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ECDYSOZOA: ARTHROPODS      363





                        Box 14.1 Classifi cation of arthropods

               There are currently differences in the status given to the main arthropod groups. If the Arthropoda
               is in fact a superphylum, the following groupings are phyla. However, some authorities have assigned
               the following superclass status within the phylum Arthropoda. The classification here is a compro-

               mise. Basal to the phylum are a number of minor but evolutionarily important groups, such as the
               tardigrades (water bears), that are now known from the Cambrian.

               Subphylum TRILOBITOMORPHA
               •  Trilobites and their relatives; animals with a cephalon, thorax and pygidium; the body, length-
                  wise, has an axial lobe and two lateral pleural lobes
               • Cambrian to Permian
               Subphylum CHELICERATA
               •  Large group with a body divided into two tagmata; the prosoma (which bears six pairs of append-
                  ages, the first being the chelicerae or pincer-like appendages, giving the group its name), and the

                  opisthosoma with an extended tail or telson
               • ?Cambrian to Recent
               Subphylum MYRIAPODA

               • Includes the flexible centipedes together with the millipedes
               •  ?Ordovician, Silurian to Recent
               Subphylum HEXAPODA
               •  Highly-diverse group, with a head, thorax and abdomen and six legs; includes the ants, beetles,

                  dragonfl ies, flies and wasps
               • Devonian to Recent
               Subphylum CRUSTACEA
               •  Includes the bivalved phyllocarids; Early Paleozoic taxa were ancestral to the crabs, shrimps and
                  lobsters
               • Cambrian to Recent




             Cambrian Burgess Shale and related deposits     of morphological disparity not really exceeded
             (see Box 14.8); some have even been assigned    during the next 500 million years of evolu-
             to new phyla, emphasizing the expansive         tion. Moreover, our knowledge of the Cam-
             nature of the explosion, truly evolution’s “big   brian arthropod record, particularly that of
             bang”. Stephen Jay Gould, in his bestseller     soft-bodied organisms, is probably not nearly
             Wonderful Life argued that morphological        as complete as that of the modern fauna and
             disparity during the Cambrian was greater       we should expect further surprises as more
             than at any time since. Nevertheless, cladistic,   Cambrian Lagerstätten are investigated (Box
             and phenetic analyses of both morphological     14.2).
             and taxonomic criteria suggested otherwise
             (Briggs et al. 1993). Rather, the morphologi-
             cal disparity among the Cambrian arthropods     SUBPHYLUM TRILOBITOMORPHA
             is not markedly different from that seen across   The trilobitomorphs are highly derived arthro-
             living taxa, they just look stranger to us. But   pods lacking specialized mouthparts, and
             it is nonetheless remarkable that very early in   with tagmata comprising the cephalon, thorax
             their history arthropods attained high levels   and  pygidium, together with trilobitomorph
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