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H.M. Anderson et al. (2008): Stems with attached Dicroidium leaves from the Ipswich Coal Measures, Queensland, Australia. PDF file, Memoirs of the Queensland Museum 52: 1-12. Snapshot taken by the Internet Archive´s Wayback Machine.

E. Antevs (1914): Lepidopteris ottonis (GÖPP.) SCHIMP. und Antholithus zeilleri NATHORST. Kungl. Svenska Vetenskapsakademiens Handlingar Ny Följd, 51: 1-18. Uppsala & Stockholm.

S. Archangelsky (1968): Studies on Triassic fossil plants from Argentina. IV. The leaf genus Dicroidium and its possible relation to Rhexoxylon stems. PDF file, Palaeontology, 11: 500-512.
A version archived by Internet Archive Wayback Machine.

E. Artabe et al. (1999): Rhexoxylon brunoi Artabe, Brea et Zamuner, sp. nov., a new Triassic corystosperm from the Paramillo de Uspallata, Mendoza, Argentina. In PDF, Rev. Palaeobot. Palynol., 105: 63–74.

Brian J. Axsmith et al. (2007): The "New Approach to Corystospermales" and the Antarctic Fossil Record: A Critique. Ameghiniana, 44. See also here (PDF file).

B.J. Axsmith et al. (2000): New perspectives on the Mesozoic seed fern order Corystospermales based on attached organs from the Triassic of Antarctica. Free access, American Journal of Botany, 87: 757-768.
Note Fig. 21: Reconstruction of an Umkomasia uniramia cupulate organ.

M. Backer et al. (2019): Frond morphology and epidermal anatomy of Compsopteris wongii (T. Halle) Zalessky from the Permian of Shanxi, China. Open access, PalZ.

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

B. Bomfleur et al. (2018): Polar Regions of the Mesozoic-Paleogene Greenhouse World as Refugia for Relict Plant Groups. Chapter 24, 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.
Note figure 24.2: Distribution of Dicroidium through space and time.

B. Bomfleur et al. (2012): Modified basal elements in Dicroidium fronds (Corystospermales). In PDF, Review of Palaeobotany and Palynology, 170: 15-26.

B. Bomfleur et al. (2011): Systematics and paleoecology of a new peltaspermalean seed fern from the Triassic polar vegetation of Gondwana. In PDF, International Journal of Plant Sciences, 172: 807-835.

L.D. Boucher et al. (1995): Dicroidium compression floras from southern Victoria Land. PDF file, Antarctic Journal, 41.

Philippe Choler, Laboratoire de Biologie des Populations d'Altitude, Université Joseph Fourier, Grenoble: Biologie Evolutive Végétale. Concepts and methods in evolutionary biology (in French).
Navigate from "Plan du cours" (access to about 335 slides).
Go to: Corystospermaceae, or Caytoniales.
These expired links are available through the Internet Archive´s Wayback Machine.

C.J. Cleal et al. (2015): Pennsylvanian fossil flora from the Velebit Mountains and Lika region (SW Croatia). In PDF, Bulletin of Geosciences 90: 721-742.

José Alejandro D´angelo (2006): Analysis by Fourier transform infrared spectroscopy of Johnstonia (Corystospermales, Corystospermaceae) cuticles and compressions from the Triassic of Cacheuta, Mendoza, Argentina. Ameghiniana, 43.

! W.A. DiMichele et al. (2006): Paleoecology of Late Paleozoic pteridosperms from tropical Euramerica. In PDF, The Journal of the Torrey Botanical Society, 133: 83-118. See also here.

W.A. DiMichele et al. (2005): THE PERMIAN PELTASPERM RADIATION: EVIDENCE FROM THE SOUTHWESTERN UNITED STATES. PDF file, p. 67-79, In: Lucas, S.G. and Zeigler, K.E., (eds.): The Nonmarine Permian, New Mexico Museum of Natural History and Science Bulletin No. 30.

! A.B. Doweld (2012): Proposals to conserve the name Scytophyllum Bornem. (fossil Pteridospermae, Peltaspermopsida) against Scytophyllum Eckl. & Zeyh. (Celastraceae) and the name S. bergeri with a conserved type. In PDF, Taxon, 61: 1128-1129.

! A. Elgorriaga et al. (2019): Relictual Lepidopteris (Peltaspermales) from the Early Jurassic Cañadón Asfalto Formation, Patagonia, Argentina. Abstract, Int. J. Plant Sci., 180. See also here (in PDF), and there.
"... and its youngest species, Lepidopteris ottonis, has been used as a Rhaetian marker for several European, Greenlandic, and American localities ..."
"... Lepidopteris scassoi represents the youngest occurrence of the genus by more than 20 Myr. Lepidopteris and Dicroidium lineages, dominant in Southern Hemisphere Triassic ecosystems, show a similar overall pattern of origination (Late Permian), diversification (late Early-Middle Triassic), and decline (Late Triassic), with relict occurrences during the Early Jurassic. ..."

A. Elgorriaga et al. (2019): Southern Hemisphere Caytoniales: vegetative and reproductive remains from the Lonco Trapial Formation (Lower Jurassic), Patagonia. Open access, Journal of Systematic Palaeontology, DOI: 10.1080/14772019.2018.1535456

A. Hamad et al. (2008): A Late Permian flora with Dicroidium from the Dead Sea region, Jordan. In PDF, Review of Palaeobotany & Palynology 149: 85-130.

! T.M. Harris (1933): A new member of the Caytoniales. New Phytologist, 32: 97–114. In PDF, See also here (abstract).

X. He et al. (2017): Peltaspermalean seed ferns with preserved cuticle from the Upper Triassic Karamay Formation in the Junggar Basin, northwestern China. Abstract, Review of Palaeobotany and Palynology, 247: 68-82. 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.

! J. Hilton and R.M. Bateman (2006): Pteridosperms are the backbone of seed-plant phylogeny. In PDF, Journal of the Torrey Botanical Society, 133: 119-168.

E.V. Karasev (2009): A New Genus Navipelta (Peltaspermales, Pteridospermae) from the Permian/Triassic Boundary Deposits of the Moscow Syneclise. PDF file, Paleontological Journal, 43: 1262-1271.

H. Kerp et al. (2006): Typical Triassic Gondwanan floral elements in the Upper Permian of the paleotropics. In PDF.

Hans Kerp, Abdallah Abu Hamad, Klaus Bandel & Birgit Niemann: A new Upper Permian flora from the Middle East with typical Triassic Gondwana elements. Abstract, The 15th Plant Taphonomy Meeting, Naturalis, National Museum of Natural History, Leiden, The Netherlands, 12-13th November 2004. Provided by the Internet Archive´s Wayback Machine.

Sharon D. Klavins et al. (2002): Anatomy of Umkomasia (Corystospermales) from the Triassic of Antarctica. American Journal of Botany, 89: 664-676. See also here. Abstract, Botany 2001, August 12 - 16, 2001; Albuquerque, New Mexico.

Michael Krings et al. (2006): Frond architecture of Odontopteris brardii (Pteridospermopsida, ?Medullosales): new evidence from the Upper Pennsylvanian of Missouri, U.S.A. Journal of the Torrey Botanical Society, 133: 33-45.

! E. Kustatscher and J.H.A. van Konijnenburg-van Cittert (2013): Seed ferns from the European Triassic - an overview. In PDF, In: Tanner, L.H., Spielmann, J.A. and Lucas, S.G., (eds.): The Triassic System. New Mexico Museum of Natural History and Science, Bulletin 61.
See also here from

E. Kustatscher et al. (2011): Scytophyllum waehneri (Stur) nov. comb., the correct name for Scytophyllum persicum (Schenk) Kilpper, 1975. In PDF, Zitteliana, A 51.

! T. Linnell (1933): Zur Morphologie und Systematik triassischer Cycadophyta, II. Über Scytophyllum Bornem., eine wenig bekannte Cycadophyten-Gattung aus dem Keuper. Svensk Botanisk Tidskrift, 27: 310-331. See also here.

C. Mays and S. McLoughlin (2019): Caught between mass extinctions - the rise and fall of Dicroidium. In PDF.

S. McLoughlin et al. (2018): Pachytestopsis tayloriorum gen. et sp. nov., an anatomically preserved glossopterid seed from the Lopingian of Queensland, Australia. Chapter 9, 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.

S. McLoughlin (2017): Antarctica’s Glossopteris forests. In PDF, In: 52 More Things You Should Know About Palaeontology,eds. A. Cullum, A.W. Martinius. Nova Scotia: Agile Libre, p. 22-23. See also here.

S. McLoughlin (2011): Glossopteris - insights into the architecture and relationships of an iconic Permian Gondwanan plant. In PDF, J. Botan. Soc. Bengal 65: 1-14.

Stephen McLoughlin et al. (2008): Seed ferns survived the end-Cretaceous mass extinction in Tasmania. PDF file, American Journal of Botany, 95: 465-471.

B. Meyer-Berthaud et al. (1993): Petrified Stems Bearing Dicroidium Leaves from the Triassic of Antarctica. In PDF, Palaeontology, 36.

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

V. Mosbrugger, Institut für Geologie und Paläontologie, Eberhard-Karls-Universität Tübingen: Lecture notes about plant palaeobiology. PDF files, in German. Provided by the Internet Archive´s Wayback Machine. Go to:
Introduction, Progymnosperms (4 MB),
Wood, Seed evolution (1.2 MB),
Pteridosperms I, Lyginopterids (1.2 MB),
Pteridosperms II, Medullosans (2.1 MB),
Glossopterids, Caytoniales (1.3 MB),

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, PTERIDOSPERMS OR SEED FERNS. Link list page with rankings and brief explanations. Images of Lyginopteris, Lagenostoma, Sphenopteris crepinii, Lagenostoma lomaxii, Pachytesta, Trigonocarpus, Bernaultia, Schopfipollenites, Bernaultia formosa, Whittleseya microphylla, Medullosa noei, Myeloxyleon, Neuropteris, Alethopteris, Sphenopteris, Alethopteris decurrens, Alethopteris lonchitica, Alethopteris sullivantii, Neuropteris obliqua, Eusphenopteris, Mariopteris, Mariopteris muricata, Reticulopteris, Cyclopteris, Lescuropteris genuina, Alethopteris zeilleri.
Still available via Internet Archive Wayback Machine.

! Palaeobotanical Research Group, Münster, Westfälische Wilhelms University, Münster, Germany. History of Palaeozoic Forests, CALLIPTERIDS. Link list page with rankings and brief explanations. Images of Autunia conferta, Rhachiphyllum schenkii.
This expired link is available through the Internet Archive´s Wayback Machine.

Dennis C. Murphy, ("Devonian Times", a paleontology web site featuring Red Hill): Who's Who at Red Hill, Early Seed Plants (lyginopterids).

! S.V. Naugolnykh (2012): Vetlugospermum and Vetlugospermaceae: A new genus and family of peltasperms from the Lower Triassic of Moscow syneclise (Russia). In PDF, Geobios, 45: 451-462.
Embedment of plant remains in block-diagram reconstructions!

H. Nishida, K.B. Pigg and J. F. Rigby, Swimming sperm in an extinct Gondwanan plant. Glossopteris´ simple mode of reproduction. PDF file, Nature, 422: 396-397; 2003.

G.A. Pattemore (2016): The structure of umkomasiacean fructifications from the Triassic of Queensland. Acta Palaeobotanica, 56: 17–40. See also here.

G.A. Pattemore et al. (2015): Triassic-Jurassic pteridosperms of Australasia: speciation, diversity and decline. In PDF, Boletín Geológico y Minero, 126: 689-722.

G.A. Pattemore et al. (2015): The Mesozoic megafossil genus Linguifolium Arber 1917. In PDF, Acta Palaeobotanica, 55: 123-147.

G.A. Pattemore et al. (2014): Palissya: A global review and reassessment of Eastern Gondwanan material. In PDF, Review of Palaeobotany and Palynology, 210: 50-61. See also here.

Kathleen B. Pigg, Department of Plant Biology, Arizona State University: Plant Fossils and Evolution. Go to: Laboratory 11. Paleozoic Seed Ferns, Cordaites & Early Conifers, Gondwana groups.

K.P. Pigg and S. McLoughlin (1997): Anatomically preserved Glossopteris leaves from the Bowen and Sydney Basins, Australia. PDF file, Rev. Palaeobot. Palynol. 97: 339-359.

R. Prevec (2011): A structural re-interpretation and revision of the type material of the glossopterid ovuliferous fructification Scutum from South Africa. In PDF, Palaeont. afr., 46: 1–19.
See also here and there (abstract).
Please take notice of the sketch in fig 3 on PDF page 6, showing depressed seed scars of the apical portion of a Scutum leslii fructification.

Rosemary Prevec, Geology Department Rhodes University, South Africa (website by Science in Africa): The power of plants: how ancient forests drive SA´s economy. About Glossopteris forests and coal. This expired link is available through the Internet Archive´s Wayback Machine.

R. Prevec et al. (2009): Portrait of a Gondwanan ecosystem: A new late Permian fossil locality from KwaZulu-Natal, South Africa. Abstract, Review of Palaeobotany and Palynology, 156: 454-493. See also here (PDF file). About Glossopteris.

G.J. Retallack (2002): Lepidopteris callipteroides, an earliest Triassic seed fern of the Sydney Basin, southeastern Australia. In PDF, Alcheringa, Alcheringa 26:475–500.

! G.J. Retallack and D.L. Dilcher (1988): Reconstructions of Selected Seed Ferns. In PDF, Annals of the Missouri Botanical Garden. 75: 1010–1057. See also here.
! Note fig. 1: Reconstructions of Stamnostoma huttense.
! Note fig. 3: Reconstructions of Lyrasperma scotia.
! Note fig. 4: Reconstructions of Calathospermum fimbriatum.
! Note fig. 5: Reconstructions of Lagenostoma lomaxii.
! Note fig. 6: Reconstructions of Pachytesta illionensis.
! Note fig. 7: Reconstructions of Callospermanion pusillum.
! Note fig. 8: Reconstructions of Dictyopteridium sporiferum.
! Note fig. 9: Reconstructions of Peltaspermum thomasii, Triassic.
! Note fig. 10: Reconstructions of Umkomasia cranulata, Triassic.
! Note fig. 11: Reconstructions of Caytonia nathorstii.

G. Shi et al. (2019): Diversity and homologies of corystosperm seed-bearing structures from the Early Cretaceous of Mongolia. Abstract, See also here (in PDF).
Note figure 12: Reconstruction of a shoot of Umkomasia mongolica.
Note figure 13: Reconstructions of the seed-bearing units of Umkomasia mongolica, Umkomasia corniculata and Umkomasia trilobata.

! G. Shi et al. (2016): Early Cretaceous Umkomasia from Mongolia: implications for homology of corystosperm cupules. In PDF, New Phytologist, 210: 1418–1429. See also here.

M. Slodownik et al. (2021): Fossil seed fern Lepidopteris ottonis from Sweden records increasing CO2 concentration during the end-Triassic extinction event. Open access, Palaeogeography, Palaeoclimatology, Palaeoecology, 564. See also here (in PDF).

D. Soltis et al. (2017): Phylogeny and Evolution of the Angiosperms. Book announcement. See also here (Google books). Worth checking out:
! Relationships of Angiosperms to Other Seed Plants. In PDF.
Note figure 1.12: Reconstructions of Caytoniales.
Note figure 1.13: Reconstruction of Bennettitales.
Note figure 1.14: Reconstructions of Pentoxylon plants.
Note figure 1.15: Reconstructions of glossopterids.

A.K. Srivastava and R. Srivastava (2016): Glossopteridales: An intricate group of plants. In PDF, The Palaeobotanist, 65: 159–167.

A.K. Srivastava et al. (2010): Dicroidium: no more a Triassic Gondwana index fossil. PDF file.

Hans Steur, Ellecom, The Netherlands: Hans´ Paleobotany Pages. Plant life in the Silurian, Devonian, Carboniferous, Permian and Cretaceous. Go to: Seed ferns from the Piesberg and Ibbenbüren. See also: The Permian flora of Lodève (France), The seed ferns.

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.

Edith L. Taylor (1996): Enigmatic gymnosperms? Structurally preserved Permian and Triassic seed ferns from Antarctica. PDF file, Review of Palaeobotany and Palynology. See also here (abstract).

Edith L. Taylor and Thomas N. Taylor (2009): Seed ferns from the late Paleozoic and Mesozoic: Any angiosperm ancestors lurking there? PDF file, American Journal of Botany, 96: 237-251. See also here.

Edith L. Taylor et al. (2006): Mesozoic seed ferns: Old paradigms, new discoveries. PDF file, Journal of the Torrey Botanical Society, 133: 62-82. Provided by the Internet Archive´s Wayback Machine.

R. Tewari et al. (2017): The Glossopteris flora of Manuguru Area, Godavari Graben, Telangana, India. In PDF, Palaeobotanist, 66: 17–36.

R. Tewari et al. (2015): Glossopteris flora in the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica: Implications for paleogeography, paleoclimatology, and biostratigraphic correlation. Abstract, GR Focus Review, Gondwana Research, 28: 905-932. See also here (in PDF).

! H.H. Thomas(1933): On some pteridospermous plants from the Mesozoic rocks of South Africa. Open access, Philosophical Transactions of the Royal Society, B. 222: 193–265.

! John A. Townrow (1966): The Peltaspermaceae, a pteridosperm family of Permian and Triassic age. PDF file, Palaeontology, 3: 333–361.

M. Wan et al. (2016): A typical Euramerican floral element from the Shanxi Formation (Cisuralian, lower Permian) in the Wuda Coal Field, Inner Mongolia, North China. Palaeobiodiversity and Palaeoenvironments, 96: 507–515.

WANG Jun, H.W. Pfefferkorn, SUN Bainian & LIU Lujun: Discovery of organic connection of Chiropteris Kurr and Nystroemia Halle from Early Permian of western Henan, China. PDF file (32 MB !), Chinese Science Bulletin, 2003, Vol. 48, No. 20, p. 2248-2252.

Wikipedia, the free encyclopedia:
! Pteridospermatophyta.
Samenfarne (in German).
! Caytoniales.
Caytoniales (in German).

N. Zavialova and J.H.A. van Konijnenburg-van Cittert (2011): Exine ultrastructure of in situ peltasperm pollen from the Rhaetian of Germany and its implications. Abstract, Review of Palaeobotany and Palynology, 168: 7–20. See also here (in PDF).

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Last updated March 12, 2021

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