Isolated cuticle fragments, spores, and tubes are found in Lower to Middle Ordovician rocks, and extend through the Silurian into the Lower Devonian (Edwards et al., 1979; Wellman, 2001; Edwards, 1986). Although even less is known about these fragments than the enigmatic fossils described earlier, they constitute an important source of paleobiological data because they occur at a point in geologic time that coincides with the origin of a terrestrial biota (T. Taylor, 1982a, 1988a; Kenrick, 2003; Steemans and Wellman, 2004).
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