A Reconsideration of Hyenia elegans Kräusel et Weyland and Hyenia `complexa' Leclercq: Two Middle Devonian Cladoxylopsids from Western Europe

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Abstract
Two taxa of Middle Devonian Cladoxylopsida from localities in Belgium and Germany are redescribed. A counterpart of one of the missing type specimens of Hyenia elegans Kräusel et Weyland from Kirberg, Germany, demonstrates that rather than having simple forked fertile appendages (as apparent from the type description), more complex fertile units were present; these consisted of two daughter segments (resulting from a proximal dichotomy), each of which bore three pairs of elongate sporangia on short recurved stalks proximal to a terminal bifurcation. Specimens from Oé, Belgium, informally referred to as Hyenia `complexa' by Leclercq, were shown to have a similar fertile organ that lacked the tuft of filiform extensions previously believed to be present. The parallel axes of this plant are now interpreted to be derived from a digitate branch rather than from a horizontal rhizome. Both taxa, H. elegans and H. `complexa,' therefore demonstrate fertile appendage morphology and other characters that are typical of the Middle Devonian cladoxylopsid Calamophyton primaevum Kräusel et Weyland, which has priority.