Morphologic evolution of New World marsupials (Mammalia, Didelphimorphia)

Author(s)

Harley Sebastião da Silva

• Document type: Doctoral Thesis
• Press: São Paulo.
• Institution: Universidade de São Paulo (USP). Instituto de Biociências (IBIOC/SB)
• Defense date: 2010-11-19

Examining board members

Gabriel Henrique Marroig Zambonato; Glauco Machado; Diego Astúa de Moraes; Paulo Inácio de Knegt López de Prado; Rui Cerqueira Silva

• Advisor: Gabriel Henrique Marroig Zambonato

Abstract

One of the central goals in evolutionary biology is related to how evolutionary processes, mainly natural selection and genetic drift, shaped living organisms. The combined use of Morphologial Integration and Quantitative Genetics give us powerful tools to accomplish this goal. Morphological Integration is concerned of how characters are related, as well as their underlying genetics/developmental relationship, while Quantitative Genetics have methodologies designed to explore the phenotypic forces underlying diversity among organisms. Using Didelphimorphia marsupials as a study group, I combined these two approaches to study its morphologic cranial diversification. In Quantitative Genetics, the genetic additive covariance matrix (G) resume the genetic variation underlying resemblance among relatives, which is the portion of the variance responsive to selection. Initially developed to microevolutionary scale studies, it can be extended to a macroevolutionary scale if it remains relatively similar in that time scale. However, as G matrix estimations require a huge number of related specimens with known genealogy, I used its phenotypic (P) counterpart which was more easily obtained. In the first chapter I showed high similarities among Didelphimorphia marsupials covariance and correlation P matrices. On the other hand, integration magnitudes (which measure the average correlation among traits) vary among taxa. Neither phylogeny nor morphologic distances showed any association with the similarity in patterns and magnitudes of integration. In the second chapter, I did these same analyses, but throughout genera ontogeny. Again, there was a high similarity among taxa in patterns of integration, both when I analyzed the ontogeny for each genus separately or against each other (at different age class). Morphological integration magnitudes showed the same variation obtained for adults, with a tendency to decrease at older ages. Taking these results into account, I compared the phenotypic correlation matrices to theoretical matrices, based on hypotheses of shared developmental and functional units. I searched for modularity in the two main skull regions (face and neurocranium) and five sub regions (cranium base and vault, face, nasal, and oral). I also looked for modularity concerning somatic development (Neurocranium vs. Face) and total modularity, as the 200 summation of the five sub-regions. Only Face and its sub-regions nasal and oral, showed significant correlations to the phenotypic genera matrices. Despite integration magnitude differences, all evolutionary responses produced by taxa were highly similar. These results, combined with a huge size variation (or size related variation - allometry) across taxa, lead me to search for the evolutionary consequences due to size variation. In the third chapter, I compared evolutionary response directions produce by each genera matrices before and after size removal under a random selection simulation. Allometry strongly affect these skulls, turning them into highly integrated structures with lower modularity (skull modules are not easily distinguished). Because of this, modules cannot evolve relatively independent of other modules and evolutionary responses will strongly affect the whole cranium. This is related to the variation along the lines of least evolutionary resistance. This line is the multivariate direction of greatest genetic or phenotypic variation (the combination of a suite of traits that displays the maximum within-population variance). In Didelphimorphia marsupials, this line is aligned with size variation and regardless the selection direction, evolutionary change is usually aligned to this least resistance line. The removal of size variation diminish the magnitude of integration while increases modularity. Consequently, skulls become able to respond to selection in more directions as modules become relatively more independent of each other. In the last chapter I compared size and shape differences between ontogenetic trajectories of two sister genera Didelphis and Philander. Using traditional and geometric morphometric analysis plus allometric coefficient analysis, I could show that bigger differences between them are size related. Despite similarities, ontogenetic trajectory in Didelphis is longer, leading to bigger specimens. (AU)