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362  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD











                                                                                         Crustacean:
                                                                                          decapod
                                                       Chelicerate:    Crustacean:
                                   Trilobitomorph:      scorpion        decapod
                                     trilobite






                                     Crustacean:       Crustacean:
                                      ostracode         barnacle         Insect:
                                                                       palaeopteran         Insect:
                                                                                          coleopteran
                      Figure 14.1  Some of the main arthropod groups: a variety of forms based on a simple body plan of a
                      tough exoskeleton and jointed limbs.



                                                                      not usually mineralized. Although many
                      ARTHROPODS: INTRODUCTION
                                                                      arthropods undergo metamorphosis, virtually
                      Arthropods are a very common and spectacu-      all the main groups grow by molting or
                      larly diverse group of legged invertebrates     ecdysis; first the endoskeleton is dissolved and

                      accounting for about three-quarters of all      second the old exoskeleton is detached along
                      species living on the planet today, largely     sutures while the new exoskeleton is gener-
                      because of the phenomenal abundance of the      ated.  Exuviae, or cast-off coverings, are all
                      insects. The basic body plan – conspicuously    that remain of the previous skeleton or cuticle
                      segmented, with jointed appendages adapted      of the animal: one arthropod can thus produce
                      for feeding, locomotion and respiration –       many potential skeletal fossils in its lifetime.
                      together with a tough  exoskeleton, fi rst         During a geological history of at least 540
                      appeared during the Early Cambrian and has      million years, the fi ve subphyla of arthropods
                      since been exploited by a huge variety of living   (Box 14.1) have adapted to life in marine,
                      and fossil arthropods that pursue many life-    freshwater and terrestrial environments. For
                      styles. All members of this phylum have both    a long time their closest living relatives were
                      segmented bodies and appendages (Fig. 14.1);    thought to be the segmented annelid worms,
                      moreover the animal is differentiated into a    but new studies show that the closest sister
                      head, thorax and abdomen, with often the        group of arthropods is a clade of unsegmented
                      head and thorax fused to form the cephalo-      worms that includes the priapulids and the
                      thorax. The possession of mandibles, or hard    nematodes or round worms. Their segmenta-
                      mouthparts, equipped many arthropods with       tion may thus either have arisen independently
                      the ability to process a wide variety of        to that of the annelids, or may have been
                      foods.                                          inherited from a very deep ancestor to both
                        The arthropod exoskeleton is constructed      groups.
                      mainly from the organic substance chitin.
                      This is often hardened or sclerotized by
                      calcium carbonate or calcium phosphate, so      EARLY ARTHROPOD FAUNAS
                      the potential for preservation is excellent     A huge variety of bizarre arthropod types
                      across the group. The exoskeleton acts as a     formed much of the basis for the Cambrian
                      base for the attachment of locomotory           explosion (see p. 249). Over 20 groups of
                      muscles, permitting rapid movement, and is      arthropod have been described from the Mid
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