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178  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                                                                      including a substantial extinction phase bet-
                      EXTINCTION THEN AND NOW
                                                                      ween the Middle and Late Permian, some
                      Extinction events                               10 myr before the PT event. This Middle–Late
                                                                      Permian extinction, the  end-Guadalupian
                      Somewhere between background extinction         event, may turn out to be a mass extinction
                      and mass extinction have been many times        in its own right. Numerous marine and non-
                      when rather large numbers of species have       marine groups were hard-hit at that time, and
                      died out, but perhaps only in one part of the   it has been hard to identify until recently
                      world, or perhaps affecting only one or two     because its effects were sometimes confused
                      ecological groups. These medium-sized extinc-   with the end-Permian event, because of lack
                      tions are often classed together as extinction   of clarity about dating.
                      events, but clearly each one is different. Many   There were further such events at the end
                      extinction events have been identifi ed (see Fig.   of the Early Triassic and in the Late Triassic.
                      7.2), and some of the better-known ones are     The Late Triassic extinction event, more com-
                      noted briefl y here.                             monly called the Carnian-Norian event (after
                        The fi rst  is  the  Ediacaran  event, about   the stratigraphic stages) occurred some 15–
                      542 Ma, which is ill defined in terms of timing,   20 myr before the end-Triassic mass extinc-

                      but it marks the end of the Ediacaran animals   tion. The Carnian-Norian event was marked
                      (see pp. 242–7). Some Ediacaran beasts may      by turnovers among reef faunas, ammonoids
                      have survived into the Cambrian, but the        and echinoderms, but it was particularly
                      majority of those strange quilted jellyfi sh-like,   important on land. There were large-scale
                      frond-like and worm-like creatures disap-       changeovers in floras, and many amphibian

                      peared, and the way was cleared for the dra-    and reptile groups disappeared, to be followed
                      matic radiation of shelly animals at the        by the dramatic rise of the dinosaurs and
                      beginning of the Cambrian. Because of the       pterosaurs. At this time, many modern groups
                      antiquity of this proposed mass extinction, it   arrived on the scene, such as turtles, crocodil-
                      is hard to be sure that all species became      ians, lizard ancestors and mammals. The
                      extinct at the same time, and some would        cause of these events may have been climatic
                      argue that this was not a mass extinction at    changes associated with continental drift. At
                      all. Causes are equally debated, with some      that time, the supercontinent Pangaea (see p.
                      evidence for a nutrient crisis or a major tem-  48) was beginning to break up, with the
                      perature change. An older putative mass         unzipping of the Central Atlantic between
                      extinction, at the start of the Ediacaran, some   North America and Africa.
                      650 Ma, might have been triggered by global       Extinctions during the Jurassic and Creta-
                      cooling, the “snowball Earth” model (see p.     ceous periods were minor. The Early Jurassic
                      112), but this is equally debated.              and end-Jurassic events involved losses of
                        An extinction at the end of the Early Cam-    bivalves, gastropods, brachiopods and ammo-
                      brian marked the disappearance of previously    nites as a result of major phases of anoxia.
                      widespread archaeocyathan reefs (see p. 268).   Free-swimming animals were unaffected, and
                        A series of extinction events occurred        the events are undetectable on land – they
                      during the Late Cambrian, perhaps as many       may be partly artificial results of incomplete


                      as five, in the interval from 513 to 488 Ma.     data recording. Events have been postulated
                      There were major changes in the marine          also in the Mid Jurassic and in the Early Cre-
                      faunas in North America and other parts of      taceous, but they are hard to determine. The
                      the world, with repeated extinctions of trilo-  Cenomanian-Turonian extinction event some
                      bites. Following these, animals in the sea      94 Ma, associated with extinctions of some
                      became much more diverse, and groups such       planktonic organisms, as well as the bony
                      as articulated brachiopods, corals, fi shes, gas-  fishes and ichthyosaurs that fed on them, is

                      tropods and cephalopods diversifi ed dramati-    probably associated with sea-level change.
                      cally during the great Ordovician radiation       Extinctions since the KT event have been
                      (see p. 253).                                   more modest in scope. The Eocene-Oligocene
                        There were many further extinction events     events 34 Ma were marked by extinctions
                      or turnover events in the Paleozoic, between    among plankton and open-water bony fi shes
                      the Late Devonian and PT mass extinctions,      in the sea, and by a major turnover among
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