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364  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                      appendages that have lateral branches devel-    14.3), with the facial or cephalic suture sepa-

                      oped from the walking limbs. The trilobito-     rating the inner fixed and the outer free cheeks.
                      morphs include mainly the trilobites and over   Although many trilobites lacked eyes, this
                      15,000 species are known. Trilobites were a     may be a secondary condition; despite loss
                      unique and very successful arthropod group,     of vision the cephalic sutures remained. The
                      common throughout the Paleozoic until their     sutures themselves are very important for
                      extinction at the end of the Permian. There is   understanding the functional morphology

                      no doubt that the trilobites are one of the     and classification of the group. There are four
                      most attractive fossils groups, much prized by   main types of suture (Fig. 14.5): the proparian
                      both amateur and professional collectors        mode extends posteriorly in front of the genal
                      alike. Some of the earliest arthropods were     angle, whereas the opisthoparian mode cuts
                      trilobites, and marine Cambrian strata (as      the posterior margin of the cephalon behind
                      well as many later deposits) are usually cor-   the genal angle, and the gonatoparian suture
                      related on the basis of trilobite assemblages.   bisects the genal angle and lateral sutures
                      The group formed an important part of the       follow the margin of the cephalon. In the
                      mobile benthos, although a few groups were      rarer metaparian condition the suture extends
                      adapted to pelagic life modes.                  from near the genal angle on the posterior
                                                                      margin, around the eye to fi nish farther along
                                                                      the same margin.
                      Trilobite morphology
                                                                        On the ventral surface, underneath the
                      The trilobite exoskeleton (Fig. 14.3), as the   cephalon, three plates were associated with
                      name trilobite (“three-lobed”) suggests, is     the anterior soft parts including the mouth.
                      divided longitudinally into three lobes; the    The  rostral plate is situated at the anterior
                      axial lobe protects the digestive system,       margin. Posterior to the rostral plate, the
                      whereas the two pleural lobes cover append-     hypostome, a plate of variable shape and size,
                      ages. In virtually all trilobites a well-defi ned   is usually sited under the glabella. The shape
                      cephalon, thorax and pygidium are devel-        and position of the hypostome is of great
                      oped; the trilobite exoskeleton is composed     help in classifying the group. The small metas-
                      almost entirely of calcite.                     toma is known from only a few taxa and
                        The cephalon has a raised axial area, the     apparently lay behind the mouth. The dorsal
                      glabella, with a series of glabellar furrows.   margin was protected by a ventral fl ange or
                      Eyes are commonly developed laterally (Box      doublure.








                                 Box 14.2 Ediacaran arthropods?


                        Are they or aren’t they? Some paleontologists believe they can identify some of the Ediacaran animals
                        as arthropods or proto-arthropods; others dispute this. Parvancorina (Fig. 14.2), for example, is a
                        possible candidate, with its shield-shaped outline, strong axial ridge and arched anterior lobes,

                        together with a convex profile. It really looks like a juvenile trilobite molt stage, but did not have a
                        mineralized skeleton. Not convinced? Beautifully preserved fossils from a new Cambrian Lagerstätte,


                        the Chinese Kaili fauna seem to confirm it. Specimens of the genus Skania, first described from the
                        Burgess Shale, have many similarities to Parvancorina but this genus has an exoskeleton and a better-
                        defined cephalon and dorsal trunk (Lin et al. 2006). Skania together with Parvancorina and Primi-

                        caris may have formed a sister group to the Arachnomorpha (which includes the spiders). Moreover

                        this relationship establishes a Proterozoic root for the Arthropoda and is the first arthropod crown
                        group that demonstrably ranges through the Precambrian–Cambrian boundary. A Proterozoic origin
                        for the arthropods may help pin down more precisely their time of divergence from the last common
                        ancestor of the arthropods and priapulid worms.
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