Diversity of large woody lignophytes preceding the extinction of Archaeopteris: New data from the middle Tournaisian of Thuringia (Germany)

Anatomically preserved axes representing three lignophyte species occur in the middle Tournaisian deposit of Kahlleite in Thuringia. One is characterized by a small oval eustele, short uniseriate rays, and alternate distichous phyllotaxy. It is assigned to the progymnosperm genus Protopitys. The two others share a eustelic primary vascular system comprising a parenchymatous pith and numerous xylem strands in a peripheral position. The secondary xylem comprises rays that are mostly uniseriate and rarely exceed 20 cells in height. One is referred to as Eristophyton sp.; the second, characterized by ray cells showing a wide range of sizes and shapes is assigned to Aporoxylon primigenium. These records extend the stratigraphical range of Protopitys and Eristophyton down to the middle Tournaisian and confirm their great longevity through most of the Mississippian. They suggest that the diversity of putative arborescent lignophytes co-occurring with Archaeopteris around the D/C boundary but surviving successfully above this limit has been underestimated.