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The multiple skeletal components of the skull originate asynchronously and their developmentalschedule varies across amniotes. Here we present the embryonic ossificationsequence of 134 species, covering all major groups of mammals and their close relatives. Thiscomprehensive data set allows reconstruction of the heterochronic and modular evolution ofthe skull and the condition of the last common ancestor of mammals.We show that the modeof ossification (dermal or endochondral) unites bones into integrated evolutionary modules ofheterochronic changes and imposes evolutionary constraints on cranial heterochrony. However,some skull-roof bones, such as the supraoccipital, exhibit evolutionary degrees offreedom in these constraints. Ossification timing of the neurocranium was considerablyaccelerated during the origin of mammals. Furthermore, association between developmentaltiming of the supraoccipital and brain size was identified among amniotes. We argue thatcranial heterochrony in mammals has occurred in concert with encephalization but withina conserved modular organization

Mammalian skull heterochrony reveals modular evolution and a link between cranial development and brain size