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THE BASAL METAZOANS: SPONGES AND CORALS  293





















               (a)                                          (b)
             Figure 11.35  Devonian reefs of the Canning Basin, Australia: (a) main face, and (b) Windjana Gorge.
             The fore-reef slope in the foreground has large blocks of unbedded reef material in the background; the
             reef is prograding over the fore-reef toward the viewer. (Courtesy of Rachel Wood.)


                                                             building and non-reef-building groups in
             Distribution: corals through time
                                                             shallow- and deep-water environments, respec-
             Although some coral-like forms have been        tively. Scleractinian evolution was marked by
             described from the Cambrian, most lack          a number of morphological trends: solitary life
             typical zooantharian structures.  Cothonion,    strategies were eventually superseded by a
             for example, with poorly integrated corallite-  dominance of colonial forms that display tran-
             like clusters and opercula was probably a       sitions from low levels of integration in phace-
             Cambrian experiment with coralization (Fig.     loid growth modes to higher levels in meandroid
             11.36). The first tabulates appeared during      styles, common in modern reefs.

             the Early Ordovician with cerioid growth          Corals have been used effectively for the
             modes; tabulae were rare and septa and mural    correlation of Silurian (tabulates) and Devo-
             pores were absent. Nevertheless, by the Mid     nian (rugose) strata but they have been proved
             and Late Ordovician the more typical charac-    most useful for Carboniferous biostratigra-
             ters of the Tabulata had evolved when they      phy. During the early 1900s, Arthur Vaughan
             dominated coral faunas. Some workers have       studied in detail the distribution of Lower
             removed the heliolitids, with individual coral-  Carboniferous corals in Belgium and Britain
             lites mutually separated by extensive coenos-   and he argued they would be of great value
             teum, from the Tabulata, as a distinct order.   in Carboniferous biostratigraphy. Corals are
             The group was common until the Early Silu-      very common, often widespread, usually dis-
             rian when the more open structures of the       tinctive, and well preserved in the Lower Car-
             favositids with massive cerioid colonies began   boniferous rocks of Europe. However, more
             to dominate, although they were already         modern studies on Carboniferous biostratig-
             abundant in the Ordovician.                     raphy using microfossils such as conodonts
               The Rugosa appear during the Mid              and foraminiferans, together with sequence
             Ordovician. Many of the evolutionary trends     stratigraphy, have shown that the occurrences
             across the order have been repeated many        of corals are controlled as much by rock facies
             times in different families. In general terms the   as by time, and so they cannot be used for
             group evolved more complex, heavier skeletons   global correlation. Nevertheless, many corals
             prior to extinction at the end of the Permian.  are still useful for local correlations, and

               The first scleractinians were established by   Lower Carboniferous stratigraphy (Fig. 11.37)

             the Mid Triassic, derived from multiple ances-  has been refined on the basis of Vaughan’s
             tors among the sea anemones. The Triassic       pioneer work and more modern techniques
             taxa were probably photosymbiotic, forming      (Riley 1993).
             patch reefs in parts of the Tethyan belt. The     But was coloniality amongst the bilaterians
             group, however, expanded signifi cantly during   a derived condition or, more controversially,
             the Jurassic with the radiation of both reef-   a primitive state (Box 11.10)?
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