Testing the hypothesis of morphological integration on a skull of a vertebrate with a biphasic life cycle: a case study of the alpine newt.

In this article, we explore the possible influences of the developmental and functional relationships between skeletal elements on the pattern of morphological integration in the adult skull of the alpine newt. Like many tailed amphibians, the alpine newt has a biphasic life cycle, which implies the possibility that two distinct sets of constraints on development and function of the cranial skeleton may act at different times. We study how trait covariation, resulting from processes early in development, affects patterns of covariation at the adult stage. We test whether the observed patterns of integration are consistent with those predicted from three a priori hypothesized sources of integration: developmental timing, hormonally mediated growth/remodeling during metamorphosis, and developmental and functional relationships. The analyses of the covariation among the landmarks in the dorsal and ventral alpine newt craniums yield somewhat contrasting results. Our results do not indicate a clear correspondence between the observed variations in the skull shape and any of the three proposed hypotheses. No traceable reflection of hypothesized developmental relationships in the pattern of morphological integration/modularity in the adult skull indicate that covariation structure is continually restructured by overlaying variation introduced through developmental and environmental factors at different stages of development. This finding supports the recently elaborated palimpsest view of morphological integration. Also, our results indicate that the allometry-free shape data have an even higher level of morphological integration than the data that contain the allometric component of the shape variation.