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DEUTEROSTOMES: ECHINODERMS AND HEMICHORDATES  405





                      Box 15.6  Into the deep: but not until the Late Cretaceous?

               A number of animal groups common in the Paleozoic evolutionary fauna, such as the brachiopods
               and crinoids, are common in deep-water environments. This reinforces the view that the deep sea is
               some sort of refuge for archaic taxa that have been forced down the continental slope by predation
               or unsuccessful competition on the shelf. Andrew Smith and Bruce Stockley (2005) have developed
               a quite different model, however, based on molecular clock estimates and the phylogeny of echinoid
               taxa. Results show that the modern deep-sea omnivore fauna appeared gradually over the last 150–
               200 myr; detritivores, however, were in place during a much shorter time span between 75 and
               50 Ma (Fig. 15.13). This 25 myr window of seaward migration appears to be associated with marked
               increases in seasonality, continental sediment discharge and surface productivity. The increased avail-
               ability of organic carbon and nutrients in the deep sea provided the means for habitat expansion
               rather than an escape from competition and predation on the shelf.



                                              Paleoproductivity  K/T
                                                   –2
                                                      –1
                                               (gCm  yr )
                                                      160
                                                      140
                                                      120
                                           detritivores
                                                      100
                                                                            20
                                           carnivores
                                                       80
                                                       60                   15  Number of deep-sea lineages
                                                      OAEs                  10

                                                                            5


                                     200      150       100      50
                                                     Time (Ma)
               Figure 15.13  Events in the deep sea: cumulative frequency polygons for maximum and minimum
               times of origin of 38 clades of extant, carnivore and detritivore deep-sea echinoids (Smith &
               Stockley 2005). K/T, Cretaceous–Tertiary boundary; OAEs, oceanic anoxic events.




                                                             abundant. The sparse early record of the
                           time

                 Turonian     M. gibbus  Coniacian – Santonian  group might reflect a relatively fragile skele-
              inferred depth of burrowing  M. leskei  M. decipiens  not a common element of the Paleozoic
                                                             ton that quickly disintegrated after death; on
                                                             the other hand the echinoids were probably
                                                 sea bottom
                                                             benthos. The enigmatic Bothriocidaris,
                                                             described from the Ordovician of Estonia and
                                                             from southwest Scotland, has been variously
                                                             classified as an echinoid, cystoid or holothu-

                                        M. coranguinum
                                                             rian. Some authorities consider that Bothrio-
             Figure 15.14  Evolution of the Late Cretaceous   cidaris and Eothuria might be unclassifi able
             heart urchin, Micraster. (Based on Rose, E.P.F. &   echinoids, hopeful monsters that arose during
             Cross, N.E. 1994. Geol. Today 9.)               the rapid Ordovician radiation of the group.
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