The oldest bryozoan reefs: a unique Early Ordovician skeletal framework construction

The oldest bryozoan reefs occur in the Lower Ordovician (late Tremadocian) Fenhsiang Formation of the Three Gorges area, South China. These reefs show a unique type of bryozoan (Nekhorosheviella) framework, and were constructed as follows: the first stage involved colonization by lithistid sponges, which acted as a baffler to trap sediments, providing bryozoans with a stable substrate for attachment. The bryozoans then grew as an encruser on the surfaces of sponges, showing a preferential downwards and lateral growth within the sponge scaffolding to avoid biological and physical disturbance. Finally, these biotic combinations among skeletal organisms formed a rigid, threedimensional skeletal framework. This mode of bryozoan growth in association with lithistid sponges is remarkable and unique in its growth direction, and the appearance of such reefs, just prior to the widespread development of skeletal-dominated reefs as part of the Great Ordovician Biodiversification Event, provides an excellent example of the earliest attempts by skeletal organisms to form frameworks by themselves. This find significantly enhances our understanding of the initial stages of skeletal-dominated reef evolution and the ensuing development of reefs during the Middle-Late Ordovician.