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372  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD






                                                                             pelagic – swimming
                                                                             and floating



                                                                                  molting
                                       mobile
                                       nektobenthos                feeding

                                                                                 resting
                                                enrolled       molt stage






                                                  infaunal – living in burrows
                      Figure 14.9  Lifestyles of the trilobites: a mosaic of selected Lower Paleozoic trilobites in various life
                      attitudes.


                      abundance and diversity. Not surprisingly the     Trilobites show a number of evolutionary
                      Trilobitomorpha has been a major source of      trends. Through time, for example, those tri-
                      evolutionary data and there have been many      lobites that adopted enrolment as a defensive
                      studies on the functional morphology of the     strategy became better at it: the spines and
                      group (e.g. Bruton & Haas 2003).                sockets around their exoskeletons came to fi t
                        Trilobites have provided key evidence in      and lock better and better. Early trilobites
                      studies of macroevolution, especially in the    probably rolled up into a rough ball, but
                      controversy over punctuated equilibrium and     could be prized apart by a persistent
                      phyletic gradualism (see Chapter 5). Trilo-     predator; later enrolling trilobites were impen-
                      bites have complex morphologies that can be     etrable. There was a reduction in the size of
                      easily measured and analyzed statistically      the rostral plate and in some groups there was
                      (Box 14.4). The studies of Niles Eldredge and   an increase in spinosity and a trend from
                      Stephen Jay Gould on the number of lens fi les   micropygy to isopygy. The evolution of schizo-
                      of the Devonian trilobite Phacops rana formed   chroal visual systems appeared, by pedomor-
                      the basis for their punctuated equilibrium      phosis, during the Early Ordovician in the
                      model. On the other hand Peter Sheldon’s        phacopids.
                      investigation of over 15,000 specimens of
                      Mid Ordovician trilobites demonstrated
                      gradual changes in the number of pygidial       Trilobite abnormalities and injuries
                      ribs, possibly a slower, adaptive, fi ne-tuning   Trilobites have left a rich record of abnor-
                      to more stable environments. Euan Clarkson’s    malities and injuries, some evidence that they
                      survey of microevolutionary change in Upper     faced problems during ecdysis and that they
                      Cambrian olenid trilobites from the Alum        were attacked by predators (Fig. 14.14). There
                      Shales of Sweden provided evidence of similar   are three main types of abnormality (Owen
                      gradual change (Fig. 14.13). Macroevolution-    1985):
                      ary change in trilobites was effected by heter-
                      ochrony (see p. 145). Pedomorphosis during      1  Injuries sustained during molting.
                      ontogeny of the animal as a whole or applied    2  Pathological conditions resulting from
                      to particular organs such as the eyes gener-       disease and parasitic infestations.
                      ated new species and new biological             3  Teratological effects arising through some
                      structures.                                        embryological or genetic malfunctions.
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