The Voltziales are an extinct group of conifer-like gymnosperms which appear in the Pennsylvanian and disappear during the Cretaceous extinction. There are generally two types of Voltziales: walchian and voltzian. The former had needle-like leaves and branches growth parallel to the ground; the latter with scale-like leaves and more upright branches. It is possible that the Voltziales were intermediate between the true conifers and the cordaites.
Two types of Voltiales: Walchian and Voltzian types
Plagiotropic branching (branches grow parallel to the ground)
Needle-like leaves
Dwarf shoots with radially-arranged scales
Pollen monosaccate
Orthotropic branching (branches grow upright; perpendicular to ground)
Leaves are broad, dwarf shoots bearing flattened and partially fused scales
Pollen bisaccate
Fused female reproduction organs
Reproduction involves a pollen tube
└Voltziales †
Mostly monoecious
Differentiated megasporangiate (♀) and microsporangiate (♂)
Ovuliferous scales
Highly-condensed, fertile shoot
Ovuliferous scales clustered into fertile area or cone
Utrechtiaceae
Thucydiaceae
Emporiaceae
Majonicaceae
Ullmanniaceae
Bartheliaceae
Ferugliocladaceae
Buriadiaceae
Krassiloviaceae
Vegetative branches
Vegetative branches
S. daohugouensis (Zhang et al. 2011)
S. liasokeuperianus (Braun 1847)
Bilobate cone scales from the Late Triassic to Early Cretaceous of Europe & northern Asia
Jongmans & Gothan 1925, 1935
Early Permian (Asselian) from Sumatra
Cylindrical female cones with deeply split cone scales of a possible early Voltzian cone
Seed scales and bract scales are helically- and loosely-arranged on cone axis (Harris 1979)
Above: Reconstruction of Tobleria bicuspis (Van Wessen 2019)