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THE BASAL METAZOANS: SPONGES AND CORALS  261


             Until nearly the end of the Neoproterozoic,     eggs hatching within the parent sponge, and
             some 80% of geological time, the oceans of      larvae released into the water.
             the world were mainly occupied by rather          There are over 10,000 species of sponge.
             simple, usually unicellular, organisms. By the   All are aquatic, and most are marine. Sponges
             Ediacaran it was clear that this simple exis-   are part of the sessile benthos, fi xed  to  the
             tence was not enough, and more complex          seabed, pumping large volumes of water – in
             body plans were soon to develop their own       extreme cases over 1000 L per day – through

             ecosystems. Two groups, the Porifera and the    their fixed but commonly fl exible  bodies,

             Cnidaria, form the basal parts of the meta-     which act as filters for nutrients. The group
             zoan tree, diverging during the Neoprotero-     has a remarkable range of morphologies; the
             zoic. Despite their origins in deep time and    more specialized, stalked forms live in deep-
             their relative simplicity, both maintained      water environments and fl attened,  dumpy
             high diversity, notably as colonial organisms,   forms prefer shallower-water, high-energy
             throughout the Phanerozoic, frequently          environments. Despite the apparent simplicity

             becoming important parts of the planet’s reef   of the sponges, the classification of the phylum
             ecosystems.                                     has recently undergone considerable revision
                                                             (Box 11.1). Some well-established calcifi ed
                                                             groups, such as the “chaetetids” and “sphinc-
                                                             tozoans”, are probably polyphyletic, merely
             PORIFERA
                                                             representing convergence towards common
               So he dissected sea sponges by night,         grades of organization. The well-established
               winter night after winter night  .  .  .  adult   and diverse Demospongea, the common
               and embryo human body parts by day,           sponges, may too be polyphyletic. Despite
               adult and larval sponge body parts by         their relative simplicity, the complex relation-
               night.                                        ships of “sponge-grade” animals have yet to
                                                             be resolved.
                  Rebbeca Stott (2003) Darwin and the
                Barnacle, on the sponge doctor, Robert
                                                Grant
                                                             Morphology: examining a typical sponge
                                                             A typical sponge individual is not particularly
             Most of us have used a bath sponge, probably    complex or intellectually demanding to under-
             a synthetic replica of the real thing. But ancient   stand; it is nonetheless a remarkable organ-
             peoples used sponge skeletons as an aid to      ism. It is sac-shaped with a central cavity or
             bathing and possibly exfoliation in some of     paragaster, which opens externally at the top
             the world’s earliest and most exclusive health   through the osculum (Fig. 11.1). The sponge
             farms. Most considered they were some form      is densely perforated by  ostia,  small holes
             of plant until proper biological study in the   marking the entrances to minute canals
             mid and late 1700s suggested they were          through which pass the inhalant currents. In
             animals – and at first they were classifi ed as   simple terms, there are three main cell types:


             corals. It was in fact Dr Robert Grant (1793–   (i) flattened epithelial cells; (ii) collar cells or
             1874), one time mentor to Charles Darwin,       choanocytes, which occupy the internal cham-
             who later established the Porifera as a unique   bers and move water along by beating their
             group in its own right. The poriferans or       fl agella; and (iii) amoeboid cells, which have
             sponges have a unique porous structure and      digestive, reproductive and skeletal functions.
             a body plan based at the cellular level of orga-  Amoeboid cells can actually irreversibly
             nization; they are said to lack true tissues.   change into other cell types with other func-
             Most lack symmetry, true differentiated         tions. Nutrient-laden water is thus sucked

             tissues, and organs, although their cells, like   through the ostia, flagellated by the choano-
             those of the protists, can switch function.     cytes and processed by the amoeboids. Waste
             They reproduce both asexually (by budding)      products and spent water, together with repro-
             and sexually with different cells expelling     ductive products when in season, are ejected
             clouds of eggs and sperm out through an         upwards through the paragaster into the
             opening; some are even viviparous, with the     water column.
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