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The results of evolution can be inferred from comparative studies of related organisms. In this doctoral thesis, I use phylogenetic comparative methods along side geometric morphometrics to analyse shape variation in order to infer evolution of the caecilian skull. Caecilians are elongate, limbless amphibians that superficially resemble snakes or earthworms, and use their head as a locomotory organ. I characterise large-scale patterns of cranial morphological diversity and quantify variation across the main family-level clades by describing patterns relating to phylogeny, disparity and ecology. Then I examine the origins and evolution of morphological variation in the skull by describing patterns relating to morphological integration and modularity. This thesis demonstrates a variety of existing statistical techniques that can be used to infer processes from large-scale evolutionary patterns in morphological data using non model organisms. Throughout the thesis, I show that the evolution of the caecilian skull to be multifaceted. On the patterns of diversity, the most striking is a "starburst" arrangement in shape space, which suggests that early in caecilian evolution ancestral lineages traversed greater expanses of the shape space, while subsequent phylogenetic divergence within the main clades entailed less morphological diversification. This may be related to early diversification into different ecological-niches, yet more data are needed to test this. The clades differ considerably in their cranial disparity, but there appears to be no unified pattern across the whole order that indicates disparity is coupled with clade age or speciation events. I show that aquatic species are more diverse than their terrestrial relatives, and that there is convergence of cranial shape among dedicated burrowers with eyes covered by bone. On the patterns of morphological integration and modularity, another remarkable finding is the caecilian cranium is modular with respect to two functional regions, the snout and the back of the cranium. Modularity is important for understanding the evolution of this structure. The main elements of the caecilian anterior skeleton, the cranium, mandible and atlas vertebra, reveal different patterns of morphological integration, suggesting different developmental and evolutionary processes are involved in sorting and maintaining new variation of each structure. Allometry is an important component of integration in each of the structures. Covariation of the cranium-mandible after size correction is significant and follows the same direction of shape change across all levels and as shown for allometry. In contrast, covariation of the cranium-atlas follows different directions at each level. These results suggest the two main joint of the caecilian skull differ substantially in their origin and evolution. I discuss the contribution made in this thesis to caecilian and evolutionary biology and offer an outlook of how theses findings can be used to initiate future studies to better understand of the evolution of the caecilian skull. |
