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146  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD



                                     Rec.
                                     Ple.
                                      Miocene                    Pliocene  N. nigricans  T. doederleini













                                                       T. thomsoni

                                      Oligocene  N. antipoda



                                                                  T. coelata



                                      Paleocene                          Eocene  ontogeny  T. squamosa
                                                              pedomorphocline








                                                           T. boongeroodaensis
                                           juvenile
                                                                                         adult

                      Figure 6.7  Heterochronic evolution in the Cenozoic brachiopods Tegulorhynchia and Notosaria.
                      Adults of more recent species are like juveniles of the ancestor. Hence, pedomorphosis (“juvenile
                      formation”) is expressed in this example. (Based on McNamara 1976.)


                        In studying heterochrony, it is necessary to   sis, and hyperomorphosis in particular (devel-
                      have a robust phylogeny of the organisms        opmental time is longer, but rate of morpho-
                      in question, an adequate fossil record of       logical development is not faster). Thus,
                      the group, and a sound set of ontogenetic       heterochronic changes can occur in different
                      sequences for each species. This allows the     directions in different characters, a phenome-
                      paleontologist to compare juveniles and adults   non called mosaic evolution.
                      throughout the phylogeny. A classic example       In a classic study, McNamara (1976) sug-
                      is human evolution. It seems obvious that       gested that species of the Cenozoic brachio-
                      human adults look like juvenile apes, with      pod  Tegulorhynchia evolved into  Notosaria
                      their flat faces, large brains and lack of body   by a process of heterochrony (Fig. 6.7). The

                      hair. These would imply a pedomorphic           main changes were a narrowing of the shell,
                      change in humans with respect to the human/     a reduction in the number of ribs in the shell
                      ape ancestor. However, other characters do      ornament, a smoothing of the lower margin,
                      not fi t this pattern. For example, developmen-  and an enlargement of the pedicle foramen
                      tal time in humans is far longer than in apes   (the opening through which a fl eshy  stalk
                      and ancestral forms, a feature of peramorpho-   attaches the animal to a rock). These changes
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