its host. This hints at an intricate set of relationships between the members of the coral holobiont that have been developing as evolution of these members occurs. A study published in 2018 revealed evidence of phylosymbiosis between corals and their tissue and skeleton microbiomes. The coral skeleton, which represents the most diverse of the three coral microbiomes, showed the strongest evidence of phylosymbiosis. Coral microbiome composition and richness were found to reflect coral phylogeny. For example, interactions between bacterial and eukaryotic coral phylogeny influence the abundance of Endozoicomonas, a highly abundant bacterium in the coral holobiont. However, host-microbial cophylogeny appears to influence only a subset of coral-associated bacteria. Reefs Many corals in the order Scleractinia are hermatypic, meaning that they are involved in building reefs. Most such corals obtain some of their energy from zooxanthellae in the genus Symbiodinium. These are symbiotic photosynthetic dinoflagellates which require sunlight; reef-forming corals are therefore found mainly in shallow water. They secrete calcium carbonate to form hard skeletons that become the framework of the reef. However, not all reef-building corals in shallow water contain zooxanthellae, and some deep water species, living at depths to which light cannot penetrate, form reefs but do not harbour the symbionts. There are various types of shallow-water coral reef, including fringing reefs, barrier reefs and atolls; most occur in tropical and subtropical seas. They are very slow-growing, adding perhaps one centimetre (0.4 in) in height each year. The Great Barrier Reef is thought to have been laid down about two million years ago. Over time, corals fragment and die, sand and rubble accumulates between the corals, and the shells of clams and other molluscs decay to form a gradually evolving calcium carbonate structure. Coral reefs are extremely diverse marine ecosystems hosting over 4,000 species of fish, massive numbers of cnidarians, molluscs, crustaceans, and many other animals. Evolution At certain times in the geological past, corals were very abundant. Like modern corals, their ancestors built reefs, some of which ended as great structures in sedimentary rocks. Fossils of fellow reef-dwellers algae, sponges, and the remains of many echinoids, brachiopods, bivalves, gastropods, and trilobites appear along with coral fossils. This makes some corals useful index fossils. Coral fossils are not restricted to reef remnants, and many solitary fossils are found elsewhere, such as Cyclocyathus, which occurs in England's Gault clay formation. Early corals Corals first appeared in the Cambrian about .