capability for fertilization until after the release of polar bodies. This delay, and possibly some degree of self-incompatibility, likely increases the chance of cross-fertilization. A study of four species of Scleractinia found that cross-fertilization was actually the dominant mating pattern, although three of the species were also capable of self-fertilization to varying extents. Evolutionary history There is little evidence on which to base a hypothesis about the origin of the scleractinians; plenty is known about modern species but very little about fossil specimens, which first appeared in the record in the Middle Triassic (). It was not until 25 million years later that they became important reef builders, their success likely a result of teaming up with symbiotic algae. Nine of the sub-orders were in existence by the end of the Triassic and three more had appeared by the Jurassic (200 million years ago), with a further suborder appearing in the Middle Cretaceous (100 million years ago). Some may have developed from a common ancestor, either an anemone-like coral without a skeleton, or a rugose coral. A rugose coral seems an unlikely common ancestor because these corals had calcite rather than aragonite skeletons, and the septa were arranged serially rather than cyclically. However, it may be that similarities of scleractinians to rugosans are due to a common non-skeletalized ancestor in the early Paleozoic. Alternatively, scleractinians may have developed from a Corallimorpharia-like ancestor. It seems that skeletogenesis may have been associated with the development of symbiosis and reef formation, and may have occurred on more than one occasion. DNA sequencing appears to indicate that scleractinian corals are a monophyletic group. The earliest scleractinians were not reef builders, but were small, phaceloid or solitary individuals. Scleractinian corals were probably at their greatest diversity in the Jurassic and all but disappeared in the mass extinction event at the end of the Cretaceous, about 18 out of 67 genera surviving. Recently discovered Paleozoic corals with aragonitic skeletons and cyclic septal insertion – two features that characterize Scleractinia – have strengthened the hypothesis for an independent origin of the Scleractinia. Whether the early scleractinian corals were zooxanthellate is an open question. The phenomenon seems to have evolved independently on numerous occasions during the Tertiary, and the genera Astrangia, Madracis, Cladocora and Oculina, all in different families, each have both zooxanthellate and non-zooxanthellate members. The fact that zooxanthellate coral make up only about half of the