Evolutionary changes in the number of digits and other limb elements appear
to be severely constrained, probably as a result of a low level of modular
ity during limb development. Reduced limb structures typically develop thro
ugh a process of construction followed by destruction and amniotes have evo
lved many digit-like structures rather than actual extra digits. In amniote
s, limb development occurs during the crucial phylotypic stage, when many i
nductive interactions are occurring throughout the body. As a result, chang
es in limb development usually engender changes in other body parts. Thus,
mutations that change the number of limb bones are expected to have many pl
eiotropic effects, which severely reduces the chance of such mutations bein
g successful. In amphibians with aquatic larvae, limb development occurs af
ter the phylotypic stage and limb development is decoupled from the interac
tivity of the phylotypic stage. The constraint of pleiotropic effects is, t
herefore, expected to be weaker. This expectation agrees with the larger va
riability in the number of hand and foot structures in amphibians, with fro
gs even occasionally possessing six toes. These facts once again emphasize
the importance of pleiotropic effects as constraints to evolutionary change
, including their role in the conservation of body plans.