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THE BASAL METAZOANS: SPONGES AND CORALS  279






                             discoid
                                                                            scolecoid
                                                       trochoid


                            patellate









                            turbinate    ceratoid      cylindrical        calceoloid   pyramidal

             Figure 11.19  Terminology for the main modes of solitary growth in corals. (From Treatise on
             Invertebrate Paleontology, Part F. Geol. Soc. Am. and Univ. Kansas Press.)

             arrangement. At the circumference, the septa    grams (Fig. 11.20). Moreover colonies with
             are joined to the theca or skeletal wall, which   imperforate walls may exhibit phaceloid,
             extends longitudinally from the apex of the     cateniform, cerioid and meandroid forms,
             corallum to the  calice where the polyp is      whereas those with perforate walls have only
             attached. During growth the polyp may secrete   phaceloid and cerioid growth modes together
             a series of horizontal sheets, or tabulae,      with coenenchymal structures in some taxa,
             together with smaller curved or angled plates   such as the sarcinulids. These growth modes
             or dissepiments. The columella, usually arising   are variably developed across the rugosans,
             from the fusion of the axial edges of the septa,   tabulates and scleractinians – but meandroid
             occupies the core region of the corallum. The   and hydnophoroid modes were developed
             vertical walls or septa radiate outwards from   during the Mesozoic and are thus restricted
             the columella and divide the corallite. Despite   to the scleractinians.
             the apparent simplicity of the coral skeleton,    Colonial integration usually involves a loss
             there is a great deal of variation in both soli-  of individuality. Many organisms display a
             tary and colonial growth programs and the       transition from solitary growth modes,
             end result is a remarkable array of shapes and   through morphologies with asexually budded
             sizes of corals.                                modules, to a fully integrated colony with the
               The three main subclasses of stony corals     growth or astogeny of the compound struc-
             have colonial or compound growth modes          ture showing little variation across the indi-
             whereas only the Scleractinia and Rugosa        vidual corallites. The degree of integration of
             have solitary skeletons. The solitary growth    a colony is usually measured by the amount
             forms include conical, ceratoid or horn-        of cohesion between the individual skeletal
             shaped, calceoloid, cylindrical, discoid, patel-  parts and soft tissues and by the range of form
             late, scolecoid, trochoid and turbinate         observed between individual components.
             skeletons (Fig. 11.19). Colonial corals with    Clearly there is a spectrum from phaceloid
             corallites have adopted either fasciculate or   modes with little or no integration to thamn-
             massive growth modes. Fasciculate styles        asteroid and meandroid (and coenenchymal)
             exhibit either dendroid or phaceloid strategies   modes with high levels of integration. Indi-
             with either no or poor integration. The hal-    vidual polyps are no longer separated by cor-
             ysitid or cateniform chain-like growth strat-   allite walls and may share a common enteron
             egy is a further variation on this pattern. The   and nervous system. This suggests a high
             massive colonies are much more varied, with     degree of integration where the colony
             cerioid, astraeoid, aphroid, thamnasteroid,     approaches the body plan of a typical meta-
             meandroid and hydnophoroid together with        zoan. These modes have varied through time
             coenenchymal or coenostoid growth pro-          (Fig. 11.21).
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