Page 115 - Introduction to Paleobiology and The Fossil Record
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102  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                      been promoted by storms, providing fresher-       In marine environments acceleration of the
                      water conditions for short periods of time.     height, complexity and stratifi cation of benthic
                                                                      tiering was later matched by increases in the
                                                                      depth and sophistication of infaunal tiering
                      Ecological patterns and trends through time
                                                                      as, particularly in Mesozoic and Cenozoic
                      During the last 600 myr, both animal and        faunas, many more organisms adopted bur-
                      plant communities expanded and diversifi ed      rowing lifestyles and the benthos switched

                      (Box 4.5). In simple terms the number of        from filter to deposit feeding with signifi cantly
                      Bambachian megaguilds multiplied through        more predators. The Cambrian evolutionary
                      the Cambrian (nine megaguilds), Paleozoic       fauna occupied, more or less, only the surface
                      (14) and Modern (20) evolutionary faunas.       of the seabed, but by the Ordovician crinoids
                      The focus in the Cambrian was on marine         had developed tiers over a meter above the
                      animals that were either attached or mobile     seabed and burrowing had already com-
                      with suspension- or deposit-feeding strategies,   menced into the sediment. Terrestrial environ-
                      such as the eocrinoids and trilobites. The      ments, initially dominated by small green
                      morphologies of individual organisms were       plants, various arthropods and snails, together
                      rather plastic as were their community com-     with diverse amphibian faunas in the Mid to
                      positions and structures. Relatively few class-  Late Paleozoic, changed signifi cantly  during
                      level taxa were included in each ecological     the Mesozoic, with the diversifi cation of veg-
                      box (Fig. 4.21). By the Ordovician, however,    etation and eventually fl owering plants, and
                      the number of megaguilds had expanded,          terminating, for now, in the high and elabo-
                      with an overall numerical dominance of sus-     rate canopies we see today in the tropical rain
                      pension feeders, such as the brachiopods,       forests (see p. 505).
                      bryozoans, corals and crinoids. The Paleozoic     The Modern fauna was also characterized
                      fauna was characterized by sedentary organ-     by something rather special, an arms race
                      isms. The Modern fauna, by contrast, was        (Harper 2006). During the so-called Meso-
                      dominated by deposit-feeding, essentially       zoic marine revolution, predators, such as

                      mobile animals bound into a process of esca-    bony fishes, crustaceans, marine reptiles and

                      lation, or ever-increasing competition, and the   starfishes began to develop better and better
                      first intense arms race on the planet. The term   ways of crushing or opening shells. The

                      arms race is used by ecologists to describe     Modern world was a much more dangerous
                      ever-intensifying interactions between preda-   place and in order to survive, potential prey
                      tors and prey, for example.                     had to develop thicker, more elaborately
                        Throughout the Phanerozoic there seems to     ornamented shells with smaller apertures
                      have been an offshore movement in marine        (Box 4.6) and devise more cunning evasive
                      faunas. New communities and taxa may have       strategies such as greater mobility or deeper
                      occurred in nearshore, high-energy environ-     and deeper burrowing. Unfortunately expo-
                      ments first, before migrating into deeper        sure to intense predation and a much more


                      water. Thus older, more archaic groups tended   bioturbated seafloor was no place for many
                      to characterize deeper-water habitats. For      groups of epifaunal animals such as the bra-
                      example during the Ordovician radiation (see    chiopods, some groups of bivalves and echi-
                      p. 253), typical members of the Paleozoic       noderms. But as prey developed more armor
                      fauna (brachiopods, bryozoans and crinoids)     and better evasive strategies, the hunters
                      expanded and migrated into deeper-water         developed better weaponry. Together this
                      habitats, while their place in shallow water    escalation and increased tiering set the
                      was taken by components of the Modern           Modern fauna quite apart from those of the
                      fauna (bivalves and gastropods). But why?       Cambrian and Paleozoic. Perhaps the whole
                      Are nearshore habitats particularly harsh,      ecosystem functioned in a different way,
                      driving innovative communities and taxa into    allowing biodiversity to continue to expand
                      deep water, or can innovative organisms arise   way beyond the plateau of the Paleozoic
                      at any depth and those in shallower-water       fauna (see p. 541).
                      environments are just more resistant to extinc-   Unlike biodiversity change, where we have
                      tion and can readily migrate into deeper water   numbers of taxa to count and monitor, eco-
                      (Jablonski & Bottjer 1990)? Perhaps it was a    logical change is much more diffi cult to describe
                      combination of both.                            and quantify. Since some changes are much
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