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142  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD


                                                                      positive allometry is in the antlers of the Irish
                        100
                                                                      deer (Box 6.2), and indeed in many other
                                                                      sexually selected features that are minute or
                                                                      absent in the juvenile, but very large in the
                       Orbit length (cm)  10                          adult.
                                                                        Allometry is commonly considered only in
                                                                      the context of ontogeny, the growth from egg
                                                                      or embryo through juvenile to adult. But
                                                                      studies of form may compare species, and
                                                                      shape variation can be accounted for in an
                                                                      evolutionary context too. For example, a
                         1           10                   100
                                                                      comparison of species of antelope would
                                        Skull length (cm)
                      (a)                                             show positive allometry in leg width: scaled
                                                                      against body length, the sturdiness or width
                         50                                           of the leg increases positively allometrically.
                                                                      This is because of the well-known biological
                                                                      scaling principle: some organs and functions
                        Skull length (cm)                             sional measure), whereas others relate to body
                                                                      relate to the mass of an animal (a three-dimen-
                                                                      length or body outline (one- and two-dimen-
                         10
                                                                      sional measures). As body mass (three-dimen-
                                                                      sional) increases, the diameter of the legs
                                                                      (two-dimensional) increases in proportion to
                                                                      support the added weight. So, in body outline,
                          10                           100
                                                                      small antelope have extremely slender legs,
                                     Backbone length (cm)             and larger ones have relatively more massive
                      (b)
                                                                      legs.
                      Figure 6.4  Tests of allometry in the ichthyosaur   These aspects of allometry may be under-
                      Ichthyosaurus. (a) Plot of orbit length against   stood in terms of the allometric equation,
                      skull length, and (b) plot of skull length against
                      backbone length. The Somerset embryo (Fig.                        y = kx a
                      6.3b) is indicated by a solid circle. Both graphs
                      show negative allometry (orbit diameter = 0.355   where y is the measurement of interest (e.g.
                      (skull length) 0.987 ; skull length = 1.162 (backbone   head length, eye diameter), x is the standard
                      length) 0.933 ), confirming that embryos and     of comparison (e.g. body length), k is a con-

                      juveniles had relatively large heads and eyes.   stant and a is the allometric coeffi cient. The
                      (Courtesy of Makoto Manabe.)                    constant k is calculated using the allometric
                                                                      equation for each particular case. The allome-

                                                                      tric coeffi cient  a defines the nature of the
                      ratio of eye diameter to body length dimin-     slope: if  a  = 1, the slope is at 45˚ and this
                      ishes as the animal approaches adulthood.       defines a case of isometric growth; if a > 1,

                      This is an example of  allometric (“different   we have positive allometry, and if a < 1, we
                      measure”) growth. If there is no change in      have negative allometry (see Fig. 6.4).
                      proportions during growth, the feature is said    After the nature of any allometric change
                      to show isometric (“same measure”) growth.      of parts or organs has been established quan-
                        Allometric growth is commoner than iso-       titatively, it is possible to investigate why such
                      metric. Positive allometry is when the organ    changes might occur. The large eyes and small
                      or feature of interest increases faster than the   noses of babies are said to make them look
                      isometric expectation, and negative allometry   cute so their parents will look after them, and
                      is when growth of the structure of interest is   feed them. But the fundamental reason is pre-
                      slower than isometry. Head and eye size         sumably because the eye is complex and is at
                      usually show negative allometry, starting rela-  nearly adult size in the baby for functional
                      tively large in the juvenile, and becoming rela-  reasons, and the relatively large head of a
                      tively smaller in the adult. An example of      human baby is to accommodate the large
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