Early heliolitine tabulate corals from the Sandbian (Upper Ordovician) in the Yunnan-Sichuan border area, Southwest China

Heliolitine tabulate corals are extremely rare in pre-Katian (Upper Ordovician) rocks worldwide, hindering the understanding of their origin and early evolution. Here, we systematically study new heliolitine material from the upper member (Sandbian) of the Huadan Formation in the Yunnan-Sichuan border area, Southwest China, which includes Protaraea ningnanensis, Plasmoporella kiaeri, Wormsipora sp. and Heliolites spinosus. The findings indicate that Protaraea and Heliolites are ancestral forms of coccoserids and typical tubular heliolitines, respectively, and that the absence of trabecular septa is probably a primitive feature of plasmoporellids. The refined Ordovician stratigraphy of this area implies that the coccoserid Protaraea ningnanensis Lin, 1965 from the lower member (late Darriwilian) of the Huadan Formation is the earliest known heliolitine, which further indicates that all heliolitines might have evolved from Protaraea. The comparison of Protaraea with other early tabulate corals also suggests an independent heliolitine origin from auloporinids and sarcinulinids.