On the horizon of Protopteryx and the early vertebrate fossil assemblages of the Jehol Biota

Protopteryx, a monotypic fossil bird discovered from the Sichakou basin in Fengning, Hebei, is the most primitive enantiornithine currently known. The bird-bearing strata do not contain the index fossils of the Yixian Formation in western Liaoning; the fish and bird fossils have more primitive features than the related forms found in the Yixian Formation, and the conchostracans are those usually distributed in the Dabeigou and Dadianzi formations in northern Hebei. Besides, the Protopteryx-bearing strata underlie the deposits bearing the index fossils of the Yixian Formation in the neighboring basin. Thus, it could be confirmed that the horizon of Protopteryx should be lower than the Yixian Formation, and is approximately equivalent to the Dadianzi Formation in northern Hebei. This is the lowest horizon of the known fossil birds in China and Mesozoic enantiornithine birds in the world. Accompanying Protopteryx, there are other birds, acipenseriform fishes, salamanders, and mammals, which compose the Peipiaosteus fengningensis-Protopteryx fengningensis assemblage. This new assemblage traces the vertebrate evolution history of the Jehol Biota back to 130.7 Ma before. It is suggested that the demarcation of the Jehol Biota should be based on the large-scale tectonic-sedimentary cycles, and Peipiaosteus, instead of Lycoptera, could be taken as the vertebrate representative of the Jehol Biota.