Developmental structure in brain evolution

First, we clarify the central nature of our argument: our attempt is to apportion variation in brain size between developmental constraint, system-specific change, and "mosaic" change, underlining the unexpectedly large role of developmental constraint, but making no case for exclusivity. We consider the special cases of unusual hypertrophy of single structures in single species, regressive nervous systems, and the unusually variable cerebellum raised by the commentators. We defend the description of the cortex (or any developmentally-constrained structure) as a potential spandrel, and weigh the implications of the spandrel concept for the course of human evolution. The empirical and statistical objections raised in the commentary of Barton are discussed at length. Finally, we catalogue and comment on the suggestions of new ways to study brain evolution, and new aspects of brain evolution to study.