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106  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD





                               Box 4.6 Shell concentrations

                        Shell concentrations of various types can tell us a huge amount about environments of deposition
                        but also can act as a proxy for biological productivity through time (Kidwell & Brenchley 1994).
                        Moreover, it is possible that evolutionary changes in the diversity and ecology of organisms that
                        produce and destroy calcareous skeletons suggest that the nature of these concentrations may have
                        changed through the Phanerozoic. Data from marine siliciclastic rocks, silicate-based clastic sedi-

                        ments, of Ordovician-Silurian, Jurassic and Neogene ages show a significant increase in the thickness
                        of densely packed bioclastic concentrations, from thin-bedded brachiopod-dominated concentrations
                        in the Ordovician-Silurian to a mollusk-dominated record with more numerous and thicker shell
                        beds in the Neogene (Fig. 4.22). Jurassic shell beds vary in thickness depending on whether they

                        have Paleozoic or modern affinities as the main components. This suggests that the Phanerozoic
                        increase in shell-bed thickness was not controlled by diagenesis or by a shift in taphonomic condi-
                        tions on the seafl oor, but rather by the evolution of biogenic clast producers, themselves – i.e. groups

                        with, firstly, more durable low-organic skeletons, secondly, greater ecological success in high-energy
                        environments, and thirdly higher rates of carbonate production. These results indicate that (i) repro-
                        ductive and metabolic output has increased in benthic communities over time; and (ii) the scale of
                        time averaging in benthic assemblages has increased owing to greater hard-part durability of modern
                        groups. New data, however, suggest that brachiopods were probably just as durable as mollusks,
                        but their communities simply did not produce so many shells. The frequency and thickness of shell
                        beds through time may simply be down to the relative biological productivity of different groups of
                        organisms.



                                                8
                                                        Modern fauna
                                                        Paleozoic fauna
                                                4
                                                                        oysters  oysters other epifaunal bivalves  infaunal bivalves  cheilostomes
                                               Thickness (m)  2       other mollusks


                                                       brachiopods  archaeogastropods stenolaemate bryozoans  brachiopods  cephalopods

                                                1


                                                0
                                                   0     100%    0     100%    0     100%
                                                    Ord-Sil        Jurassic     Neogene
                                                    (n = 85)      (n = 139)     (n = 218)
                                                                  % of shellbeds
                        Figure 4.22  Thicknesses of shell concentrations during the Ordovician-Silurian, Jurassic and
                        Neogene. Thick shell beds are a phenomenon of the Modern fauna, mainly generated by
                        bivalves. (From Kidwell & Brenchley 1994.)
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