New materials from the Upper Devonian (Famennian) Wutong Formation of Chizhou District, Anhui Province, South China, allow description of fertile and sterile characters of Archaeopteris halliana. This plant has penultimate axes attached by sterile leaves and paired ultimate branches in the same ontogenetic spiral. Sterile leaves are narrowly cuneate in shape and bear distal margins dissected deeply. These leaves are spirally arranged on sterile ultimate axes. Non-laminated sporophylls occur spirally on the ultimate axes and bifurcate once or twice. Elongate sporangia with longitudinal dehiscence are borne adaxially below or above the dividing points of the sporophylls. Despite the leaf shape resembling that of Archaeopteris macilenta, our plant is assigned to A. halliana because of its non-laminated sporophylls. Fertile characters should be given priority or enough consideration in identifying species of Archaeopteris.
Well-preserved specimens of Hamatophyton verticillatum collected from the Upper Devonian (Famennian) Wutong Formation of Chaohu district, Anhui Province, South China, display more complete fertile axes in three orders and multiple divisions. Comparisons indicate that Hamatophyton possibly does not have palmate planate sterile leaves but hook-like linear ones with rare divisions. We propose seven definitive characters of Sphenophyllales: (1) completely whorled lateral organs; (2) sterile leaves; (3) strobili; (4) "sporangiophores" or stalks with reflexed tips bearing sporangia; (5) three- or four-ribbed primary xylem; (6) exarch maturation of primary xylem; and (7) secondary xylem. The Sphenophyllales probably originated from the Iridopteridales based on similarities in whorled lateral organs, ribbed primary xylem and peripheral protoxylem strands. In transition from Iridopteridales to Sphenophyllales, morphological changes involve partially whorled to completely whorled lateral organs, sterile ultimate appendages to leaves, and fertile ultimate appendages to "sporangiophores"/stalks with bracts; anatomical modifications include configuration and maturation of primary xylem, and presence of secondary xylem.