, Plants. Go to: Plants - Pteridophyta.

G. Holzhüter et al. (2003): Structure of silica in Equisetum arvense. In PDF, Anal. Bioanal. Chem., 376: 512-517.
Snapshot taken by the Internet Archive´s Wayback Machine.

SHARON D. KLAVINS et al.: MATONIACEOUS FERNS (GLEICHENIALES) FROM THE MIDDLE TRIASSIC OF ANTARCTICA. Abstract, Journal of Paleontology, 2004; v. 78; no. 1; p. 211-217.

John A. Knouse, Athens, Ohio: Ferns and Fern Allies. See also: Fern Book Bibliography. Periodicals and books dedicated to pteridology.

E. Kustatscher et al. (2012): Danaeopsis Heer ex Schimper 1869 and its European Triassic species. Abstract, Review of Palaeobotany and Palynology, 183: 32-49.

E. Kustatscher & J.H.A. van Konijnenburg-van Cittert (2011): The ferns of the Middle Triassic flora from Thale (Germany). Abstract, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 261: 209-248.

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

S. Lehtonen (2011): Towards Resolving the Complete Fern Tree of Life. In PDF.

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

The Los Angeles International Fern Society: FERN BASICS.

Department of Botany, University of Wisconsin, Madison: Plant Systematics Collection. This web site provides structured access to a teaching collection of plant images representing over 250 families and 1000 genera of vascular plants. Go to: Phylum Pterophyta. The Ferns.

Eugene Marinus, Department of Biodiversity and Conservation Biology, University of the Western Cape: Ferns in the Carboniferous Period (Powerpoint presentatation).

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, FOSSIL FERNS. Link list page with rankings and brief explanations. Images of Psaronius, Psaronius melanedrus, Tietea singularis, Pecopteris, Pecopteris cyathea, Pecopteris oreopteridia, Pecopteris arborescens, P. (Senftenbergia) plumosa, Asterotheca, Scolecopteris, Scolecopteris (P.) mertensiodes, Ptychocarpus (P.) unita, Ptychocarpus (P.) unita, Senftenbergia crenata, Alloiopteris coralloides, Anachoropteris involuta, Ankyropteris, Botryopteris, Etapteris, Oligocarpia gutbieri, Pseudosporochnus, Saccopteris cristata, Stauropteris.
Snapshot taken by the Internet Archive´s Wayback Machine.

Andrew G. Murdock (2008): Phylogeny of marattioid ferns (Marattiaceae): inferring a root in the absence of a closely related outgroup. PDF file, American Journal of Botany, 95: 626-641.

Dennis C. Murphy, ("Devonian Times", a paleontology web site featuring Red Hill): Who's Who at Red Hill, Gillespiea randolphensis (Early "Fern"), and Rhyacophyton ceratangium (Early "Fern"). See also: More about Ferns.

Nature Hills Nursery, USA: What is a Fern?

Nederlandse Varenvereniging (in Dutch):

Karl J. Niklas & Tom Silva, Department of Plant Biology, Cornell University, Ithaca, NY: Introductory Botany. Review Topics, Review of Algae, Bryophytes, Pteridophytes- Common Links Between Each Group of Plants.
These expired links are available through the Internet Archive´s Wayback Machine.

! C.N. Page (2002): Ecological strategies in fern evolution: a neopteridological overview. Abstract, Review of palaeobotany and palynology, 119: 1-33. See also here and there (in PDF).

C.J. Phipps et al. (1998): Osmunda (Osmundaceae) from the Triassic of Antarctica: an example of evolutionary stasis. PDF file, American Journal of Botany, 85: 888-895.

A.R.G. Plackett et al. (2015): Ferns: the missing link in shoot evolution and development. Front. Plant Sci., 6.

! PPG I (2016); This project was organized by Eric Schuettpelz: A community-derived classification for extant lycophytes and ferns. Free access, Journal of Systematics and Evolution.

! K.M. Pryer and E. Schuettpelz (2009): Ferns. PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

! Kathleen M. Pryer, Alan R. Smith and Carl Rothfels (2009): The Tree of Life Web Project, Ferns, Polypodiopsida Cronquist, Takht. & Zimmerm. 1966.

! Kathleen M. Pryer et al. (2004): Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. PDF file, American Journal of Botany, 91: 1582-1598. See also here (abstract).

K.M. Pryer et al. (2001): Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Abstract, Nature, 409: 618-622.
! See also here (in PDF).

! Kathleen M. Pryer, Department of Botany, The Field Museum, Chicago, and Alan R. Smith, University Herbarium, University of California, Berkeley (part of Tree of Life, the University of Arizona): Leptosporangiate Ferns. Next to the flowering plants, the leptosporangiate ferns are the most diverse group of living land plants. Recent estimates place their diversity at about 12,000 species in 300 genera.

J. Pšenièka and E.L. Zodrow (2006, starting on PDF page 18 (Sec1:16)): Pennsylvanian fern taxonomy: New approach through the compact model. In PDF, Newsletter on Carboniferous Stratigraphy, 24.

The Pteridophyte Phylogeny Group (PPG), Smithsonian National Museum of Natural History. PPG aims to produce, and continually update, a community-derived classification for lycophytes and ferns - based on the understanding of phylogeny - at the family and genus levels.

L. Regalado et al. (2017): A fossil species of the enigmatic early polypod fern genus Cystodium (Cystodiaceae) in Cretaceous amber from Myanmar. Scientific Reports, 7.

J.W.F. Reumer et al. (2020): The Rhaetian/Hettangian dipterid fern Clathropteris meniscioides Brongniart found in erratics in the eastern Netherlands and adjacent Germany. In PDF, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 295: 297–306.

! R. Rößler (2000): The late Palaeozoic tree fern Psaronius - an ecosystem unto itself. In PDF, Review of Palaeobotany and Palynology, 108: 55-74.

! C.J. Rothfels et al. (2012): A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). In PDF, Taxon, 61: 515-533.

Gar W. Rothwell, Edith L. Taylor and Thomas N. Taylor: Ashicaulis woolfei n. sp.: additional evidence for the antiquity of osmundaceous ferns from the Triassic of Antarctica. Abstract, American Journal of Botany. 2002; 89: 352-361.

Scott Russell, Department of Botany and Microbiology, University of Oklahoma, College of Arts and Sciences, Norman: Morphology of Vascular Plants. Lecture notes, chiefly PDF files, including palaeobotany topics. Scroll down to: Lecture / Lab 12 - The Eusporangiate Ferns, Click: "Lab Images". See also: Lecture / Lab 12 - The Eusporangiate Ferns (PDF file).

H. Schneider (2019): Celebrating Research Devoted to Seed-Free Land Plants. Free access, Journal of Systematics and Evolution, 57: 303-304. See also here (table of contents, free access).

H. Schneider et al. (2016): Burmese amber fossils bridge the gap in the Cretaceous record of polypod ferns. In PDF, Perspectives in Plant Ecology, Evolution and Systematics, 18: 70–78. See also here (abstract).

H. Schneider and E. Schuettpelz (2016): Systematics and evolution of lycophytes and ferns: Editorial. Abstract, Journal of Systematics and Evolution, 54: 561–562. See also here (in PDF).

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

Harald Schneider, Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Georg-August-Universität, Göttingen: Vielfalt der Farne entwickelte sich im "Schatten" der neuen Blütenpflanzen (in German).

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

E. Schuettpelz and K.M. Pryer (2007): Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. In PDF, Taxon, 56: 1037–1050. See also here (abstract).

! C.S. Shi et al. (2013): Characterization of the stem anatomy of the Eocene fern Dennstaedtiopsis aerenchymata (Dennstaedtiaceae) by use of confocal laser scanning microscopy. Free access, American Journal of Botany, 100: 1626–1640.

J.E. Skog (2001): Biogeography of Mesozoic leptosporangiate ferns related to extant ferns. In PDF, Brittonia, 3: 236-269.

Andrew C. Scott et al. (2009): Scanning Electron Microscopy and Synchrotron Radiation X-Ray Tomographic Microscopy of 330 Million Year Old Charcoalified Seed Fern Fertile Organs. PDF file, Microsc. Microanal., 15: 166-173. See figure 4, SEM of charcoalified pteridosperm ovule from the mid-Mississippian (Carboniferous). See also here.

M.K.A. Smith et al. (2015): Mesozoic Diversity of Osmundaceae: Osmundacaulis whittlesii sp. nov. in the Early Cretaceous of Western Canada. Abstract, Journal of Plant Sciences, 176: 245-258. See also here (in PDF).

! A.R. Smith et al. (2008): Fern classification. In PDF.

! A.R. Smith et al. (2006). A classification for extant ferns. PDF file, Taxon 55: 705-731.

! K.L. Sporne (1962): The morphology of pteridophytes; the structure of ferns and allied plants (PDF file). See also here.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Plant life in the Silurian, Devonian, Carboniferous, Permian and Cretaceous. Go to: The tree fern Psaronius, and The tree fern Tempskya.

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
BOT335 Lecture Schedule.
Psaronius: a Carboniferous tree-fern;
Snapshots taken by the Internet Archive´s Wayback Machine.

M. Taghi Badihagh and D. Uhl (2019): The first occurrence of Phlebopteris dunkeri and P. woodwardii (Matoniaceae) from the middle Jurassic of Iran, In PDF, # Journal of Palaeogeography. See also here.

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

Herbarium, Department of Biology, Texas A&M University, College Station: Field Systematic Botany, PTERIDOPHYTES (ferns and allies). An overview and link list.

N. Tian et al. (2016): A systematic overview of fossil osmundalean ferns in China: Diversity variation, distribution pattern, and evolutionary implications. Abstract, Palaeoworld, 25: 149–169. See also here (in PDF).

J.H.A. Van Konijnenburg-van Cittert et al. (2020): Ferns and fern allies in the Rhaetian flora of Wüstenwelsberg, Bavaria, Germany. In PDF, Review of Palaeobotany and Palynology 273. See also here.

J.H.A. van Konijnenburg-van Cittert et al. (2018): Phialopteris heterophylla (Sternberg ex Göppert, 1836) comb. nov., A rare schizaeaceous fern from the Early Jurassic of Bavaria. In PDF, Fossil Imprint, 74: 5–64. See also here.

! J.H.A. Van Konijnenburg-Van Cittert (2002): Ecology of some Late Triassic to Early Cretaceous ferns in Eurasia. In PDF, Review of Palaeobotany and Palynology, 119: 113-124.

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

! A. Vasco et al. (2013): The evolution, morphology, and development of fern leaves. In PDF, Frontiers in plant science.
This expired link is now available through the Internet Archive´s Wayback Machine.
See also here (abstract).

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

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.

Y. Wang et al. (2015): Fertile structures with in situ spores of a dipterid fern from the Triassic in southern China. In PDF, Journal of Plant Research, 128.

Thomas R. Warne and Leslie G. Hickok, Department of Botany, University of Tennessee Knoxville, TN (supported by the National Science Foundation (NSF-DUE): C-Fern. Ceratopteris as a model plant system.

Biology Department, Western Washington University, Bellingham, Washington:
! Seedless Vascular Plants (Ferns, etc.) Powerpoint presentation. See also here, or there.

Wikipedia, the free encyclopedia: Fern. See also here (in German).
Fern spike.

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Last updated February 02, 2021

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