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122  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                                          geographic barrier

                                     species B       species C
                                                                                   species A         species B











                       Time




                                                                                           isolation
                                                                                               environmental crisis
                                                                                               or random event
                                             species A

                                                                                    species A


                      (a)                                                      (b)
                      Figure 5.2  Allopatric speciation models, occurring either symmetrically (a), where the parent species is
                      divided into two roughly equal halves by a geographic barrier, or asymmetrically (b), where a small

                      peripheral population is isolated by a barrier. In the first case, two new species may arise; in the second,
                      the parent species may continue unaltered, and the peripheral population may evolve rapidly into a new
                      species.


                                                                      These two models of evolution seem so dis-
                                                                      tinctive, both in the shape of phylogenies, and
                                                                      in their interpretation, that it should be pos-
                                                                      sible to test between them by observations
                                                                      from the fossil record.


                                                                      Testing punctuated equilibrium: problems
                      (a)                        (b)
                                                                      Eldredge and Gould (1972) argued that many
                      Figure 5.3  Two models of speciation and lineage   test cases of the pattern of evolution at the
                      evolution. (a) Phyletic gradualism, where       species level could be studied from the fossil
                      evolution takes place in the lineages, and      record. These should have the following
                      speciation is a side effect of that evolution.   features:
                      (b) Punctuated equilibrium, where most
                      evolution is associated with speciation events,   1 Abundant specimens.
                      and lineages show little evolution (stasis).
                                                                      2  Fossils with living representatives, so that
                                                                         species can be identifi ed clearly.
                      2 In the punctuated equilibrium model (Fig.     3  Information on geographic variation, so
                         5.3b), with rectangular branches, almost        that rapid speciation events (punctuations)
                         no evolution takes place within species         could be distinguished from migrations in
                         lineages (they show stasis), and evolution      or out of the area.
                         is concentrated in the speciation events     4  Good stratigraphic control, in terms of
                         that coincide with major sideways shifts.       long continuous sequences of rocks
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