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394  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD





                               Box 15.3 Columnal classifi cation

                        The majority of crinoid assemblages are represented by disarticulated ossicles. Conventional taxon-
                        omy based on a description of complete, articulated specimens is thus not possible. Nevertheless,

                        ossicles have many distinctive features, arguably with more well-defined characteristics than many
                        groups of macrofossils (Fig. 15.3). Single stems consist of many ossicles with a central canal or lumen
                        usually carrying nerve fibers. Both the ossicles and lumens have distinctive shapes that are the basis

                        of a form taxonomy of the group. Form taxonomy helps us classify the shapes of fossils, in the same
                        way that we can classify nuts and bolts. It is a useful method of organizing our data, but since it is
                        not biologically meaningful, cannot be used in phylogenetic analyses. Stems may be either homeo-
                        morphic, composed of similarly shaped ossicles, or heteromorphic with a variety of different-shaped
                        ossicles. Moreover stems may be subdivided into zones that may be internally homeomorphic or
                        heteromorphic. Columnal taxonomy has proved useful in describing taxa (so-called col. taxa) of
                        pelmatozoan, particularly crinoid, ossicles of stratigraphic signifi cance.

















                                     (a)                            (b)














                                                    (c)
                        Figure 15.3  Some crinoid ossicle types. (a) Articular facet of a columnal of the bourgueticrinid
                        Democrinus (?) sp., with a fulcral ridge of the synarthrial articulation; the lumen opens at the
                        bottom of the “8”-shaped depression (×15). (b) Cirral scar on a nodal of the isocrinoid Neocrinus
                        with well-preserved stereom microstructure and knob-like synarthrial fulcrum (×18). (c) Articular
                        facet of a columnal of the isocrinoid Neocrinus with symplectial articulation around the fi ve
                        petal-like areola areas (×9). (Courtesy of Stephen Donovan.)




                      units within a deep-water mudstone sequence.
                      Entire crinoids, cystoids, echinoids and cal-   Crinoidea
                      cichordates were carried downslope and          Although the crinoids, famously called “sea
                      rapidly buried on the unstable slopes of a      lilies”, look more like plants than animals,
                      submarine fan system.                           there is no doubt they are animals, and echi-
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