The crocodylian skull and osteoderms: A functional exaptation to ectothermy?

The crocodylians are ectothermic semi-aquatic vertebrates which are assessed to have evolved from endothermic terrestrial forms during the Mesozoic. Such a physiological transition should have involved modifications in their cardio-vascular system allowing to increase the heat transfers with the surrounding environment by growing a peripheral vascularization which would be mainly located in the dermal skeleton: the dermatocranium and the osteoderms. In order to assess the implication of these anatomical regions in thermal exchanges, we have recorded the temperature above a set of representative skin areas in order to draw comparisons between the skull, the osteoderms, and the rest of the body parts which present either none or residual dermal ossification. We computed the data after the specimens were successively laid in different stereotyped environmental conditions which involved significant variations in the environmental temperature. Our results show that the osteoderms collect the external heat during the basking periods as they become significantly warmer than the surrounding skin; they further release the heat into the core of the organism as they turn out to be colder than the surrounding skin after a significant cooling period. In disregard of the environmental temperature variations, the skull table (which encloses the braincase) remains warmer than the rest of the cranial regions and shows less temperature variations than the osteoderms; a result which has lead us to think that the braincase temperature is monitored and controlled by a thermoregulatory system. Therefore, as hypothesized by previous authors regarding the ectothermic diapsids, we assume that the crocodylian skull possesses shunting blood pathways which tend to maintain both the braincase and the main sensory organs at the nearest to the optimal physiological temperature depending on the external temperature variations. Concerning the skin vascularization, the study of an albino Alligator mississippiensis specimen permitted to observe the repartition of the superficial blood vessels by transparency through the skin. We thus testify that the skin which covers either the skull or the osteoderms is more vascularized than the skin which does not present any subjacent dermal ossification. We consequently deduce that the significant contrast in the thermal behavior between the dermal skeleton and the rest of the body is indeed correlated with a difference in the relative degree of skin vascularization. This last assessment confirms that the development of the dermal skeleton should have played a functional role in the crocodylian transition from endothermy to ectothermy through the set-up of a peripheral vessel network.