The evolution of modern corals and their early history

Scleractinians are a group of calcified anthozoan corals, many of which populate shallow-water tropical to subtropical reefs. Most of these corals calcify rapidly and their success on reefs is related to a symbiotic association with zooxanthellae. These one-celled algal symbionts live in the endodermal tissues of their coral host and are thought responsible for promoting rapid calcification. The evolutionary significance of this symbiosis and the implications it holds for explaining the success of corals is of paramount importance. Scleractinia stands out as one of the few orders of calcified metazoans that arose in Triassic time, long after a greater proliferation of calcified metazoan orders in the Paleozoic. The origin of this coral group, so important in reefs of today, has remained an unsolved problem in paleontology. The idea that Scleractinia evolved from older Paleozoic rugose corals that somehow survived the Permian mass extinction persists among some schools of thought. Paleozoic scleractiniamorphs also have been presented as possible ancestors. The paleontological record shows the first appearance of fossils currently classified within the order Scleractinia to be in the Middle Triassic. These earliest Scleractinia provide a picture of unexpectedly robust taxonomic diversity and high colony integration. Results from molecular biology support a polyphyletic evolution for living Scleractinia and the molecular clock, calibrated against the fossil record, suggests that two major groups of ancestors could extend back to late Paleozoic time. The idea that Scleractinia were derived from soft-bodied, ‘‘anemone-like’’ ancestors that survived the Permian mass extinction, has become a widely considered hypothesis. The 14-million year Mesozoic coral gap stands as a fundamental obstacle to verification of many of these ideas. However, this obstacle is not a barrier for derivation of scleractinians from anemone-like, soft-bodied ancestors. The hypothesis of the ephemeral, ‘‘naked coral’’, presents the greatest potential for solution of the enigma of the origin of scleractinians. It states that different groups of soft-bodied, unrelated ‘‘anemone-like’’ anthozoans gave rise to various calcified scleractinian-like corals through aragonitic biomineralization. Although there is evidence for this phenomenon being more universal in the mid-Triassic interval, following a lengthy Early Triassic post-extinction perturbation, it appears to have occurred at least three other times prior to this interval. This idea suggests that, because of ephemeral characteristics, the skeleton does not represent a clade of zoantharian evolution but instead represents a grade of organization. In the fossil record, skeletons may have appeared and disappeared at different times as some clades reverted to soft-bodied existence and these phenomena could account for notable gaps in the taxonomic and fossil record. A fuller understanding and possible solution to the problem of the origin of modern corals may be forthcoming. However, it will require synthesis of diverse kinds of data and an integration of findings from paleobiology, stratigraphy, molecular biology, carbonate geochemistry, biochemistry and invertebrate physiology. D 2002 Elsevier Science B.V. All rights reserved.