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330  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD





                                 Box 13.2  Kimberella and Odontogriphus join the mollusks

                        A modest-sized, disk-shaped fossil from the Late Precambrian, named Kimberella in 1959, has suf-
                        fered mixed fortunes. First described from the Ediacaran rocks of Australia as a jellyfi sh and later
                        a cubozoan, Mikhail Fedonkin and Ben Waggoner (1997) then reconstructed Kimberella as a bilater-
                        ally symmetric, benthic crawler with a non-mineralized, single shell, on the basis of new material
                        from the White Sea, Russia. Kimberella is linked with a variety of trace fossils suggesting mobility
                        and a feeding strategy that must have involved a radula. The body fossils and trace fossils place
                        Kimberella near the base of the molluskan clade and suggest a deep origin for the phylum (Fig. 13.2),

                        and for the bilateralians, significantly earlier than the Cambrian explosion. But who were its closest
                        relatives? A new investigation by Jean-Bernard Caron and his colleagues (2006) offers some clues.
                        They studied another enigmatic animal, Odontogriphus from the Burgess Shale. Odontogriphus had
                        previously been allied with the brachiopods, bryozoans, phoronids and even early vertebrates. The
                        new study shows that Odontogriphus possesses a radula, a broad foot and a stiffened dorsum, so
                        placing it firmly within the mollusks, close to Kimberella, together with Wiwaxia (another enigmatic

                        soft-bodied organism covered with possible scierites), which also possesses a radula, and another
                        enigma, Halkieria (Box 13.3).





















                               (a)            (b)
                                                                                         Annelida
                                    total-group Mollusca
                                                          Kimberella?
                                    putative range extension                  Odontogriphus
                                    stem-group Mollusca                       Wiwaxia
                                    and known fossil range                    Halkieriid
                                    crown-group Mollusca                 ???            Neomeniomorpha
                                    and known fossil range                              Polyplacophora
                                                                                        Other crown-group
                                                                                        Mollusca
                                                     Firm                       Substrate                               Soft
                               Cambrian substrate revolution:  Mat-based ecology
                                                                    Vertical burrowers
                                                               N-D T A B/T  Mid  Late
                                         Ediacaran                 Early                  Ordovician
                                                                      Cambrian
                                                        555        542                                                501          488                     Time (Ma)
                               (c)
                        Figure 13.2  The early mollusks (a) Kimberella, (b) Odontogriphus and (c) phylogeny and
                        stratigraphic ranges of early mollusks mapped onto some ecological changes. N-D, Nemakit-
                        Daldynian; T, Tommotian; A, Atdabanian; B/T, Botomian. (a, courtesy of Ben Waggoner; b, c,
                        courtesy of ten-Bernard Caron.)
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