Paleontological Journal

, Volume 43, Issue 10, pp 1272–1280

Diversity of Mesozoic gnetophytes and the first angiosperms



A provisionary system of gnetophytes is proposed, including recently discovered Mesozoic members. The following principal characters are used, in descending order of significance: strobilar structure of ovulate organs (eugnetophytes) as opposed to flowerlike cupules and their aggregates (anthognetophytes), radial vs. bilateral symmetry of seed cupules, and morphological distinctions in the pollen organs, pollen micromorphology and exine ultrastructure. The fossil members show mosaic associations of morphological traits, which attests to different evolution rates of their ovulate and pollinate structures. The system includes two superorders and eight orders, several of which are new. A dichotomous key is provided to differentiate between genera of fossil gnetophytes based on ovulate structures. Two stages in evolution of gnetophyte morphological diversity are recognized: first in the Triassic and then in the Early Cretaceous, the latter was associated with the origin of new type herbaceous wetland communities. Angiosperms or some of them might have originated as elements of gnetophyte diversity in such communities.

Key words

gnetophytes Mesozoic plants plant morphology and taxonomy origin of angiosperms 


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  1. 1.
    M. A. Akhmetiev and V. A. Krassilov, “Recollected Proangiosperms and Correlation of Upper Mesozoic Lacustrine Deposits in East Asia,” Stratigr. Geol. Korrelyatsiya 10(4), 105–109 (2002) [Stratigr. Geol. Correlation 10 (4), 414–418 (2002)].Google Scholar
  2. 2.
    J. M. Anderson and H. M. Anderson, Heyday of the Gymnosperms: Systematics and Diversity of the Late Triassic Molteno Fructifications: Strelitzia 15 (Natl. Bot. Inst., Pretoria, 2003).Google Scholar
  3. 3.
    J. M. Anderson, Anderson M. H., and C. J. Cleal, Brief History of the Gymnosperms: Classification, Biodiversity, Phytogeography and Ecology: Strelitzia 20 (Natl. Bot. Inst., Pretoria, 2007).Google Scholar
  4. 4.
    S. Ash, “Dinophyton, a Problematic New Plant Genus from the Upper Triassic of the South-western United States,” Paleontology 13(4), 646–663 (1970).Google Scholar
  5. 5.
    M. Barbacka, J. Ziala, E. Wcislo-Luraniec, and M. Reymanówna, “Hirmeriella muensteri (Schenk) Jung from Odrow (Poland), with Female and Male Cones, and In Situ Classopolles Pollen Grains,” Acta Palaeobot. 47(2), 339–357 (2007).Google Scholar
  6. 6.
    G. Cladera, Del G. M. Fueyo, L. Villar de Seoane, and S. Archangelsky, “Early Cretaceous Riparian Vegetation in Patagonia, Argentina,” Rev. Mus. Argentino Scienc. Nat. N. S. 91(1), 49–58 (2007).Google Scholar
  7. 7.
    C. J. Cleal, “Gymnospermophyta,” in The Fossil Record, Ed. by M. J. Benton (Capman Hall, London, 1993), Vol. 2, pp. 165–194.Google Scholar
  8. 8.
    J. A. Clement-Westerhof and J. H. A. van Konijnenburgvan Cittert, “Hirmeriella muensteri. New Data on the Fertile Organs Leading to a Revised Concept of the Cheirolepidiaceae,” Rev. Palaeobot. Palynol. 68(1–2), 147–179 (1991).CrossRefGoogle Scholar
  9. 9.
    B. Cornet, “The Reproductive Structures and Leaf Venation of a Late Triassic Angiosperm, Sanmiguelia lewisii,” Evol. Theory 7, 231–309 (1986).Google Scholar
  10. 10.
    B. Cornet, “Late Triassic Angiosperm-like Pollen from the Richmond Rift Basin of Virginia, U.S.A.,” Palaeontogr. Abt. B 213, 37–87 (1989).Google Scholar
  11. 11.
    B. Cornet, “Dicot-like Leaf and Flowers from the Late Triassic Tropical Newarc Supergroup Rift Zone, U.S.A.,” Modern Geol. 3, 81–99 (1993).Google Scholar
  12. 12.
    B. Cornet, “A New Gnetophyte from the Late Carnian (Late Triassic) of Texas and Its Bearing on the Origin of the Angiosperm Carpel and Stamen,” in Flowering Plant Origin, Evolution and Phylogeny. Chapter 3, Ed. by D. W. Taylor and L. J. Hickey (Chapman and Hall, London, 1996), pp. 32–67.CrossRefGoogle Scholar
  13. 13.
    P. R. Crane and R. Upchurch Jr., “Drewria potomacensis gen. et sp. nov., an Early Cretaceous Member of Gnetales from the Potomac Group of Virginia,” Am. J. Bot. 74, 1722–1736 (1987).CrossRefGoogle Scholar
  14. 14.
    D. L. Dilcher, V. A. Krassilov, and J. G. Douglas, “Angiosperm Evolution: Fruits with Affinities to Ceratophyllales from the Lower Cretaceous,” in Abstr. 5th Conf. Int. Org. Palaeobot., Santa Barbara (1996), p. 23.Google Scholar
  15. 15.
    D. L. Dilcher, M. E. Bernerdes-de-Oliveira, D. Pons, and T. A. Lott, “Welwitschiaceae from the Lower Cretaceous of Northeastern Brazil,” Am. J. Bot. 92, 1294–1310 (2005).CrossRefGoogle Scholar
  16. 16.
    J. G. Douglas, “The Mesozoic Flora of Victoria, pts. 1, 2,” Mem. Geol. Surv. Vic. 28, 1–310 (1969).Google Scholar
  17. 17.
    B. Gomez, V. Daviero-Gomez, C. Martin-Glossas, and M. Fuente, “Montsechia vidali, an Early Aquatic Angiosperm from the Barremian of Spain,” in Abstr. 7th European Palaeobot. Palynol. Conf. Prague (2006), p. 49.Google Scholar
  18. 18.
    O. Heer, “Beiträge zur Jura-Flora Ostsibiriens und des Amurlandes,” Mém. Acad. Sci. St. Petersbourg 22, 1–122 (1876).Google Scholar
  19. 19.
    N. F. Hughes and A. B. McDougal, “Record of Angiospemoid Pollen Entry into the English Early Cretaceous Succession,” Rev. Palaeobot. Palynol. 50(3), 255–272 (1987).CrossRefGoogle Scholar
  20. 20.
    V. A. Krassilov, “On the Ovuliferous Organ of Hirmerella,” Phyta (Pant Commemorative Volume), 141–144 (1982a).Google Scholar
  21. 21.
    V. A. Krassilov, “Early Cretaceous Flora of Mongolia,” Palaeontogr. Abt. B 208(1–3), 1–43 (1982b).Google Scholar
  22. 22.
    V. A. Krassilov, “New Floral Structure from the Lower Cretaceous of Lake Baikal Area,” Rev. Palaeobot. Palynol. 47, 9–16 (1986).CrossRefGoogle Scholar
  23. 23.
    V. A. Krassilov, “Palaeobotany of the Mesophyticum: State of the Art,” Rev. Palaeobot. Palynol. 50, 231–254 (1987).CrossRefGoogle Scholar
  24. 24.
    V. A. Krassilov, Angiosperm Origins: Morphological and Ecological Aspects (Pensoft, Sophia, 1997).Google Scholar
  25. 25.
    V. A. Krassilov, “Order Eoanthales Krassilov, And. et And. nov.,” in J. M. Anderson, H. M. Anderson, and C. J. Cleal Brief History of the Gymnosperms: Classification, Biodiversity, Phytogeography and Ecology: Strelitzia 20 (Natl. Bot. Inst., Pretoria, 2007), p. 209.Google Scholar
  26. 26.
    V. A. Krassilov and S. R. Ash, “On Dinophyton—Protognetalean Mesozoic Plant,” Palaeontogr. Abt. B 208(1–3), 33–38 (1988).Google Scholar
  27. 27.
    V. A. Krassilov and S. R. Ash, “Order Dinophytonales Krassilov et Ash nov.,” in J. M. Anderson, H. M. Anderson, and C. J. Cleal Brief History of the Gymnosperms: Classification, Biodiversity, Phytogeography and Ecology: Strelitzia 20 (Natl. Bot. Inst., Pretoria, 2007), p. 206.Google Scholar
  28. 28.
    V. A. Krassilov and E. V. Bugdaeva, “Achene-like Fossils from the Lower Cretaceous of the Lake Baikal Area,” Rev. Palaeobot. Palynol. 36, 279–295 (1982).CrossRefGoogle Scholar
  29. 29.
    V. A. Krassilov and E. V. Bugdaeva, “Gnetalean Plants from the Jurassic of Ust-Balej, East Siberia,” Rev. Palaeobot. Palynol. 53, 359–374 (1988).CrossRefGoogle Scholar
  30. 30.
    V. A. Krassilov and E. V. Bugdaeva, “An Angiosperm Cradle Community and New Proangiosperm Taxa,” Acta Palaeobot., Suppl. 2, 111–127 (1999).Google Scholar
  31. 31.
    V. A. Krassilov and E. V. Bugdaeva, “Gnetophyte Assemblage from the Early Cretaceous of Transbaikalia,” Palaeontogr. Abt. B 253(4–6), 139–151 (2000).Google Scholar
  32. 32.
    V. A. Krassilov and V. I. Burago, “New Interpretation of Gaussia (Vojnovskyales),” Rev. Palaeobot. Palynol. 32, 227–237 (1981).CrossRefGoogle Scholar
  33. 33.
    V. A. Krassilov, V. V. Zherikhin, and A. P. Rasnitsyn,“Classopollis in the Guts of Jurassic Insects,” Palaeontology 40, 1095–1101 (1997).Google Scholar
  34. 34.
    V. A. Krassilov, D. L. Dilcher, and J. G. Douglas, “New Ephedroid Plant from the Lower Cretaceous Koonwarra Fossil Bed, Victoria, Australia,” Alcheringa 22, 123–133 (1998).CrossRefGoogle Scholar
  35. 35.
    V. A. Krassilov, Z. Lewy, and E. Nevo, “Controversial Fruit-like Remains from the Lower Cretaceous of the Middle East,” Cret. Res. 25, 697–707 (2004).CrossRefGoogle Scholar
  36. 36.
    V. Krassilov, Z. Lewy, E. Nevo, and N. Silantieva, Turonian Flora of Southern Negev, Israel (Pensoft, Sophia, 2005).Google Scholar
  37. 37.
    J. Kvaček and B. Pacltová, “Bayeritheca hughesii gen. et sp. nov., a New Eucommiidites-bearing Pollen Organ from the Cenomanian of Bohemia,” Cret. Res. 22, 695–704 (2001).CrossRefGoogle Scholar
  38. 38.
    B. A. R. Mohr and E. M. Friis, “Early Angiosperms from the Lower Cretaceous Crato Formation (Brazil), a Preliminary Report,” Int. J. Plant. Sci. 161(Suppl. 6), 155–167 (2000).CrossRefGoogle Scholar
  39. 39.
    K. R. Pedersen, P. R. Crane, and E. M. Friis, “Pollen Organs and Seeds with Eucommiidites Pollen,” Grana 28, 279–294 (1989).CrossRefGoogle Scholar
  40. 40.
    C. Rydin, B. Mohr, and E. M. Friis, “Cratonia cotyledon gen. et sp. nov.: A Unique Cretaceous Seedling Related to Welwitschia,” Biol. Letters R. Soc. Lond. 270, 29–32 (2003).CrossRefGoogle Scholar
  41. 41.
    C. Rydin, K. R. Pedersen, P. R. Crane, and E. M. Friis, “Former Diversity of Ephedra (Gnetales): Evidence from Early Cretaceous Seeds from Portugal and North America,” Ann. Bot. 98, 123–140 (2006).CrossRefGoogle Scholar
  42. 42.
    G. Sun, Zheng S., D. L. Dilcher, et al., Early Angiosperms and Their Associated Plants from Western Liaoning, China (Sci. Technol. Educ. Publ. House, Shanghai, 2001). 227 p.Google Scholar
  43. 43.
    M. V. Tekleva, V. A. Krassilov, J. Kvaček, and J. H. A. van Konijnenburg-van Cittert, “Eucommiidites: Ultrastructure and Affinities,” Acta Palaeobot. 46(2), 137–155 (2006).Google Scholar
  44. 44.
    J. H. A. Van Konijnenburg-van Citteret, “An Enigmatic Jurassic Microsporophyll Yielding Ephedripites Pollen,” Rev. Palaeobot. Palynol. 71, 101–109 (1992).Google Scholar
  45. 45.
    Wang Z.-Q., “A New Permian Gnetalean Cone As Fossil Evidence for Supporting Current Molecular Phylogeny,” Ann. Bot. 94(2), 181–188 (2004).CrossRefGoogle Scholar
  46. 46.
    R. Wettstein, “Die Entwicklung der Blüte der angiospermen Pflanzen aus derjenigen der Gymnospermen,” Wissen für alle 45, 705–708 (1907).Google Scholar
  47. 47.
    Yang Y., Geng B.-Y., D. L. Dilcher, Chen Z.-D, T. A. Lott, “Morphology and Affinities of an Early Cretaceous Ephedra (Ephedraceae) from China,” Am. J. Bot. 92(2), 231–241 (2005).CrossRefGoogle Scholar

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© Pleiades Publishing, Ltd. 2009

Authors and Affiliations

  1. 1.Borissiak Paleontological Institute of the Russian Academy of SciencesMoscowRussia

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