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

T. Agarwal (2018): Pentoxylales A Remarkable Order in the Evolution of the Plants. Research and Reviews: Journal of Pharmacognosy In PDF, and Phytochemistry.

! J.M. Anderson and H.M. Anderson (2023): Molteno Kannaskoppia: Mid-Triassic gymnosperm case study for whole-plant taxonomy. In PDF, 82 MB!. Palaeontologia africana, 57 (Special issue). Annals of the Evolutionary Studies Institute University of Witwatersrand.
See likewise here.
"... The flora from the Upper Triassic Molteno Formation, southern Africa, is the most extensively collected and documented macro-flora in the Gondwana Triassic
[...] In this volume, the genus Kannaskoppia and affiliates, in the order Petriellales, are described in greater detail
[...] Whole-plant species from the Molteno have been recognized, based on considerations of affiliation and taphonomy ..."

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

C.C. Baskin and J.M. Baskin (2023): The rudimentary embryo: an early angiosperm invention that contributed to their dominance over gymnosperms. Open access, Seed Science Research, 1–12. https://doi.org/10.1017/ S0960258523000168.
Note table 1: Information about fossil ovules, seeds and embryos of gymnosperms from the Upper Devonian to Late Cretaceous.

! L.M. Bowe et al. (2000): Phylogeny of seed plants based on all three genomic compartments: Extant gymnosperms are monophyletic and Gnetales' closest relatives are conifers. In PDF, PNAS, 97: 4092–4097. See also here.

D.M. Bruns Stockrahm, Biosciences Department, Minnesota State University Moorhead: Organismal Biology, Gymnosperms. Powerpoint presentation.

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.

! Maarten J.M. Christenhusz et al. (2011): A new classification and linear sequence of extant gymnosperms. PDF file, Phytotaxa, 19: 55-70.

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

Peter Crane, The Tree of Life Web Project (coordinator and editor: David R. Maddison): Spermatopsida, Seed Plants.

M.J. Dallwitz Australia (DELTA – DEscription Language for TAxonomy).
Descriptions, illustrations, interactive identification, and information retrieval from DELTA databases.

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

I. Degani-Schmidt and M. Guerra-Sommer (2019): Epidermal morphology of the cordaitalean leaf Noeggerathiopsis brasiliensis nom. nov. from the southern Paraná Basin (Lower Permian, Rio Bonito Formation) and paleoenvironmental considerations. In PDF, Braz. J. Geol., 49. See also here.

A.B. Doweld (2022): (2889) Proposal to conserve the name Podozamites against Preissleria (fossil Pinophyta: Podozamitales). Free access, Taxon, 71: 484–485.
"... The fossil-generic name Podozamites, so widely used in modern systematic palaeobotany, should not be rejected for purely nomenclatural reasons. In order to stabilize palaeobotanical nomenclature in current use by legitimizing the use of Podozamites, it is formally proposed to conserve Podozamites against the “nomen oblitum”, Preissleria. ..."

! 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 Gymnosperm Database Site Map All Taxa on the Gymnosperm Database.

H.J. Falcon-Lang and A.R. Bashforth (2005): Morphology, anatomy, and upland ecology of large cordaitalean trees from the Middle Pennsylvanian of Newfoundland. PDF file, Review of Palaeobotany and Palynology, 135: 223-243.

Debbie Folkerts, Auburn University, Auburn, Alabama: Kingdom Plantae: Gymnosperms. Powerpoint presentation.
The link is to a version archived by the Internet Archive´s Wayback Machine.

! Y. Fragnière et al. (2015): Fighting their last stand? A global analysis of the distribution and conservation status of gymnosperms. Journal of Biogeography. See also here (studied species and the full list of references from the gymnosperm database).

Anthony G. Futcher, Columbia Union College, Maryland: Plant Diversity. A lot of facts about plant groups, fungi, plant-like protists, and monerans, including taxonomy, life cycles, general structure, and representative genera. Go to:
Gymnosperms.
Still available through the Internet Archive´s Wayback Machine.

Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine: Navigate via: Notes for a Course in Paleobotany. Go to:
Gymnosperms in the Mesophytic.
These expired links are available through the Internet Archive´s Wayback Machine.

J.C. Ghildiyal (ed.; 2019):
Pteridology, Gymnosperms and Palaeobotany. Lecture notes, in PDF, Uttarakhand Open University, India.
With many black and white illustrations.
Palaeobotany part starts on PDF page 254.

! The Gymnosperm Database (edited by Christopher J. Earle). A premier source of information on conifers and their allies. Excellent!
If you have a specific question, use the search box (top of page at right). Alternatively you might also start with the Topics page.
Worth checking out: Essential Links.

! T.M. Harris (1951): The fructification of Czekanowskia and its allies. In PDF, Philos. Transactions of the Royal Society of London, B, 235: 483-508.
See also here.

M. Hrabovský (2021): Leaf evolution and classification. 3. Gymnospermopsida. In PDF, Acta Botanica Universitatis Comenianae, 57.
! Many black and white contour drawings.

W. Huang et al. (2016): New Phoenicopsis leaves (Czekanowskiales) from the Middle Jurassic Daohugou Biota, China and their roles in phytogeographic and paleoclimatic reconstruction. In PDF, Palaeoworld, 25: 388–398. See also here.

K.R. Hultine and J.D. Marshall (2001): A comparison of three methods for determining the stomatal density of pine needles. In PDF, Journal of Experimental Botany, 52: 369–373. See also here.

A.A. Klymiuk et al. (2022): A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms. In PDF, American Journal of Botany. See also here.
Note figure 30: Cupulate Mesozoic gymnosperms.

! Knox College, Galesburg, Illinois: The Seed Plants: Gymnosperms & Angiosperms. Lecture note, Powerpoint presentation.
This expired link is still available through the Internet Archive´s Wayback Machine.

E. Kustatscher et al. (2019): Did the Czekanowskiales already exist in the late Permian? Free access, PalZ.

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.

Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima.
This expired link is now available through the Internet Archive´s Wayback Machine.

A. Linkies et al. (2010): The evolution of seeds. PDF file, New Phytologist.

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

! S. McLoughlin (2021): Gymnosperms: History of Life: Plants: Gymnosperms. In PDF, p. 476-500; In: Elias, S. & Alderton, D. (eds.), Encyclopedia of Geology, Amsterdam, Elsevier. See also here.
Note fig. 10: Reconstruction of the early seed-plant Elkinsia polymorpha (Late Devonian).
Fig. 24A: Reconstruction of Cycadeoidea dacotensis, a cycadeoid bennettite.
Fig. 24C: Reconstruction of Wielandiella angustifolia, a williamsonioid bennettite.
Fig. 28: Reconstruction of the pollen-bearing organ Erdtmanitheca portucalensis (Early Cretaceous).

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, CORDAITES. Link list page with rankings and brief explanations. Images of Cordaites, Cordaites principalis, Cordaicladus, Cordaianthus, Cordaianthus williamsii, Cardiocarpus spinosus, Samaropsis,, Artisia horizontalis, Mesoxylon, Mesoxylon iowense, Amyelon.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

Dennis C. Murphy, ("Devonian Times", a paleontology web site featuring Red Hill): Who's Who at Red Hill, Archaeopteris spp. (Progymnosperm Trees), and The First Forests.

The New York Botanical Garden, Herbarium: Catalog of North American Gymnosperms. The Herbarium of The New York Botanical Garden has completed cataloging its gymnosperms from North America north of Mexico. These records represent those specimens collected in Greenland, Canada, and the United States (excluding Hawaii), and include non-native species cultivated in North America.

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

! Plantillustrations.org (by Max Antheunisse and Jan Koeman).
Plantillustrations.org is a completely non-commercial website. On top you see 2 search boxes at the right. The white one is for entering scientific names, the grey one for vernacular ones.

Ruud J. Poort, Henk Visscher, and David L. Dilcher: Zoidogamy in fossil gymnosperms: The centenary of a concept, with special reference to prepollen of late Paleozoic conifers. Article in PDF format. PNAS Vol. 93, Issue 21, 11713-11717, October 15, 1996.

Ruud J. Poort, Henk Visscher, and David L. Dilcher: Zoidogamy in fossil gymnosperms: The centenary of a concept, with special reference to prepollen of late Paleozoic conifers. The National Academy of Sciences, PNAS 1996 93: 11713-11717.

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.

! 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).

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

Vishal Sharma, Government Post Graduate College For Girls, Chandigarh: Fossils of Gymnosperms. Powerpoint presentation.

K. Stern: Introduction To Seed Plants: Gymnosperms. Powerpoint presentation.

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

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Plant life from the Silurian to the Cretaceous. Go to:
Fossil gymnosperm wood, and
The gymnospermous tree Cordaites.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Plant life from the Silurian to the Cretaceous. Go to:
Little animals in the Coal Swamp.

Biological Sciences, Teaching Development Unit, University of Sydney: Learning Modules. Provided by the Internet Archive´s Wayback Machine. Go to: Pine (Gymnosperm) Life Cycle.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Fossil plant images from the oldest land plants. Go to: Fossil gymnosperm wood.

C. Sun et al. (2015): A new species of Czekanowskia (Czekanowskiales) from from the Middle Jurassic of Inner Mongolia, China. In PDF, Botanica Pacifica, 4: 149–155.

Syllabus of Plant Families, A. Engler´s Syllabus der Pflanzenfamilien (13th edition by Wolfgang Frey):
4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)].

Botany Department, University of Toronto:
Families of Vascular Plants. An introduction to the families of vascular plants with emphasis on morphological variation in relation to identification and significant aspects of breeding systems, dispersal syndromes, and other features of their biology. Go to:
Gymnosperms. Heterospory and seeds.
Still available via Internet Archive Wayback Machine.

Jun Wang and Hermann W. Pfefferkorn (2010): Nystroemiaceae, a new family of Permian gymnosperms from China with an unusual combination of features. PDF file, Proc. R. Soc., B, 277: 301-309. See also here.

J. Wang et al. (2003): Discovery of organic connection of Chiropteris Kurr and Nystroemia Halle from Early Permian of western Henan, China. Abstract, Chinese Science Bulletin, 48: 2248-2252.

K. Unger Baillie (March 12, 2021): ‘Pompeii of prehistoric plants’ unlocks evolutionary secret. Penn Today.

J. Wang et al. (2021): Ancient noeggerathialean reveals the seed plant sister group diversified alongside the primary seed plant radiation: Open access, Proceedings of the National Academy of Sciences, 118, e2013442118.
Note fig. 2: Reconstruction of the aerial parts of Paratingia wuhaia from the early Permian of China.

K.Y. Wang et al. (2022): Anatomically preserved cordaitalean trees from the Pennsylvanian of Yangquan City, Shanxi Province, and their implication for a perhumid climate in North China Block. In PDF, Palaeoworld, 31: 294-310.
See also here.

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

! Wikipedia, the free encyclopedia: Gymnosperm, and the German Wikipedia: Nacktsamige Pflanzen (in German).

! The World List of Cycads (produced under the auspices of the IUCN/SSC Cycad Specialist Group (CSG) and sponsored by Montgomery Botanical Center).
The World List of Cycads is a working list of known cycad species names with the primary goal of providing reliable information on the taxonomy of cycads.
! Don't miss the extensive Cycad Literature List.

Y. Yang et al. (2024): The Systematics and Evolution of Gymnosperms with an Emphasis on a Few Problematic Taxa. Open access, Plants, 13. https://doi.org/10.3390/plants13162196.

B. Zhang et al. (2024): Numerical taxonomy and genus-species identification of Czekanowskiales in China based on machine learning. Free access, Palaeontologia Electronica, 27. https://doi.org/10.26879/1357.
"... accurate identification of Czekanowskiales fossils is difficult due to the similarities in some macroscopic and cuticular patterns among different genera and species
[...] This study focused on the numerical taxonomy and identification of Czekanowskiales at the generic and species levels using cluster analysis, trait selection, and supervised learning methods for machine learning ..."