Graduation Year


Document Type




Degree Granting Department

Biology (Integrative Biology)

Major Professor

Lynn B. Martin


Glucocorticoids, Invasive species, Passer domesticus, Range Expansion


Introduced species cause both considerable ecological and economic damage every year. However, not much is known about how certain species are able to establish and spread beyond the site of initial introduction, whereas others do not. Species undergoing range expansion following an introduction may prove to be a valuable resource to invasion biology, but may also be informative in light of species' responses to changing environments (i.e. global climate change). Here, I took advantage of an ongoing range expansion of an introduced vertebrate species. House sparrows (Passer domesticus) were introduced to Mombasa, Kenya in the 1950s and have subsequently expanded their range northwest-ward and now occupy most major cities in Kenya. By comparing older, established populations (i.e. those in Mombasa) with more recently colonized populations at the range edge, it might be possible to determine some of the mechanisms that underlie range expansion in some species and/or populations. In Chapter 1, the background and ideas that motivated the rest of the dissertation is summarized. In Chapter 2, I studied how exploration and glucocorticoids (a hormone released in response to stressors) changed throughout the range expansion. Exploration was greater at the range edge, which is likely to ensure greater discovery of novel resources. Glucocorticoids released in response to restraint were also highest at the range edge, which might facilitate resolution of stressors in unpredictable environments. However, chronically elevated levels of glucocorticoids are often considered maladaptive, unless an individual can appropriately cope with them. Therefore, in Chapter 3, I characterized glucocorticoid receptors (i.e. mineralocorticoid receptor (MR) and glucocorticoid receptor (GR)) in the hippocampus, an area responsible for negative feedback of glucocorticoids as well as induction of behavioral and physiological response to stressors. I found that MR density was lower relative to GR density at the range edge compared to the site of introduction (Mombasa). I speculate this pattern is a mechanism to resolve the elevated levels of glucocorticoids at the range edge. Taken together, these results indicate that individuals at the range edge have a strong glucocorticoid response to stressors to induce a rapid, strong response to resolve stressors. Subsequently, in Chapter 4, I examined the potential mechanisms of phenotypic change among Kenyan house sparrows. Typically, following an introduction event, genetic diversity undergoes a bottleneck and is greatly reduced compared to the source population; as such, genetic evolution as the main driver of changing phenotypes throughout the range expansion is unlikely. We therefore hypothesized that epigenetic mechanisms (e.g. DNA methylation) may compensate for the expected reduced genetic diversity following an introduction. Although there was no pattern of epigenetic variation among cities (i.e. variation did not increase nor decrease further from the site of introduction), epigenetic variation increased as genetic inbreeding increased (a sign of reduced genetic diversity and bottlenecks), suggesting epigenetic modifications may compensate for reduced genetic diversity following an introduction event. Overall, patterns of phenotypic variation emerged dependent on age of the population- these patterns may prove to be important in other vertebrate range expansions as well. Surprisingly, epigenetic diversity did not correlate with phenotypic variation among populations; however, within-individual studies may reveal epigenotypes are related to certain behavioral or physiological phenotypes. In the future, studies should be designed to address how phenotypic differences arise despite relatively low genetic diversity and overall high genetic admixture among individuals. In Kenyan house sparrows, maintenance of high levels of flexibility and differential developmental influences may be important factors that lead to varying phenotypes dependent on time since colonization.