The Early Ediacaran Caveasphaera Foreshadows the Evolutionary Origin of Animal-like Embryology

The Ediacaran Weng’an Biota (Doushantuo Formation, 609 million years old) is a rich microfossil assemblage that preserves biological structure to a subcellular level of fidelity and encompasses a range of developmental stages. However, the animal embryo interpretation of the main components of the biota has been the subject of controversy. Here we describe the development of Caveasphaera, which varies in morphology from lensoid, to a hollow spheroidal cage, to a solid spheroid, but has largely evaded description and interpretation. Caveasphaera is demonstrably cellular and develops within an envelope by cell division and migration, first defining the spheroidal perimeter via anastomosing cell masses that thicken and ingress as strands of cells that detach and subsequently aggregate in the polar region. Concomitantly, the overall diameter increases as does the volume of the cell mass but, after an initial phase of reductive palinotomy, the volume of individual cells remains the same through development. The process of cell ingression, detachment and polar aggregation is analogous to gastrulation; together with evidence of functional cell adhesion and development within an envelope, this is suggestive of a holozoan affinity. Maternal investment in the embryonic development of Caveasphaera and co-occuring Tianzhushania and Spiralicellula, as well as delayed onset of later development, may reflect an adaptation to the heterogeneous nature of the early Ediacaran nearshore marine environments in which early animals evolved.