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390  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                      The echinoderms and hemichordates appear        and out in the deep sea the ocean fl oors are
                      to be two very different groups of animals,     covered by brittle stars and sea cucumbers.
                      one characterized by fi ve-fold symmetry and     The phylum Echinodermata has an unusual

                      a water vascular system, the other a group of   five-fold symmetry and is uniquely equipped
                      odd stick-like colonial organisms. Surpris-     with a water vascular system in which water
                      ingly, both are closely related to each other   is forced around the plumbing by muscular
                      and, moreover, are not so distant from our-     action, while tube feet, extending from the

                      selves, the chordates. Both groups are deu-     system, are often modified for food process-

                      terostomes; the first opening to develop in the   ing, locomotion and respiration. The 6000 or
                      embryo is the anus and a second forms the       so living echinoderm species include familiar
                      mouth. The group has a dipleurula larva and     forms such as sea lilies, sea urchins, sand
                      a body cavity that developed from an exten-     dollars, starfish and sea cucumbers (Fig. 15.1).

                      sions of the embryonic gut (see p. 240).        Although many species today live in the inter-
                      Modern morphological and molecular analy-       tidal or subtidal zones, the group is most
                      ses indicate that the echinoderms and hemi-     diverse in the deep sea. Echinoderms also
                      chordates are in fact sister groups (Smith      occupied a wide range of marine environ-
                      2005). A small, extinct group – the Vetulicolia   ments and pursued a variety of life strategies
                      – so far known only from the Cambrian, has      in the geological past. Fossil echinoderms
                      also been related to the deuterostomes because   are relatively common, and because many
                      of similar gill structures and the absence of   echinoderm skeletons disintegrate rapidly

                      limbs. But although recent finds from Utah       after death, many limestones are packed with
                      have suggested that this group has more in      the distinctive skeletal debris of calcitic
                      common with the arthropods and probably         plates.
                      belongs to the ecdysozoans (see p. 361), the      Apart from the water vascular system, echi-
                      group remains an enigma (see Box 15.10).        noderms have a number of other distinctive
                                                                      features. All members of the phylum have a
                                                                      mesodermal skeleton constructed from porous
                      ECHINODERMS
                                                                      plates of calcite; each plate is usually a single
                        Clearly we stood among the ruins of           crystal of calcite and easy to recognize in thin
                        some latter-day South Kensington! Here,       sections. In addition, the plates have a unique
                        apparently, was the Palæontological           ultrastructure of rods linked to form a three-
                        Section, and a very splendid array of         dimensional lattice. This network, or stereom,

                        fossils  it  must  have  been.  .  .  .  The  place   is permeated by finger-like pieces of soft tissue
                        was very silent. The thick dust deadened      that occupy the spaces, or stroma, in the

                        our footsteps. Weena, who had been            lattice. Finally, five-rayed or pentameral sym-
                        rolling a sea urchin down the sloping         metry, occasionally modified by a secondary

                        glass of a case, presently came, as I stared   bilateral symmetry, is typical of the echino-
                        about me, and very quietly took my hand       derms. The phylum is generally split into the
                        and stood beside me. And at fi rst I was       mobile, non-stalked  eleutherozoans and the
                        so much surprised by this ancient monu-       mainly fi xed,  stalked  pelmatozoans (Box
                        ment of an intellectual age, that I gave      15.1), but the earliest forms are hard to
                        no thought to the possibilities it pre-       classify (Box 15.2).
                        sented. Even my preoccupation about the         The multiplated echinoderm skeleton disin-
                        Time Machine receded a little from my         tegrates very rapidly after death; although
                        mind.                                         individual plates or ossicles have high preser-
                                                                      vation potential, the complete skeletons do
                          H. G. Wells (1898) The Time Machine         not. Nevertheless, occasionally rapid burial or
                                                                      transportation into anoxic conditions may
                                                                      result in the preservation of complete echino-
                      Echinoderms today are one of the most abun-     derm skeletons. Starfish beds, usually charac-

                      dant marine animal groups, and as fossils they   terized by accumulations of complete
                      can sometimes be rather robust, as Weena        echinoderms, occur sporadically throughout
                      from The Time Machine found. Sea urchins        the fossil record. The Leintwardine Starfi sh
                      are common in many intertidal environments,     Bed of the England–Wales border area con-
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