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! 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. Go to: Ginkgo, Cordaites and the Conifers.

R. Barboni and T.L. Dutra (2015): First record of Ginkgo-related fertile organs (Hamshawvia, Stachyopitys) and leaves (Baiera, Sphenobaiera) in the Triassic of Brazil, Santa Maria formation. In PDF, Journal of South American Earth Sciences, 63: 417-435.

K. Bauer et al. (2014): Ginkgophytes from the upper Permian of the Bletterbach gorge (northern Italy). In PDF, see also here.

K. Bauer et al. (2013): Fossil ginkgophyte seedlings from the Triassic of France resemble modern Ginkgo biloba. In PDF, BMC Evolutionary Biology, 13.

K. Bauer et al. (2013): The ginkgophytes from the German Kupferschiefer (Permian), with considerations on the taxonomic history and use of Baiera and Sphenobaiera. In PDF, Bulletin of Geosciences, 88: 539-556.

D.J. Beerling et al. (1998): Stomatal responses of the "living fossil" Ginkgo biloba L. to changes in atmospheric CO2 concentrations. PDF file, Journal of Experimental Botany, 49: 1603-1607.

! Branko M. Begovic Bego (2011): Nature´s Miracle, Ginkgo biloba L. 1771. In PDF (40.5 MB!). Table of contents PDF page 9-11. See especially: PDF page 49: "Ancestors and relatives of Ginkgo biloba".
See also here. Access via Scribd.

B.M. Begovic Bego et al. (2010/2011): Ginkgo biloba L. 1771 • All about Ginkgo (or Maidenhair tree). In PDF.
This project is for the use of anyone anywhere at no cost and with almost no restrictions whatsoever. Slow download (ca. 41 MB).

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

B. Bomfleur et al. (2014): Habit and Ecology of the Petriellales, an Unusual Group of Seed Plants from the Triassic of Gondwana. Free access, International Journal of Plant Sciences, 175: 1062–1075.

! C.K. Boyce (2008): Seeing the forest with the leaves-clues to canopy placement from leaf fossil size and venation characteristics. In PDF, Geobiology, 7: 192-199.
See also here.

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.

! Curtis Clark, Biological Sciences Department California State Polytechnic University, Pomona: Plant Morphology. Resources, PDF files.
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

! P.R. Crane (2019): An evolutionary and cultural biography of ginkgo. In PDF, Plants, People, Planet, 1: 32–37.

dmoz, the Open Directory Project:
Science: Biology: Flora and Fauna: Plantae:
Ginkgophyta. See also:
Earth Sciences: Paleontology: Paleobotany: Taxa.
These expired links are now available through the Internet Archive´s Wayback Machine.

V-M. Dörken: Ginkgoaceae – Ginkgogewächse (Ginkgoales). PDF file, in German. Konstanz University, FB Biologie.

! 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 Ginkgoales.

M. Eberlein (2015): Bestimmungs- und Verbreitungsatlas der Tertiärflora Sachsens – Angiospermenblätter und Ginkgo. PDF file (in German). Thesis, University of Dresden (in German). First part of a reference book of the Tertiary flora of Saxony.
See also here.

A. Elgorriaga and B.A. Atkinson (2023): Zirabia cylindrica comb. nov. provides evidence of Doyleales in the Jurassic. American Journal of Botany, 110. e16182.
See also here.
"... we report a new genus of Doyleales, Zirabia gen. nov. from the Early Jurassic of Iran, that was originally described as the ginkgophyte Karkenia
[...] Our results indicate that Doyleales is significantly older than previously thought, with their stratigraphic range now extending from the Lower Jurassic to the Cretaceous ..."

T.C. Fischer et al. (2010). Permian Ginkgophyte fossils from the Dolomites resemble extant O-ha-tsuki aberrant leaf-like fructifications of Ginkgo biloba L. PDF file, BMC Evolutionary Biology, 10.

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.

Fossilworks. Fossilworks provides query, download, and analysis tools that utilize the Paleobiology Database's large relational database assembled by hundreds of paleontologists from around the world. Go to:

J.R. Foster et al. (2018): Paleontology, taphonomy, and sedimentology of the Mygatt-Moore Quarry, a large dinosaur bonebed in the Morrison Formation, western Colorado—Implications for Upper Jurassic dinosaur preservation modes. In PDF, Geology of the Intermountain West. See also here and there.

Anja Fuchs, Landshut, Germany: About Ginkgo. PDF file, in German. See also here.

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

Robert A. Gastaldo, Department of Geology, Colby College, Waterville, Maine: Gymnosperms in the Mesophytic.
The link is to a version archived by the Internet Archive´s Wayback Machine.

Google Directory:
Biology > Flora and Fauna > Plantae > Ginkgophyta.
Link directory, with ratings (Google page rank).
Website outdated. The link is to a version archived by the Internet Archive´s Wayback Machine.

K.-J. Gu et al. (2022): GinkgoDB: an ecological genome database for the living fossil, Ginkgo biloba. Free access, Database, 2022: article ID baac046; DOI:

! 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. Haworth and A. Raschi (2014): An assessment of the use of epidermal micro-morphological features to estimate leaf economics of Late Triassic-Early Jurassic fossil Ginkgoales. In PDF, Review of Palaeobotany and Palynology, 205: 1-8.

F. Herrera et al. (2017): The presumed ginkgophyte Umaltolepis has seed-bearing structures resembling those of Peltaspermales and Umkomasiales. PNAS, 114. Freely available online through the PNAS open access option. See also here (in PDF).
Reconstruction of Umaltolepis mongoliensis on PDF page 4.

Jason Hilton (2007): Living Fossils, Ginkgo biloba - its ancestors and allies. Website hosted by The International Organisation of Palaeobotany (IOP).
The link is to a version archived by the Internet Archive´s Wayback Machine.

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.

Z. Jiang et al. (2016): A Jurassic wood providing insights into the earliest step in Ginkgo wood evolution. Sci. Rep., 6.

W. Jung: Der Ginkgo-Baum, ein Unikum mit Vergangenheit. In German.

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

Jirí Kvacek et al. (2005): A new Late Cretaceous ginkgoalean reproductive structure Nehvizdyella gen. nov. from the Czech Republic and its whole-plant reconstruction. Free access, American Journal of Botany, 92: 1958-1969.

Cor Kwant, The Netherlands: The Ginkgo Pages. See especially: Fossils and Ginkgo fossils: picture gallery.
Also worth checking out: Links.

A. Leigh et al. (2011): Structural and hydraulic correlates of heterophylly in Ginkgo biloba. In PDF, New Phytologist, 189: 459-470.
See also here.

L.V. Leiz et al. (2022): New records of Late Triassic wood from Argentina and their biostratigraphic, paleoclimatic, and paleoecological implications. In PDF, Acta Palaeontologica Polonica, 67: 329–340.
See also here.
Note fig. 4: Schemes showing anatomical characters of Baieroxylon cicatricum.
Fig. 7. Schemes showing anatomical characters of Protophyllocladoxylon hilarioense.

! H.Y. Lin et al. (2022): International Biological Flora: Ginkgo biloba. In PDF, Journal of Ecology. See also here.

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 Ginkgophyta.

! S. McLoughlin (2021): Gymnosperms: History of Life: Plants: Gymnosperms. PDF file, in: Elias, S. and Alderton, D. (eds): Encyclopedia of Geology. See also here.

Q.-M. Meng et al. (2019): The natural history of oviposition on a ginkgophyte fruit from the Middle Jurassic of northeastern China. In PDF, Insect Science, 26: 171–179. See also here.

S.V. Meyen (1987): Evolution of Ginkgoopsida: from Peltaspermales to Ginkgoales, Leptostrobales and Caytoniales. In PDF, Bulletin de la Société Botanique de France. Actualités Botaniques, 134: 67-76.

Dominique Mouchel (?), France: Ginkgo biloba.

Dan Nickrent and Karen Renzaglia, Department of Plant Biology, Southern Illinois University at Carbondale: Land Plants Online, Ginkgo - Phylum Ginkgoophyta.
Now recovered from the Internet Archive´s Wayback Machine.

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

! The Paleobiology Database (PBDB).
PBDB is a public database of paleontological data that anyone can use, maintained by an international non-governmental group of paleontologists. The Paleobiology Database has been supported by many grants over the years, mostly from the National Science Foundation. You may navigate from the
Paleobiology Database Guest Menu or check out the
Frequently Asked Questions. Please also note the detailed and excellent tutorial:
! M.D. Uhen et al. (2023): Paleobiology Database User Guide Version 1.0 Free access, PaleoBios, 40: 1-56. See also here (in PDF).

Kathleen B. Pigg, Department of Plant Biology, Arizona State University: Plant Fossils and Evolution (now via wayback link). Go to: Laboratory. The Cycads, Cycadeoids (Bennettitales) and Ginkgophytes.

! (by Max Antheunisse and Jan Koeman). 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.

C. Pott et al. (2016): New Ginkgophytes from the Upper Triassic–Lower Cretaceous of Spitsbergen and Edgeøya (Svalbard, Arctic Norway): The History of Ginkgoales on Svalbard. In PDF, Int. J. Plant Sci., 177: 175–197.

! 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. Quan et al. (2010): A new Tertiary Ginkgo (Ginkgoaceae) from the Wuyun Formation of Jiayin, Heilongjiang, northeastern China and its paleoenvironmental implications. PDF file, American Journal of Botany.

! G.J. Retallack (2002):&xnbsp; Carbon dioxide and climate over the past 300 Myr. In PDF, Phil. Trans. R. Soc. Lond., A, 360: 659–673.
See also here.

Tim Rhodus, Department of Horticulture and Crop Science, Ohio State University: Ginkgo biloba.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens, OH: Vascular Plant Morphology. 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 Ginkgophytes (PDF file).

! D.L. Royer et al. (2003): Ecological conservatism in the "living fossil" Ginkgo. In PDF, Paleobiology, 29: 84–104.
See also here.
"... The living species Ginkgo biloba is phylogenetically isolated, has a relictual distribution, and is morphologically very similar to Mesozoic and Cenozoic congenerics. [...] Ginkgo is an extreme example of a geologically long-lived genus, with its one living species arguably having a temporal range of >100 Myr. ..."

P. Smarda et al. (2016): Polyploidy in a ‘living fossil’ Ginkgo biloba. Open access, New Phytologist, 212: 11–14.

! M. Steinthorsdottir et al. (2022): Key traits of living fossil Ginkgo biloba are highly variable but not influenced by climate – Implications for palaeo-pCO2 reconstructions and climate sensitivity. In PDF, Global and Planetary Change, 211. See also here.

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.

H. Süss and L. Müller (2015): Ein Stamm- und Wurzelholzfossil der Morphogattung Ginkgoxylpropinquus Savidge aus dem Tertiär der sächsischen Braunkohle, mit Bemerkungen über die Stellung der Ginkgoales innerhalb der Gymnospermen aus holzanatomischer Sicht. PDF file, in German. Geologica Saxonica, 60: 451-460.

H. Süss and K.-P. Kelber (2011): Eine neue Art der Morphogattung Baieroxylon Greguss aus dem Keuper von Franken, Deutschland. In PDF, Feddes Repertorium, 122: 257-267.

H. Süss et al. (2009): Drei neue fossile Hölzer der Morphogattung Primoginkgoxylon gen. nov. aus der Trias von Kenia. PDF file (in German), Feddes Repertorium, 120: 273 - 292. See also here (Abstract).

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.

! H. Tralau (1968): Evolutionary trends in the genus Ginkgo. Free access, Lethaia, 1: 63-101.

Alejandro Troncoso (Instituto de Biología Vegetal y Biotecnología, Universidad de Talca, Chile) & Rafael Herbst, (PRINGEPA-CONICET, Corrientes, Argentina): Ginkgoales del Triásico del norte de Chile (in Spain). Rev. geol. Chile, Dec. 1999, vol.26, no. 2.

J. Unverfärth et al. (2022): Sphenobaiera insecta from the Upper Triassic of South Australia, with a clarification of the genus Sphenobaiera (fossil Ginkgophyta) and its delimitation from similar foliage genera. In PDF, Botany Letters, DOI: 10.1080/23818107.2022.2076259.
See also here.

V. Vajda et al. (2021): Geochemical fingerprints of ginkgoales across the triassic-jurassic boundary of greenland. In PDF, Int. J. Plant Sci., 182: 649–662. See also here.
! Note fig. 2, 3: Reconstructions of selected fossil ginkgoalean taxa.

! V. Vajda et al. (2017): Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships. In PDF, Nature Ecology & Evolution. See also here and there.

L. Villar de Seoane et al. (2015): Ginkgoites patagonica (Berry) comb. nov. from the Eocene of Patagonia, Last Ginkgoalean Record in South America. In PDF, International Journal of Plant Sciences, 176: 346-363. See also here and there.

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.

Y. Wang et al. (2012): Jurassic mimicry between a hangingfly and a ginkgo from China. In PDF, Proc. Nat. Acad. Sci. USA, 109: 20514-20519.

Y. Wang et al. (2005): Cuticular anatomy of Sphenobaiera huangii (Ginkgoales) from the lower Jurassic of Hubei, China. In PDF, American Journal of Botany, 92: 709-721.

Z. Wang et al. (2017): A New Species of Ginkgo with Male Cones and Pollen Grains in situ from the Middle Jurassic of Eastern Xinjiang, China. Abstract, Acta Geologica Sinica.

! Wikipedia (a free-content encyclopedia): Spermatophyte. Go to: Ginkgo.

J.P. Wilson and A.H. Knoll (2010): A physiologically explicit morphospace for tracheid-based water transport in modern and extinct seed plants. PDF file, Paleobiology, 36: 335-355.
See also here.

SanPing XIE et al. (2009): Altitudinal variation in Ginkgo leaf characters: Clues to paleoelevation reconstruction. PDF file, Science in China Series D: Earth Sciences, 52: 2040-2046.
"The results show that leaf area, petiole length, and stomatal parameters have no obvious linear relationship with altitude (...). The results also suggest that the differences in stomatal density and stomatal index between sun and shade leaves had more influence on paleoelevation reconstruction than that in other parameters".

C. Yiotis et al. (2017): Differences in the photosynthetic plasticity of ferns and Ginkgo grown in experimentally controlled low [O2]:[CO2] atmospheres may explain their contrasting ecological fate across the Triassic–Jurassic mass extinction boundary. Free access, Annals of Botany, 119: 1385–1395.

Y. Xu et al. (2023): How similar are the venation and cuticular characters of Glossopteris, Sagenopteris and Anthrophyopsis? In PDF, Review of Palaeobotany and Palynology, 316.
See likewise here.
Note figure 1: Geologic ranges of some representative reticulate taxa.
"... Considering the putatively close relationship of glossopterids (Glossopteris), Caytoniales (Sagenopteris) and Bennettitales (here encompassing Anthrophyopsis) resolved as members of the ‘glossophyte’ clade in some past phylogenetic studies, cuticular features suggest that these groups are not closely related. In addition, anastomosing venation, superficially similar to that of Glossopteris, Sagenopteris and Anthrophyopsis appears to have arisen independently in numerous other plant groups ..."

B. Zhang et al. (2024): Numerical taxonomy and genus-species identification of Czekanowskiales in China based on machine learning. Free access, Palaeontologia Electronica, 27.
"... 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 ..."

! Y.P. Zhao et al. (2019): Resequencing 545 ginkgo genomes across the world reveals the evolutionary history of the living fossil. Open access, Nature Communications, 10.
"... investigations provide insights into the evolutionary history of ginkgo trees and valuable genomic resources for further addressing various questions involving living fossil species.
[...] It is likely that the morphological stasis of living fossils is an effective adaptation strategy in response to environmental change, although the underlying mechanisms are unclear ..."

Z. Zhiyan and W. Xiangwu (2006): The rise of ginkgoalean plants in the early Mesozoic: a data analysis. Abstract, Geo. J., 41: 363-375.

! Z.-Y. Zhou (2009): An overview of fossil Ginkgoales. Abstract, Palaeoworld, 18: 1-22.

! Zhiyan Zhou (website hosted by International Organisation of Palaeobotany): Gingko biloba: its ancestors and allies.
Now recovered from the Internet Archive´s Wayback Machine.

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