A new tree fern stem, Heilongjiangcaulis keshanensis gen. et sp. nov., from the Cretaceous of the Songliao Basin, Northeast China: a representative of early Cyatheaceae

A new genus and species of Cretaceous Cyatheacean tree fern, Heilongjiangcaulis keshanensis gen. et sp. nov., is erected for several permineralized stems collected at the Keshan County in Songliao Basin, Heilongjiang Province, Northeast China. The new taxon is characterized by a dictyostelic, erect stem with dense multicellular scales and surrounded by persistent petiole bases and adventitious roots. The stem contains a central pith lacking medullary bundles, which is surrounded by a dictyostele, and the cortex externally. Each meristele of the dictyostelic ring is enclosed by a sclerenchyma sheath. The pith and cortex are parenchymatous. The proximal petiole bases present a frond trace composed of numerous meristeles, arranged in 1 abaxial and 2 adaxial arcs, with internally projecting bundles on the upper and lateral sides. The feature combination of the new genus is nearly identical to the anatomical structures of modern scaly genera of the Cyatheaceae apart from the absence of medullary bundles. It is interpreted as a primitive representative of early Cyatheaceae, that closely resembles the modern scaly genera, which suggests that in the Cretaceous, the tree ferns in this family were already in possession of most of the anatomical characteristics observed in extant taxa. The fossil records of the stems, petioles, and spores indicate that during the Jurassic and Cretaceous, the Northeastern region of Asia may have been one of the distribution centers of early Cyatheaceae.