In this collection of papers I have summarized my investigations into the field of evolutionary genetics and more specifically into patterns of biodiversity and evolutionary processes. The lizards (and frogs) studied here share common features in that they are largely present in unique environments, which are also regions that are biologically understudied. Most of these taxa show high degrees of endemism, interesting natural history characteristics, and each group manifests distinctive adaptations of general evolutionary interest. My work in the genus Telmatobius has been a progressive approach that began in my MS program, and it first focused on alpha taxonomy, morphological variation, and species boundaries. This work led to new studies initiated and completed at BYU involving further taxonomic revision (Formas et al., 2003; Chapter 1), and then revisiting and re-evaluating species boundaries established earlier (with allozyme markers) and this time with population level molecular (mitochondrial DNA) markers (Chapter 2). Our results indicate that the striking differences in size, coloration and general appearance in the various Lake Titicaca morphotypes are not genetically based. Further, there is evidence that these morphotypes have evolved very rapidly after demographic bottlenecks eroded present genetic variability. Telmatobius frogs of Lake Titicaca are listed by the International (IUCN) as critically endangered. We support this classification and further suggest studies to explore open questions like the possibility of adaptation along ecological resource gradients. Lizards of the genus Microlophus are interesting but for different reasons, and studies of this group constitutes the bulk of my dissertation work. The genus includes both Galapagos insular species, and continental taxa distributed in a linear gradient along > 4000 km of the western coast of South America. In studying Microlophus I first tackled the unresolved phylogenetic relationships within the genus (Chapter 3) and then pay attention to phylogeographic aspects of the most speciose lizard radiation in the Galapagos Archipelago (Chapter 4). Chapter 3 is a single manuscript provisionally accepted in the journal Systematic Biology. This paper introduces the lizard genus Microlophus (“lava lizards”) as a study system, and includes a large nuclear data set accompanied by an equally large mitochondrial data set (7877 characters in total). This paper explicitly differentiates among sequence alignments of gene regions that vary in tempo and class of mutational events. We show that this recognition is important and we suggest ways to appropriately deal with the alignment of multi-locus non-coding DNA data sets. A secondary finding in this study is that mtDNA and nDNA topologies are discordant with each other but that both are strongly supported, and that the nuclear topology is concordant with species distribution patterns along coastal South America. We hypothesize that in this particular region of the tree, the nuclear genome recovers a topology that is closer to the species tree, and conflicts occur due to likely secondary contact of distantly related taxa, suggesting that unique taxonomic relationships in the mtDNA gene tree are the result of hybridization. This last point highlights the value of dense taxonomic and character sampling for teasing apart different aspects of evolutionary processes. Chapter 4 is a manuscript to be submitted to the journal Evolution; in this study we further investigate the most speciose radiation of Microlophus in the Galapagos, based on an unparalleled sampling of most islands and small islets in the Archipelago. We use mtDNA sequences to both test hypothesized between-island colonization routes, as well as the expectation that within-island phylogeographic structure should be greater on older islands. Our mtDNA gene tree is strongly supported and allows rejection of previous alternatives, and we propose a novel sequence of between-island colonization events. Our results also reject the idea of phylogeographic structure been related solely to island age. Instead, we provide evidence to suggest that active volcanism as a major player in the generation of genetic diversity in within-island environments, and this is further compounded by the seemingly stochastic nature of within-island long-distance colonization routes mediated by ocean currents. We suggest that the direction and intensity of these currents, as currently understood, are insufficient to generate a priori hypotheses of oceanic colonization routes and their influence on gene flow. We do show that the standard stepping-stone model of migration, where genetic interchange is only possible among neighboring localities, does not explain much of the within-island population genetic structure unraveled by this study. From a biological conservation perspective the study of patterns of recent evolutionary history in the Galapagos provides with a window to evolutionary processes that have shaped and continue to impact the generation of biodiversity in the Galapagos Archipelago. Islands have long been viewed as natural laboratories of evolutionary change, and thus all island isolates are or could be distinctly important components of the larger, archipelago-wide processes. We provide working hypotheses for some of the demographic processes that might be generating within- and between-island biodiversity in this clade of lizards; confirmation of these explanations with independent data will have management implications for conserving the unique patterns observed in the Galapagos biota, but also the processes that generated these patterns.



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Life Sciences; Biology



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nuclear introns, alignment, length mutations, Microlophus, secondary contact, mitochondrial-nuclear conflict, phylogenetics, Galapagos, mtDNA, phylogeography, nested clade analysis, colonization routes, volcanism, lava lizards

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