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354  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                        By analogy with modern squids, the belem-     shell beds (Fig. 13.23). These accumulations

                      nites were probably rapidly-moving predators    conform to five genetic types (Doyle & Mac-
                      living in shoals with their body level regulated   Donald 1993): (1) post-spawning mortalities
                      by the guard. The animal thus probably main-    (Fig. 13.23a); (2) catastrophic mass mortali-
                      tained a horizontal attitude within the water   ties; (3) predation concentrates, either in situ
                      column, preferring the open ocean. Data from    or regurgitated (Fig. 13.23b); (4) condensa-
                      the stomach contents of ichthyosaurs confi rm    tion deposits perhaps aided by winnowing
                      that these mollusks formed part of their        and sediment by-pass; and (5) resedimented
                      diet.                                           deposits derived from usually condensed
                        Some of the oldest records of belemnites,     accumulations. Many of these so-called bel-

                      for example  Jeletzkya from the mid-            emnite battlefields are then partly natural
                      Carboniferous of Illinois, are tentative. The   occurrences, reflecting the biology of the


                      first unequivocal belemnites are from the        animals (numbers 1–3), but it is important to
                      Middle Triassic rocks of Sichuan Province,      distinguish these from sedimentary accumula-
                      China where several species of Sinobelemnites   tions (numbers 4 and 5) that say nothing
                      occur. Belemnites became extinct at the end     about belemnite behavior.
                      of the Cretaceous; later records are reworked
                      or based on misinterpretations.
                        Some of the first supposed belemnites, like    CLASS SCAPHOPODA

                      the Carboniferous Paleoconus, were relatively   Scaphopods are generally rare as fossils. The
                      short stubby forms. In the Early Jurassic,      Scaphopoda, or elephant-tusk shells, have a
                      Megateuthis was a long, slender form, whereas   single, slightly curved high conical shell, open
                      Dactyloteuthis was laterally fl attened;  the    at both ends (Fig. 13.25a). They lack gills and
                      later Jurassic  Hibolites is spear-shaped. The   eyes, but have a mouth equipped with a radula
                      Cretaceous  Belemnitella has a large bullet-    and surrounded by tentacles; they also possess
                      shaped guard, whereas that of Duvalia has a     a foot, similar to that of the bivalves, adapted

                      flattened spatulate shape (Fig. 13.22d).         for burrowing. Scaphopods are mainly car-
                      However, despite differences in the detailed    nivorous, feeding on small organisms such as
                      morphology of the endoskeltons across           foraminiferans and spending much of their
                      genera, many authorities consider that most     life in quasi-infaunal positions within soft
                      of the Mesozoic belemnites probably looked      sediment in deeper-water environments. The
                      very similar, but there are still enough features   first scaphopods appeared during the Devo-

                      to measure on their skeletons and discrimi-     nian and apparently had similar lifestyles to
                      nate taxa (Box 13.8).                           living forms such as Dentalium.
                        The compact calcareous guards of the bel-
                      emnites have proved ideal for the analysis of
                                                   18
                                               16
                      oxygen isotope ratios (O  : O ) relating to     CLASS ROSTROCONCHA
                      paleotemperature conditions in the Jurassic     Relatively recently a small class of mollusks,

                      and Cretaceous seas. These data have indi-      superficially resembling bivalves but lacking
                      cated warm peaks during the Albian and the      a functional hinge, has been documented
                      Coniacian-Santonian (mid-Cretaceous) with a     from the Paleozoic. Over 35 genera have been
                      gradual cooling from the Campanian (Late        described; most were originally described as
                      Cretaceous) onwards. And as with many other     bivalved arthropods. The rostroconchs prob-
                      Mesozoic groups, belemnite distributions        ably had a foot that emerged through the
                      show separate low-latitude Tethyan and high-    anterior gape between the shells. However,
                      latitude Boreal assemblages.                    the two shells are in fact fused along the mid-
                        Spectacular mass accumulations of belem-      dorsal line, and posteriorly the shells are
                      nite rostra are relatively common in Mesozoic   extended as a platform or rostrum (Fig.
                      sediments and, although some authors have       13.25b). Ontogeny occurs from an initial dis-
                      used these assemblages in paleocurrent studies,   soconch with the bilobed form developing
                      few have addressed their mode of accumula-      from the disproportionate growth of shell
                      tion. Dense accumulations of bullet-shaped      from the lateral lobes of the mantle. The

                      belemnite rostra have promoted the term         group appeared first during the Early Cam-

                      “belemnite battlefields” for such distinctive    brian when, for example, Heraultipegma and
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