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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-