massive corals are commoner in quiet waters, and ‘Acropora occurs in high-energy zones (Vosburgh, 1982). The apparent structural strength of corals is therefore only one factor amongst many that control ecological zonation; ability to compete for space and to cope with sediment influx may likewise be important. Zonation varies from: réef to reef depending on various factors. The presence of resistant algal ridges, for instance, can greatly affect the patter of zona- tion. Not only the scleractinians but also the gor- gonians (in the Caribbean reefs) and octocorals (in the Indo-Pacific reefs) are ecologically zoned, and so are the various bivalves and other molluscs that form specialized communities in the very varied reef habitats (Yonge, 1974) The growth rate of corals varies greatly with the environment and the season but may reach 26 cm per year in branching corals and 1 cm per year in the more massive colonies. Reef growth as a whole may reach a maximum of 1.2 cm per year, but in the past it has generally been less. Within reefs there is not only continual construc tion by the scleractinians and encrusting algae, but also a constant attrition through wave action and, equally important, destruction by boring organisms and coral-eating fishes. These produce a great quan- tity of fine comminuted coral sand, which fills up the lagoon and is transported to deeper waters. Deep-water coral banks Deeper-water corals (ahermatypes such as Lophelia and Dendrophyllia) may form structures. differing from those of their shallow-water counterparts. Some of these deep-water corals are solitary; others are colonial and dendriform. The development of these structures (Squires, 1964) takes place in four stages (Fig, 5.174). At first there is a single colony. When this colony is joined by others it becomes a thicket a few metres across, in which the members may be all of the same species or of different species. This newly developed envi- ronment attracts other organisms, e.g, fishes, mol- luses and crustaceans, giving it a new ecological character. As the thicket spreads and matures, skele- tal debris accumulates from broken and bored coral, providing a new substratum which again attracts other animals. This stage is now a coppice and may be several metres across. Eventually, with the accu mulation of more debris and further coral growth, a Class Anthozoa bank develops: a topographic entity with a core of solid skeletal debris, a covering mat of more open debris and a capping ofllive coral. The proportion of living to dead coral decreases with time. The exposed sequences studied by Squires were Tertiary in age, but similar banks are forming today. The study of fossil thickets, coppices and banks can yield valuable palaeoecological data. Geological uses of corals Corals as stratigraphical indicators Corals are generally too long-ranged for use as zone fossils, though the widely distributed Palaeocyclus provides a fine example of a usefal acme biozonal indicator for the late Llandovery-carly Wenlock. Nevertheless, corals have been used where no shorter-ranged fossils are available, especially in the Carboniferous. This work began in Belgium and in 1905 Vaughan, working on the Lower Carboniferous of Bristol, England, erected a zonal scheme based upon the first appearance of corals and brachiopods. This scheme was later extended to the North of England and proved, though crude and inexact, to be applicable, with minor modifications, to much of Europe. Vaughan believed that the faunal sequence he erected reflected evolutionary lineages, but in fact the corals really occur as assemblage biozones with cach of the main cycles bringing in a new migrant fauna. Recently Mitchell (1989) has plotted the time ranges for 68 rugose coral species against standard Visean stages, showing that a now-refined coral stratigraphy is possible for the Lower Carboniferous, supplementing other stratigraphic data based on goniatites, conodonts and foraminiferids. New stages based upon transgressive cycles in the Lower Carboniferous are now standard, but much refine- ment is still needed. Coral zonation in the European Devonian, once elaborated in great detail by Wedekind, has proved to be overoptimistic. Corals as geochronometers In well-preserved specimens of many rugose and scleractinian corals the epithecal surface shows fine growth ridges, some 200 per cm. Each of these rep- resents a growth increment: the former position of the rim of the calice, These fine growth ridges are often grouped in prominent bands or annulations 1356 Cnidarians between which the epitheca is constricted. It is nor~ mally recognized that the fine growth ridges repre- sent daily growth increments, whereas the banding is monthly and the broader and more widely spaced annulations are yearly. Wells (1963) first established in the scleractinian Manicina that the growth ridges were diurnal, and since the Recent Lophelia pertusa, from Norwegian fjords, has-28 growth ridges (pre sumably diurnal) per band, each band corresponding to a lunar cycle, the monthly banding hypothesis seems at least possible. Wells (1963), working on Devonian corals with annual rather than monthly groupings, counted the number of growth ridges per annulation in a num- ber of species. Though the results were limited by the preservation of the corals, he concluded, mainly from observations on Heliophyllum, Eridophyllum and Favosites, that there were an average of 400 days in the Devonian year. This figure corresponded well to astronomical estimates that the Earth’s rotation has been slowing down through tidal friction by about 2 per 100.000 years. Assuming that the Earth’s annual circuit round the Sun was the same length, which seems to be fairly well substantiated, there must have been more days in the Devonian year, but they were shorter (Scruton and Hipkin, 1973). ‘Annual banding is not often found in corals, hav ing only been recorded in specimens that originally lived in waters where there were seasonal fluctua- tions. It is normally easier to work with the Rugosa that show the growth ridges grouped in monthly bands. These are more abundant and do not have to have a complete epitheca. Many Recent corals have monthly breeding cycles during which carbonate deposition is inhibited, and this may have been the case with the Rugosa. The monthly banded Middle Devonian corals studied by Scrutton (1965; Fig. 5.18) had an average of 30.6 growth ridges per monthly band, and if the figure of 399 (the astronomical estimate for length of the Devonian year) is divided by the average growth-ridge count per band, the result is consistently 13. Hence it seems that 13 bands each of about 30.6 growth ridges were laid down by these corals in the Middle Devonian year. The ‘coral clock’ provides a consistent check on estimates of the slowing of the Earth's rotation based Figure 5.18 Bren op te ech se coral #Hel sp. (x3). (Copyright photogra fae co erat ‘of Dr et ‘Scrution.) upon a biological rather than a physical system. It has, however, become increasingly clear that corals are not ideal tools for this, for while diurnal banding is clear enough on all unworn corals, clear monthly and annual cycles are harder to detect and may not be present in many corals (Scruton, 1978). Tidal thythmites (ebb-tidal deposits whose variable lamina thickness encodes a full spectrum of semidiurnal to lunar modal palaeotidal events; Williams, 1989) offer excellent potential for geochronometry, espe- cially for the unfossiliferous Precambrian. Random events may in any case have left a masking record in the growth of the skeleton. Periodic growth features in bivalves are now better understood and may prove to be more suitable for geochronometry.46 = Ontogenetic Variation adult form, especially in the study of organic evolution. To consider an evolutionary series as a sequence of adult forms is to oversimplify. Rather, evolution must be looked upon as a sequence of ontogenies. Types of Growth Organic growth is extremely complicated; it usually involves several types of change, among which are changes in cell size, number of cells, number of cell types, and relative positions of cells. During life, an organism may change in form abruptly (undergo metamorphosis) or it may change gradually. All organisms that depend on a hard skeleton—for support, for protection, or for muscle attachment —must enlarge the skeletal structure to accommodate growth of the soft body. Postembryonic growth of skeletons is accomplished in four ways. ACCRETION OF EXISTING PARTS Most shelled molluscs increase their skeleton size simply by adding new material to the shell throughout life, which has the obvious advantage of permitting continued use of skeletal material deposited at earlier ontogenetic stages. It has the disadvantage, however, that the form of the juvenile shell must be incorporated as part of the adult shell. This type of growth is well illustrated by gastropods (see Figures 2-5 and 2-6). The shell of the coiled gastropod may be looked upon as a hollow tapering tube that is coiled about an axis. As the animal inside the shell grows larger, new material is added to the aperture, or opening, of the tube. Growth lines showing increments of growth can often be seen paralleling the outline of the aperture. Most adult gastropods occupy their entire shells and thus the shape of the soft body of a gastropod is, in effect, an internal mold of the shell. Growth by simple accretion is found in many other animal groups, particularly among animals whose shell is external and serves primarily for protection and muscle attachment. Figure 3-1 illustrates growth in bivalved molluscs. The bivalve grows by accretion in much the same way that the gastropod or cephalopod does. In fact, as we shall see in subsequent chapters, the geometry of the bivalve is not substantially different from that of the other molluscan groups we have considered. We may look upon the individual valve as a hollow cone that tapers from a large, open end to asmall umbo or “beak.” The principal difference is that the “cone” shell of the bivalve is quite shallow and tapers much more sharply from opening to beak than does that of the gastropod or cephalopod. The cross-section of the shell (Figure 3-1) is marked by growth lines which can be observed in the shell as well as on the outer surface. New skeletal material is added not only at the leading edge of the shell, but also as a covering over most of the interior surface of the shell. Figure 3-1 includes an enlarged photograph of a series of growth lines. In this instance the shell was grown under