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ORIGIN OF THE METAZOANS  235


             ORIGINS AND CLASSIFICATION                      The fi rst metazoans: when and what?

             When did the first complex animals, the meta-    Life on our planet has been evolving for nearly
             zoans, appear on Earth and what did they        4 billion years. Molecular data suggest meta-
             look like? How could complex, multicelled       zoans have probably been around for at least
             animals evolve from the undifferentiated        600 myr (Fig. 10.1), during which time,
             single-celled organisms of most of the Pre-     according to some biologists, as many as 35
             cambrian? Why did they take almost 4            separate phyla have evolved. Five lines of evi-

             billion years to appear? These questions have   dence have figured prominently in the search
             puzzled scientists, including Charles Darwin,   for the earliest metazoans: body fossils, trace
             for over two centuries. In the last few decades   fossils, fossil embryos, the molecular clock
             a range of multidisciplinary techniques, from   and biomarkers.
             molecular biology to X-ray tomography, has        Much controversy still surrounds the timing
             helped generate new testable hypotheses         of their origin. Was there a long cryptic inter-
             regarding the origins of our early ancestors.   val of metazoan evolution prior to the Edia-
             Apart from the fossil evidence of metazoan      caran – a time when we do not fi nd  fossils
             body and trace fossils, the investigation of    preserved, either because the animals lacked
             minute fossil embryos, carefully calibrated     preservable bodies, or they were small, or
             molecular clocks and more recently biomark-     perhaps a combination of both? Or, as the
             ers have placed the investigation of Precam-    recalibrated molecular clocks suggest, can
             brian life at the top of many scientifi c        animal origins be tracked back only to the
             agendas.                                        Ediacaran, when there was also a sudden rise





                                                                          C              Crinozoa
                                                           535                          Deuterostomia  Eleutherozoa
              200      metazoan orders                                                   Hemichordata
                       metazoan classes
                                                                          C              Pteriomorpha
                    604  molecular clock estimates             C                         Nuculoida
              160    C molecular clock calibration  579                                  Vetigastropoda
                          feeding larva                  545            C
                          non-feeding larva             549           493               Spiralia  Sorbeochoncha
              Number of classes/orders  80  604     561  546          494               Ecdysozoa  Spionidea + Serpulidea
                          Oman biomarker record
                                                                                         Phyllodicida
              120
                                                                                         Hoplonemertea
                                                                                         Heteronemertea
                                                                                         Arthropoda
                                 634
                                                                                        “Porifera” Cnidaria  Calcispongia
                                                                                         Anthozoa
                                                         543                             Priapulida
                         664
               40                                                                        Hydrozoa

                                                            N-D T AB/T  M Late  Early M  Late  Demospongia
                       Cryogenian           Ediacaran
                                                              Early
                                                               Cambrian     Ordovician
                700               635      600    575  555 542    513 501 488  472 461  443 Ma




             Figure 10.1  Time scale and tempo of early animal evolution: the key metazoan groups are shown with
             the putative age of their last common ancestor, together with an estimate of the respective numbers of
             classes and orders indicated against a stratigraphy indicating key biological and chemical events. N–D,
             Nemakit-Daldynian; T, Tommotian; A, Atdabanian; B/T, Botomian. (Courtesy of Kevin Peterson.)
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