Links for Palaeobotanists

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

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


B.J. Axsmith et al. (2003): The enigmatic Paleozoic plants Spermopteris and Phasmatocycas reconsidered. PDF file, American Journal of Botany, 90: 1585-1595.

AXSMITH, BRIAN J1, RUDOLPH SERBET2, MICHAEL KRINGS2, THOMAS N TAYLOR2, and EDITH L TAYLOR2. 1Department of Biological Sciences, University of South Alabama, Mobile, Alabama; 2Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Research Center, University of Kansas, Lawrence. - New intepretations of the enigmatic Paleozoic plants Spermopteris and Phasmatocycas. Abstract. Botany 2001, August 12 - 16, 2001; Albuquerque, New Mexico.

M.R. Barone Lumaga et al. (2015): Epidermal micromorphology in Dioon: did volcanism constrain Dioon evolution? In PDF, Botanical Journal of the Linnean Society, 2015, 179: 236–254. See also here.

Phil and Jesse Bergman, Jungle Music Palms and Cycads, San Diego CA: Cycads. Visit the Cycad Photo Gallery. There are 437 pictures in the gallery.

Museum of Paleontology (UCMP), University of California, Berkeley: Introduction to the Cycads, Legacy of the Mesozoic. See also: Cycads: Fossil Record.

Blue Forest Safaris (Morne Fouche): Africa Cycads.

E.R. Bodor (2015): Plant reproductive organs from the Mecsek Coal Formation. Thesis, (abbreviated version?), ELTE Doctoral School of Earth Sciences, Budapest. In PDF.
Conclusion about "Dioonitocarpidium"(?).

J.G. Bornemann (1856): Über organische Reste der Lettenkohlengruppe Thüringens: Ein Beitrag zur Fauna und Flora dieser Formation, besonders über fossile Cycadeen, nebst ... (in German). Provided by Google books.

E.D. Brenner et al. (2003): Cycads: evolutionary innovations and the role of plant-derived neurotoxins. In PDF, Trends in Plant Science, 8.

! Robert Buckley Trabuco Canyon, California, (Illustrations by Douglas Henderson, John Sibbick and Mark Hallet), The Palm & Cycad Societies of Florida (PACSOF): The Fossil Cycads. See also
! here (in PDF).

Robert Buckley Trabuco Canyon, California: Inducing adventitious root growth in cycad leaves. Reprinted with permission from The Cycad Newsletter, Issue 1, 1999. This article is a 388 Kb PDF document and must be viewed using the Adobe Acrobat Reader plug-in.

Benjamin Burger, Utah State University, Vernal, Utah:
Why study fossil plants?
Invertebrate Paleontology and Paleobotany.
How did plants colonize the land, based on the fossil record?
How did the first seed plants (the Gymnosperms) evolve?
How did gymnosperms diversify during the early Mesozoic to become a modern dominate plant group?
! How good is the fossil record of Cycads?
What is the significance of the fossil record of Ginkgo?
What is the fossil record of Horsetails?
Fossil Algae.
What is an Angiosperm?
Video lectures.

! R.J. Butler et al. (2009): Testing co-evolutionary hypotheses over geological timescales: interactions between Mesozoic non-avian dinosaurs and cycads. PDF file, Biol. Rev., 84: 73-89. See also here (abstract).

Bane Cheek, Polk Community College, Winter Haven, Florida: Cycads. Now provided by the Internet Archive´s Wayback Machine. See especially: An Introduction to Cycad Reproduction.

Curtis Clark, Biological Sciences Department California State Polytechnic University, Pomona: Plant Morphology. Resources. Go to: Lab 12: Cycadophyta, Ginkgophyta.

Lynne M. Clos, Fossil News: Cycads: Living Fossils. See also: Field Adventures in Paleontology: Hunting For Fossil Cycads in Petrified Forest National Park, Arizona.
Still provided by the Internet Archive´s Wayback Machine.

F.L. Condamine et al. (2015): Origin and diversification of living cycads: a cautionary tale on the impact of the branching process prior in Bayesian molecular dating. In PDF, BMC Evolutionary Biology.

! N.R. Cúneo et al. (2010): Review of the Cycads and Bennettitaleans from the Mesozoic of Argentina. In PDF.

The Cycad Newsletter. Provided by the Cycad Society, a non-profit organization dedicated to the conservation of cycads through education and scientific research.

! The Cycad Society. A non-profit organization dedicated to the conservation of cycads through education and scientific research. Read examples of the interesting articles that have appeared in The Cycad Newsletter. Visit The Cycad Photo Gallery.

! The Cycad Society: Glossary of Cycad Terms. Consisting of two parts: Illustrated Glossary of Cycad Terms, and Expanded Glossary of Cycad Terms (PDF file).

Jose Manuel Sanchez de Lorenzo Cáceres, Palm & Cycad Societies of Florida, Inc.: The Cycads: Fossils of the Past.

Paolo De Luca, Dipartimento di Biologia vegetale and Orto Botanico, Universita di Napoli, Italia: A Historical Perspective on Cycads from Antiquity to the Present. The historical aspects of the acquisitions of cycad collections in botanical gardens are outlined, and a brief survey of the history of cycads in various cultures is given.

The Cycad Society of South Africa.

V.M. Dörken and B. Rozynek (2013): Proliferated megasporangiate strobili of Zamia furfuracea (Zamiaceae, Cycadales) and its possible evolutionary implications for the origin of cycad-megasporophylls. In PDF, Palaeodiversity, 6: 135-147.

! C.C. Davis and H. Schaefer (2011): Plant Evolution: Pulses of Extinction and Speciation in Gymnosperm Diversity. See also here (abstract).

S. Deng et al. (2014): Pollen cones and associated leaves from the Lower Cretaceous of China and a re-evaluation of Mesozoic male cycad cones: In PDF, Journal of Systematic Palaeontology, 12.

W.A. DiMichele et al. (2001): An Early Permian flora with Late Permian and Mesozoic affinities from north-central Texas. In PDF.

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

V.M. Dörken and B. Rozynek (2013): Proliferated megasporangiate strobili of Zamia furfuracea (Zamiaceae, Cycadales) and its possible evolutionary implications for the origin of cycadmegasporophylls. In PDF, Palaeodiversity 6: 135-147.

! Christopher J. Earle (server space has been provided by the Department of Botany, Rheinische Friedrich-Wilhelms-Universität Bonn, Germany): The Gymnosperm Database. Currently the database provides basic information for all species and higher-ranked taxa of the gymnosperms, i.e., conifers, cycads, and their allies. You may navigate from the species list, alphabetized by binomial.

! B. Erdei and S.R. Manchester (2015): Ctenis clarnoensis sp. n., an Unusual Cycadalean Foliage from the Eocene Clarno Formation, Oregon. In PDF, Int. J. Plant Sci., 176: 1–43.

Palmengarten, Frankfurt am Main, Das Tor zur Welt der Pflanzen: Palmfarne - weder Palmen noch Farne (in German).

! Florida Museum of Natural History, University of Florida, Gainesville:
Cycads and Ginkgo (Powerpoint presentatation).

Debbie Folkerts, Auburn University, Auburn, Alabama: Kingdom Plantae: Gymnosperms. Powerpoint presentation.

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

David R. Greenwood, Victoria University of Technology, Melbourne (Palm and Cycad Societies of Australia, PACSOA): Cycad Articles, Australian Fossil Cycads. Provided by the Internet Archive´s Wayback Machine.

Guido Grimm, Department of Palaeobotany, Swedish Museum of Natural History, Stockholm: Cladistic analyses of fossil and recent Cycadales based on morphological and molecular data. See also:
The Best Tree Cladogram, and here (abstract), and there "Phylogenie der Cycadales" (in German).

T.M. Harris (1961): The fossil cycads. PDF file.

! Jody L. Haynes (c/o Palm and Cycad Societies of Florida): Virtual Cycad Encyclopedia. Information about cycad taxonomy, biology, evolution, horticulture, conservation, etc.

M. Haworth et al. (2011): Cycads show no stomatal-density and index response to elevated carbon dioxide and subambient oxygen. Abstract, Australian Journal of Botany.

E.J. Hermsen et al. (2009): Morphology and ecology of the Antarcticycas plant. PDF file, Review of Palaeobotany and Palynology, 153: 108-123. Antarcticycas plant reconstruction on page 112.

E. Hermsen et al. (2007): Cycads from the Triassic of Antarctica: Permineralized cycad leaves. PDF file, Int. J. Plant Sci., 168: 1099-1112.

E.J. Hermsen et al. (2006): Cataphylls of the Middle Triassic cycad Antarcticycas schopfii and new insights into cycad evolution. PDF file, American Journal of Botany, 93: 724-738. See also here.

! Ken D. Hill et al. (2004): The World List of Cycads. PDF file.

! Ken D. Hill and D.W. Stevenson, Royal Botanic Gardens Sydney: The Cycad Pages. Botanical information and links, including identification and classification of cycads, ecology and conservation, cycad evolution and the fossil record, geography and distribution, as well as horticultural and general information. Excellent! Go to: John Hendricks, Ken D. Hill, Roy Osborne and Dennis Stevenson, Royal Botanic Gardens Sydney: The World List of Cycads. The lists as published gave the valid names of all known extant cycads.

L. Kessler, The New York Times: The Cult of the Cycads.

S.D. Klavins et al. (2005): Coprolites in a Middle Triassic cycad pollen cone: evidence for insect pollination in early cycads? PDF file, Evolutionary Ecology Research, 7: 479-488.

S.D. Klavins et al. (2003): Gymnosperms from the Middle Triassic of Antarctica: the first structurally preserved cycad pollen cone. In PDF, Int. J. Plant Sci., 164: 1007-1020.

E. Kustatscher and J.H.A. van Konijnenburg-van Cittert (2010): Seed ferns and Cycadophytes from the Triassic Flora of Thale. In PDF, N. Jb. Geol. Paläont. Abh.

Palm and Cycad Societies of Australia (PACSOA).

The Palm Haven. Palm Trees, Cycads, Bromeliads and other Tropical Plants. Go to: Cycads.

Conrad C. Labandeira et al. (2007): Pollination drops, pollen, and insect pollination of Mesozoic gymnosperms. PDF file, Taxon, 56:663-695.

Gerhard Leubner Lab, University Freiburg, Germany: Seed Evolution. Go to: Jurassic parc/Mesozoic era: Extinct gymnosperms and living ancient gymnosperms. Extant Cycadales and extinct Bennettitales.

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

Yuandan MA et al. (2009): Carbon storage of cycad and other gymnosperm ecosystems in China: implications to evolutionary trends. PDF file, Polish Journal of Ecology, 57: 635-646.
This expired link is available through the Internet Archive´s Wayback Machine.

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 Cycadophyta The Cycads).

SERGIUS H. MAMAY, DAN S. CHANEY, AND WILLIAM A. DIMICHELE: A Mesozoic-type flora from the Lower Permian of Texas. Abstract, 1998 Annual Meeting of the Botanical Society of America, 2-6 August, 1998 Baltimore, MD. Specimens of Dioonitocarpidium! Provided by the Internet Archive´s Wayback Machine.

! S.H. Mamay (1976): Paleozoic Origin of the Cycads. In PDF, Geological Survey Professional Paper, 934. See also here.

! Y. Miao et al. (2017): Fossil and Living Cycads Say "No More Megasporophylls". J. Morphol. Anat., 1: 107. See also here and there (in PDF).

John M. Miller (, University of California, Berkeley: Origin of Angiosperms. See also here or navigate from essay contents. Go to:
! Cycadales.

G.E. Mustoe (2007): Coevolution of cycads and dinosaurs. Cycad Newsletter.

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, CYCADS. Link list page with rankings and brief explanations. Images of modern cycads.

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

netfirms: The Cool Tropics, Research Cycads. A directory.
Snapshot provided by the Internet Archive´s Wayback Machine.

Paläontologische Museum Nierstein: Image of Dioonitocarpidium pennaeforme (website in German).

! Y. Ogura, Department of Botany, Faculty of Science, University of Tokyo, Tokyo, Japan, (page hosted by Botany online, The Internet Hypertextbook, Biological Classics in the Internet): HISTORY OF DISCOVERY OF SPERMATOZOIDS IN GINKGO BILOBA AND CYCAS REVOLUTA. PHYTOMORPHOLOGY, Vol 17, 109 - 114 (1967).

The Palm and Cycad Societies of Australia (PACSOA): PACSOA is a group of affiliated, semi-autonomous, regional Palm and/or Cycad Societies, based in Brisbane, Queensland.

The Palm and Cycad Society of Florida. The fossil Cycads. With paintings and reconstructions of Douglas Henderson, John Sibbick, and Mark Hallett.

The Palm & Cycad Societies of Florida (PACSOF): Virtual Cycad Encyclopedia. Worth to visit: Chapter 4: Cycad evolution.

Bill Parker, Arizona: Chinleana. Discussion of Late Triassic paleontology and other assorted topics. Go to:
Discovery of an Entire Fossil Cycad from the Late Triassic of China (September 07, 2009).

Joe & Karen Perner, Katherine, Northern Territory, Australia: Cycad International. A Cycad Gardens Nursery. Go to: What are Cycads, or visit the Cycads Gallery.
Snapshot provided by the Internet Archive´s Wayback Machine.

! Kathleen B. Pigg, Department of Plant Biology, Arizona State University: Laboratory. Go to: The Cycads, Cycadeoids (Bennettitales) and Ginkgophytes.

C. Pott and A. Launis (2015): Taeniopteris novomundensis sp. nov. – "cycadophyte" foliage from the Carnian of Switzerland and Svalbard reconsidered: How to use Taeniopteris? Abstract, Neues Jahrbuch für Geologie und Paläontologie, 275. See also here (in PDF).

! Christian Pott and Michael Krings (2010): Gymnosperm Foliage from the Upper Triassic of Lunz, Lower Austria: an annotated check list and identifiation key. PDF file, Geo.Alp, 7: 19-38.

! C. Pott et al. (2012): Baikalophyllum lobatum and Rehezamites anisolobus: Two Seed Plants with "Cycadophyte" Foliage from the Early Cretaceous of Eastern Asia. Paper awarded with the Remy and Remy Award 2012, Botanical Society of America.

! C. Pott et al. (2010): Late Palaeozoic foliage from China displays affinities to Cycadales rather than to Bennettitales necessitating a re-evaluation of the Palaeozoic Pterophyllum species. PDF file, Acta Palaeontol. Pol., 55: 57-168.

C. Pott et al. (2007): Pseudoctenis cornelii nov. spec. (cycadalean foliage) from the Carnian (Upper Triassic) of Lunz, Lower Austria. PDF file, Ann. Naturhist. Mus. Wien, 109 A: 1-17.

S.E. Pratt (2014): Benchmarks: September 1, 1957: Fossil Cycad National Monument is dissolved. Earth, September 01, 2014.
See also here (National Fossil Day).

Alberto Prado (2011): The Cycad Herbivores. PDF file, Bulletin de la Société d´entomologie du Québec.

Mary Ann Reihman, Biological Sciences, Natural Sciences and Mathematics, California State University, Sacramento: Biology lecture notes. Go to: Comparison of Cycadophyte Misc. Gymnosperms (Bennettitales).

Bernard Renault, 1881-1885 (provided by Gallica): Cours de botanique fossile fait au Muséum d'histoire naturelle.- Première année (1881). Cycadées, zamiées, cycadoxylées, cordaïtées, poroxylées, sigillariées, stigmariées.

! S. Renner (2009): Gymnosperms. Provided by the Internet Archive´s Wayback Machine. PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

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

B. Rozynek (2008): Schozachia donaea n. gen., n. sp., a new cycad megasporophyll from the Middle Triassic (Ladinian) of Southern Germany. PDF file, PALAEODIVERSITY 1: 1-18.

! D.E. Salas-Leiva et al. (2013): Phylogeny of the cycads based on multiple single-copy nuclear genes: congruence of concatenated parsimony, likelihood and species tree inference methods. In PDF, Annals of Botany, 112: 1263–1278. See also here (abstract).

Vincent L. Santucci, National Park Service, Kemmerer, WY, and Marikka Hughes, Peabody Museum of Natural History, Paleobotany Division, New Haven: Fossil Cycad National Monument: A Case of Paleontological Resource Mismanagement. This expired link is available through the Internet Archive´s Wayback Machine.

D. Schneider et al. (2002): Cycads: their evolution, toxins, herbivores and insect pollinators. PDF file.

! J. Schuster (1932):
Das Pflanzenreich, IV, Cycadaceae. In German, Verlag Wilhelm Engelmann, Leipzig.

Scott Singleton, Houston Gem and Mineral Society: Petrified Wood Articles by HGMS Authors and Others, Cycad Anatomy and Fossil Occurrences in Texas. PDF file, 2006.

! A.R.T. Spencer et al. (2017): New insights into Mesozoic cycad evolution: an exploration of anatomically preserved Cycadaceae seeds from the Jurassic Oxford Clay biota. PeerJ 5.
Description of a new genus of anatomically preserved gymnosperm seed from the Callovian–Oxfordian (Jurassic) Oxford Clay Formation (UK), using a combination of traditional sectioning and synchrotron radiation X-ray micro-tomography (SRXMT).

STOCKEY, RUTH1 and GAR W. ROTHWELL2. 1Department of Biological Sciences, University of Alberta, Edmonton: 2Department of Environmental and Plant Biology, Ohio University, Athens, Ohio: Permineralized cycad seeds from the Jurassic of British Columbia, Canada. Abstract. Botany 2001, August 12 - 16, 2001; Albuquerque, New Mexico.

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
! BOT335 Lecture Schedule. Some interesting chapters in terms of palaeobotany, e.g.
The First Vascular Land Plants;
Carboniferous Forests;
Arborescent Lycopods;
Psaronius: a Carboniferous tree-fern;
Carboniferous Horsetails;
Carboniferous Seed Ferns;
The Evolution of Conifers;
Cycadophytes, the True Cycads;
Mesozoic Cycadeoids;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

! T.N. Taylor (1969): Cycads: Evidence from the Upper Pennsylvanian. In PDF, Nature 164.

! J.H.A. van Konijnenburg-van Cittert et al. (2017): Differentiation of the fossil leaves assigned to Taeniopteris, Nilssoniopteris and Nilssonia with a comparison to similar genera. Abstract, Review of Palaeobotany and Palynology, 237: 100–106. See also here (in PDF).

Shi-Jun Wang et al. (2011): Cycad Wood from the Lopingian (Late Permian) of Southern China: Shuichengoxylon tianii gen. et sp. nov. PDF file, Int. J. Plant Sci., 172: 725-734.

! X. Wang et al. (2009): The discovery of whole-plant fossil cycad from the Upper Triassic in western Liaoning and its significance. Chinese Science Bulletin, 54: 3116–3119.
See also here. (in PDF).
Also worth checking out: Discovery of an Entire Fossil Cycad from the Late Triassic of China (by Bill Parker, September 07, 2009).

! T. Walters and R. Osborne (eds., 2004): Cycad Classification: Concepts and Recommendations. 288 pages. Provided by CABI Publishing through the Google Print Publisher Program. Registration procedure required. Use "More results from this book" or "Search this book" to navigate. Unfortunately, you can view two pages around your search result, but you can search again! Use Google Book Search to search the full text of books.

George Reber Wieland, (photograph). Go to: Vincent L. Santucci, National Park Service, Kemmerer, WY, and Marikka Hughes, Peabody Museum of Natural History, Paleobotany Division, New Haven: Fossil Cycad National Monument: A Case of Paleontological Resource Mismanagement. See also here.

Wikipedia, the free-content encyclopedia:
! Cycadophyta.
Fossil Cycad National Monument.

! P. Wilf et al. (2016): The last Patagonian cycad, Austrozamia stockeyi gen. et sp. nov., early Eocene of Laguna del Hunco, Chubut, Argentina. In PDF, Botany, 94: 817-829. See also here.

J.M. Zgurski et al. (2008): How well do we understand the overall backbone of cycad phylogeny? New insights from a large, multigene plastid data set. PDF file, Molecular Phylogenetics and Evolution.

! J.-W. Zhang et al. 2010): A new species of Leptocycas (Zamiaceae) from the Upper Triassic sediments of Liaoning Province, China. Abstract, Journal of Systematics and Evolution, 48: 286–301. See also here (in PDF).
See fig. 9: Reconstruction of Leptocycas yangcaogouensis, resembling like that of Dioon edule.

! Y.J. Zhang et al. (2015): Extending the generality of leaf economic design principles in the cycads, an ancient lineage. In PDF, New Phytologist.

Zhu Jia-nan and Du Xian-ming: A New Cycad - Primocycas chinensis gen. et sp. nov. discovered from the Lower Permian in Sha-nxi, China and its Significence. Abstract, Journal of Integrative Plant Biology (formerly Acta Botanica Sinica).

G. Zijlstra et al. (2016): (2438–2439) Proposal to conserve the names Taeniopteris and T. vittata with a conserved type (fossil Tracheophyta: "Taeniopterides"). Taxon, 65: 399-400.

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Last updated October 19, 2017

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