Resembled a tree fern with seeds
Stems
Form-genus Lyginopteris †
Usually manoxylic wood, but see Pitys (below)
Leaves
Frond-like, resembling a fern
Form genera Sphenopteris †, Sphenopteridium †, or Pecopteris †
Leaves are developmentally distinct from the branch system
Petiole form genus is Lyginorachis †
Leaf traces separate in the cortex into a pair of strands that fuse in the base of the petiole to form a V- or W-shaped bundle
Reproduction
Ovules
e.g., form genera Lagenostoma †, Gnetopsis †
Ovules are found inside a cupule
They are adiospermic (radially-symmetrical)
Pollen organ
Form genus e.g., Crossotheca †
└Lyginopteridales †
Jennings, 1987
Upper Mississippian stem and frond form taxon
Hirmer, 1933
Carboniferous stem form taxon
May & Matten, 1983
Late Devonian frond form taxon
Mississippian ground cover on a coastal plain
Above: Occloa
Lyginorachis
Pycnoxylic wood from the Mississippian
Parenchymatous pith with up to 50 mesarch primary xylem strands located around the periphery.
Secondary xylem consisted of tracheids with hexagonal–circular bordered pits and wide, high vascular rays
Long (1963) suggested that Lyginorachis papilio was the petiole of Pitys
Tristichia ovensii was the ovule-bearing axis arising in the bifurcation of the Lyginorachis papilio rachis
Foliage may be Sphenopteris or Sphenopteridium-type
Mississippian-Permian of Scotland
Dense wood with alternate, crowded bordered pits on the radial walls of tracheids and numerous, small pits in the cross fields.
Lyginorachis waltonii is suggested to be the leaf of Eristophyton (Long, 1987; Galtier and Scott, 1990)
Carboniferous of Scotland
Stems probably scrambling in habit and with fairly long internodes
Elliptical in transverse section owing to the very gradual disengagement of petioles, that are arranged in sub-opposite pairs and usually in four orthostichies
Marked longitudinal mesh-work of sclerenchymatous hypoderma: mesh long and narrow, except where secondary thickening of axis causes stretching of cortical tissues, and consequent widening of the hypoderma mesh.
Xylem cylinder 4-rayed normally, rarely 5-rayed, solid. Protoxylem of spiral or annular tracheids; metaxylem with scalariform and reticulate thickenings
Secondary tracheids bear reticulate thickenings on both radial and tangential walls.
Medullary rays elongate, narrow, parenchymatous, and without ray tracheids.
Hypodermal sclerenchyma, a network of elongated groups of fibers
Mesh of network elongated longitudinally to stem, and meshes very narrow, except where development of secondary wood has caused tangential stretching of the mesh.
Median zone of outer cortex with many mucilage cells, inner zone a large-celled parenchyma without mucilage elements.
Middle cortex usually badly preserved but containing nests of sclerotic elements.
Parenchyma round sclerotic nests elongated radially with respect to each.
Petioles in nearly opposite pairs, successive pairs at right angles to one another and consequently four orthostichies of these off-sets
Petiole bases much swollen and without hypoderma near junction with stem cortex
Equal dichotomy of petiole some 5 in. beyond junction with stem; first pinnae on outer sides of each arm of bifurcation and normally above that bifurcation.
No a priori reason why primary pinnae should not depart below actual dichotomy of petiole.
First pinna-trace differentiated-at a low level of the petiole trace, and might easily become detached well below the bifurcation
Above: Cross-sections of Tetrastichia
Long, 1961; Galtier & Meyer-Berthaud, 1996
Permineralization of terete protostelic stems bearing leaves
Protostele is 3-lobed with mesarch protoxylem strands near tips of ribs
When present, secondary xylem with narrow rays and tracheids with multiseriate pittings
Petiole equal in width to stem at node, with 1/3 phyllotaxis
Leaf trace tetrarch, papilionoid in transverse section
Cortex showing sclerotic nests and sparganum hypofermis
T. ovensii
Long, 1961
T. longii
Galtier, 1977
T. tripos
Unger, 1856
Upper Mississippian stem form taxon