Page 158 - Introduction to Paleobiology and The Fossil Record
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FOSSIL FORM AND FUNCTION  145


             passes through the major nodes of the clado-    of heterochronic change,  pedomorphosis
             gram of vertebrates. The synapomorphies (see    (“juvenile formation”), or sexual maturity in
             p. 130) of vertebrates appear fi rst, then those   a juvenile body, and peramorphosis (“overde-
             of tetrapods, then those of amniotes, then      velopment”), where sexual maturity occurs
             those of mammals, of primates, and of the       relatively late. These changes can each occur
             species Homo sapiens last.                      in three ways, by variation in timing of the
               Three other aspects of development throw      beginning of body growth, the timing of
             light on phylogeny. Certain developmental       sexual maturation or the rate of morphologi-
             abnormalities called  atavisms, or throw-       cal development (Table 6.1).
             backs, show former stages of evolution, such
             as human babies with small tails or excessive
             hair, or horses with extra side toes (Fig. 6.6a),
             showing how earlier horses had fi ve, four or
             three toes, compared to the modern one.
               Vestigial structures tell similar phylogenetic
             stories. These are structures retained in living
             organisms that have no clear function, and
             may simply be there because they represent
             something that was once used. So, modern
             whales have, deep within their bodies, small
             bones in the hip region that are remnants
             of their hindlegs (Fig. 6.6b). Whales last
             had functioning hindlegs over 50 Ma in the
             Eocene, and the vestigial remnants are still
             there, even though they serve no further                     (a)
             purpose in locomotion, and only support
             some muscles associated with the penis.                                      femur pelvis
               The third aspect of development that forms
             links with phylogeny is the observation that
             ontogenetic patterns themselves have evolved.
             In particular the timing and rate of develop-
             mental events has varied between ancestors            (b)
             and descendants, often with profound effects.   Figure 6.6  Hints of ancestry in modern animals.
             This phenomenon is termed heterochrony.         (a) Extra toes in a horse, an example of an
                                                             atavistic abnormality in development, or a
                                                             throw-back, to earlier horses which had more
             Heterochrony: are human adults juvenile apes?
                                                             than one toe; normal horse leg (left), extra toes
             Heterochrony means “different time”, and        (right). (b) The vestigial hip girdle and hindlimb
             includes all aspects of changes of timing and   of a whale; the rudimentary limb is the rudiment
             rates of development. There are two forms       of a hindlimb that functioned 50 Ma.




             Table 6.1  The processes of heterochrony: differences in the relative timing and rates of development.
                                  Onset of growth   Sexual maturation   Rate of morphological development
              Pedomorphosis
               Progenesis                –                Early                        –
               Neoteny                   –                 –                        Reduced
               Postdisplacement       Delayed              –                           –
              Peramorphosis
               Hypermorphosis            –               Delayed                       –
               Acceleration              –                 –                        Increased
               Predisplacement         Early               –                           –
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