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398  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD





                                 Box 15.4  The age of crinoids: an Early Carboniferous diversity spike

                        Early Carboniferous (Mississippian) crinoids were abundant and diverse, so much so that this inter-
                        val is often called the “Age of crinoids”. Limestones of this age often consist of over 50% pelmato-
                        zoan debris, and are known as encrinites. Why then were crinoids so abundant at this time? Two
                        factors seem to have contributed to these extensive shoals of crinoids (Kammer & Ausich 2006).

                        Firstly, five major groups were in various states of recovery after the Frasnian-Famennian extinction
                        event, particularly the advanced cladids (Fig. 15.6). Secondly, with the disappearance of the shelf-
                        edge coral–stromatoporoid buildups at the end of the Devonian, platform geometries were quite
                        different. There was improved and unimpeded water circulation, which promoted stenohaline condi-
                        tions that encouraged the growth of crinoid communities. With new ecospace and a lack of predation
                        pressures, crinoid diversity exploded. Sadly, the good times came to an end with regression and the
                        cooler-water conditions associated with the Late Carboniferous glaciation. Crinoids were never again
                        so diverse.


                                                 200
                                                        2005
                                                        2002
                                                 160

                                               Number of genera  120


                                                 80


                                                 40


                                                  0
                                                     Loch  Prag  Emsi  Eife  Give  Fras  Fame  Tour  Vise  Serp  Bash  Mosc  Step


                        Figure 15.6  Diversity of Early Carboniferous crinoids. (From Kammer & Ausich 2006.)



                      wide range of strategies from a fi xed sessile   the Late Devonian, and were probably
                      mode to free-living recumbent styles. The dip-  replaced by the better adapted blastoids during
                      loporites were very widespread from the Early   the Silurian and Devonian.
                      Ordovician to the Early Devonian and prob-
                      ably evolved from a Late Cambrian blasto-       Blastoids
                      zoan ancestor.
                                                                      The extinct blastoids were small, pentamer-
                      Rhombifera  The    rhombiferans   appeared      ally symmetric animals with short stems
                      during the Late Cambrian equipped with bra-     and hydrospires adapted for respiration (Fig.
                      chioles and distinctive rhombic patterns of     15.8). They are represented by over 80 genera
                      respiratory pores crossing thecal plate sutures   in rocks of Silurian to Permian age. The
                      (Fig. 15.7). They are classified according to    blastoid cup or theca is usually globular and

                      the pattern and shape of their pores; these     composed of a ring of three basal plates, sur-
                      separate the order Dichoporita from the Fis-    mounted by a circle of fi ve larger radial plates.
                      tulipora. The rhombiferans became common        The mouth is often surrounded by fi ve large
                      during the Early Ordovician and continued       openings or  spiracles associated with the
                      with a near cosmopolitan distribution until     respiratory system. Although relatively rare,
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