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

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Annette Becker et al. (2002): Ancestry and diversity of BEL1-like homeobox genes revealed by gymnosperm (Gnetum gnemon) homologs. PDF file, Dev Genes Evol, 12: 452-457.
Now provided by the Internet Archive´s Wayback Machine.
See also here (in German).

! 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: 40924097. See also here.

Gregory T. Chandler, The Australian National University:
Introduction to Taxonomy. Go to:
Gnetophyta (in PDF).

Shu-Miaw Chaw et al. (2000): Seed plant phylogeny inferred from all three plant genomes: Monophyly of extant gymnosperms and origin of Gnetales from conifers. PDF file, PNAS, 97. See also here.

! dmoz, the Open Directory Project:
Science: Biology: Flora and Fauna: Plantae:
Gnetophyta. See also:
Earth Sciences: Paleontology: Paleobotany: Taxa. This expired link is now available through the Internet Archive´s Wayback Machine.

M.J. Donoghue and J.A. Doyle (2000): Seed plant phylogeny: Demise of the anthophyte hypothesis?. Free access, Current Biology, 10: R106-R109.
See also here.
"... Recent molecular phylogenetic studies indicate, surprisingly, that Gnetales are related to conifers, or even derived from them ..."

! J.A. Doyle (1998): Molecules, morphology, fossils, and the relationship of angiosperms and Gnetales. In PDF, Molecular phylogenetics and evolution.

! 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 Gnetales.
These expired links are now available through the Internet Archive´s Wayback Machine.

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.

! W.E. Friedman (1996): Introduction to biology and evolution of the Gnetales. International Journal of Plant Sciences 157(6) (supplement): 1-2.

! E.M. Friis et al. (2013): New Diversity among Chlamydospermous Seeds from the Early Cretaceous of Portugal and North America. Free accesss, International Journal of Plant Sciences, 174: 530558.
"... The material is based on numerous charcoalified and lignitic specimens recovered from Early Cretaceous mesofossil floras [...]
! Attenuation-based synchrotron-radiation x-ray tomographic microscopy (SRXTM) and phase-contrast x-ray tomographic microscopy (PCXTM) were carried out [...]
! Volume rendering (voltex), which provides transparent reconstructions, was also used for the virtual sections ..."

! E.M. Friis et al. (2011): Early Flowers and Angiosperm Evolution. Abstract, Cambridge University Press.
See also here (in PDF, long download time) and there (Google books). See especially chapter 5: Angiosperms in context: extant and fossil seed plants
Also worth to check out: Book Review, by P.J. Rudall, Botanical Journal of the Linnean Society, 170. In PDF.
"... This long-awaited book represents not only a remarkable tour de force of palaeobotanical literature, but also a potentially enduring biological textbook. ..."

Else Marie Friis et al. (2007): Phase-contrast X-ray microtomography links Cretaceous seeds with Gnetales and Bennettitales. Abstract, Nature 450: 549-552.
! See also here (in PDF).

Thorsten Hebben, Department of Biological Sciences, University of Alberta, Edmonton, Canada: Gnetophyta: An Enigmatic Group of Seed Plants. A slideshow.
Now recovered from the Internet Archive´s Wayback Machine.

! S.M. Ickert-Bond and S.S. Renner (2016): The Gnetales: Recent insights on their morphology, reproductive biology, chromosome numbers, biogeography, and divergence times. In PDF, Journal of Systematics and Evolution.

N. Jürgens et al. (2021): Welwitschia: Phylogeography of a living fossil, diversified within a desert refuge. Open access, Scientific Reports, 11.
"... With a fossil record of 112 My and phylogenetically isolated within the order Gnetales, the monotypic genus Welwitschia has survived only in the northern Namib Desert in Angola and Namibia. Despite its iconic role, the biogeography, ecological niche, and evolutionary history of the species remain poorly understood. Here we present the first comprehensive map of the strongly disjunct species range, and we explore the genetic relationships among all range fragments based on six SSR markers. ..."

V.A. Krassilov (2009): Diversity of Mesozoic Gnetophytes and the First Angiosperms. PDF file, Paleontological Journal, 43: 1272-1280. This expired link is available through the Internet Archive´s Wayback Machine.

L. Kunzmann et al. (2011): A putative gnetalean gymnosperm Cariria orbiculiconiformis gen. nov. et spec. nov. from the Early Cretaceous of northern Gondwana. In PDF, Review of Palaeobotany and Palynology, 165: 7595. 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 Gnetophyta (Vessel-bearing gymnosperms). Ephedra, Gnetum, Welwitschia.
These expired links are now available through the Internet Archive´s Wayback Machine.

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

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

Dan Nickrent and Karen Renzaglia, Department of Plant Biology Southern Illinois University at Carbondale: Land Plants Online, Phylum Gnetophyta. The Gnetophytes, list of families and genera.
This expired link is available through the Internet Archive´s Wayback Machine.

! J.-H. Ran et al. (2018): Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms. Abstract.

Catarina Rydin, Department of Palaeobotany, Swedish Museum of Natural History, Stockholm: Phylogeny and evolution of Gnetales. A research project.
This expired link is available through the Internet Archive´s Wayback Machine.

! R.A. Stockey, Department of Biological Sciences, University of Alberta: PALEOBIOLOGY OF ANGIOSPERM ORIGINS. This course explores recent advancements toward resolution of the evolutionary origin of flowering plants. Go to:
Presentation on Gnetophyta: An Enigmatic Group of Seed Plants (Author: Thorsten Hebben).
Websites outdated. Links provided by the Internet Archive´s Wayback Machine.

X. Wang and S. Zheng (2010): Whole fossil plants of Ephedra and their implications on the morphology, ecology and evolution of Ephedraceae (Gnetales). Anstract, Chinese Science Bulletin, 55: 1511-1519.

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

Y. Yang (2014): A systematic classification of Ephedraceae: living and fossil. In PDF, Phytotaxa, 158: 283-290.

Y. Yang (2010): A Review on Gnetalean Megafossils: Problems and Perspectives. In PDF, Taiwania, 55: 346-354.

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Last updated May 14, 2022

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