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350  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD





                            Era     Period
                          0       Quaternary
                              Cenozoic  Tertiary       Nautiloidea  Teuthoidea  Sepioidea  Octopoda  Ammonoidea
                         1.5
                          60
                                  Cretaceous                                  Belemnitida   Lytoceratida  Ammonitida  Phylloceratida
                              Mesozoic  Jurassic
                         130                                                         Ancyloceratida

                         190
                                   Triassic
                         220                                                     Aulacoceratida             Ceratitida
                       Time (Ma)  280  Carboniferous                                                     Prolecanitida  Goniatitida
                                   Permian



                         340                       Actinoceratoidea           Bactritida  Clymeniida  Anarcestida
                              Paleozoic  Silurian
                         400       Devonian
                         440                   Endoceratoidea
                                  Ordovician
                         500                                                    Plectronoceras
                                   Cambrian
                         570
                      Figure 13.19  Stratigraphic ranges of the main ammonite taxa together with the other main cephalopod
                      groups. (Based on Ward, P. 1987. Natural History of Nautilus. Allen & Unwin, Boston.)


                      Ammonoid ecology and evolution                  broad cadicones. Evolute planulates and ser-
                                                                      penticones, together with small planulates,
                      The pioneer work by Arthur Trueman (Univer-     were probably pelagic in the upper parts of the
                      sity of Glasgow) on the buoyancy and orienta-   water column. However, most oxycones were
                      tion of the ammonite shell established the      restricted to shallow-shelf depths. Some het-
                      probable life attitudes for even the most bizarre   eromorphs were nektobenthonic, whereas a
                      heteromorph forms (Fig. 13.21a). Theoreti-      few floated in the surface waters.

                      cally, at least, virtually all ammonoids could    In many ammonite faunas the consistent
                      favorably adjust their attitude and buoyancy    co-occurrence of large and small similarly
                      in the water column. Most ammonoids were        ornamented mature shells at specifi c horizons
                      probably part of the mobile benthos, although   suggests that the macroconch and microconch
                      after death their gas-filled shells could be     may be related sexual dimorphs (the male and

                      widely distributed by oceanic currents. Many    female of the species). The macroconch was
                      groups of ammonoids were endemic, and the       probably the female, though this may not
                      shovel-like jaws of some groups were most       always have been the case. The ammonoids

                      efficient at the sediment–water interface.       probably originated from the bactritid ortho-
                      Richard Batt’s studies (1993) on Cretaceous     cone nautiloids, with protoconchs and large
                      ammonite morphotypes from the United States     body chambers, during the earliest Devonian.
                      have established a series of shell types related   The anarcestide goniatites, with simple sutural
                      to life modes and environments (Fig. 13.21b).   patterns, were relatively scarce during the
                      For example, evolute heavily ornamented         Mid Devonian. However, by the Famennian,
                      forms were probably nektobenthonic, as were     other groups such as the clymeniids, with a
                      spiny cadicones and spherocones, nodose         dorsally situated siphuncle, were common.
                      spherocones and platycones, together with       The goniatitides expanded during the
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