Page 320 - Introduction to Paleobiology and The Fossil Record
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SPIRALIANS 1: LOPHOPHORATES  307














                                (a)












                                (b)
             Figure 12.8  Teeth of articulated brachiopods: (a) deltidiodont and (b) cyrtomatodont dentition.


               The brachiopods experienced fi ve  main          Not unexpectedly, the end-Permian mass
             extinction events followed by recoveries and    extinction saw the demise of over 90% of
             radiations of varying magnitudes. The end-      brachiopod species, including some of the
             Ordovician event occurred in two phases         most ecologically and taxonomically diverse
             against a background of glaciation and          groups. The post-extinction fauna was fi rst
             accounted for the loss of almost 80% of bra-    dominated by a variety of disaster taxa (see
             chiopod families. The recovery and subse-       p. 179), including lingulids; nevertheless the
             quent radiation is marked by the decline of     brachiopod fauna later diversifi ed  within  a
             deltidiodont groups such as the orthides and    relatively few clades dominated by the rhyn-
             strophomenides, whereas the spire-bearing       chonellides and terebratulides. The end-Trias-
             atrypides, athyridides and the spiriferides     sic event removed the majority of the remaining
             with cyrtomatodont dentition (Fig. 12.8),       spiriferides and the last strophomenides. The
             together with the pentamerides, achieved        agenda set by the end-Permian event, involv-
             greater dominance, particularly in carbonate    ing the subsequent dominance of rhynchonel-
             environments. Late Devonian events, at the      lide and terebratulide groups, was continued
             Frasnian–Famennian Stage boundary, were         after the end-Triassic event. The end-Creta-
             also associated with climate change and         ceous event may have been responsible for the
             removed the atrypides and pentamerides and      loss of about 70% of chalk brachiopod faunas
             severely affected the orthides and stropho-     in northwest Europe; nevertheless, many
             menides, whereas the spiriferides and           genera survived to diversify again in the
             rhynchonellides survived in deeper-water        Danian limestones. Despite the post-Permian
             environments and staged an impressive recov-    decline of the phylum, Modern brachiopods
             ery. A particular feature of the post-Frasnian   exhibit a remarkable range of adaptations
             fauna was the diversity of recumbent brachio-   based on a simple body plan and a well-

             pod megaguilds (see p. 91), dominated by the    defined role in the fi xed, low-level benthos.
             productides. The Carboniferous and particu-
             larly the Permian were intervals of spectacular
             experimentation: some brachiopods mim-          Ecology: life on the seabed
             icked corals or developed extravagant clusters   Living and fossil brachiopods have developed
             of spines while a number of groups reduced      a wide range of lifestyles (Fig. 12.9). Most
             their shells, thus presenting soft tissues to the   were attached by a pedicle cemented to a hard
             outside environment.                            substrate or rooted into soft sediment. A
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