Graduation Year


Document Type




Degree Granting Department

Biology (Integrative Biology)

Major Professor

Peter Stiling


enemies hypothesis, gallformers, interguild competition, Torymus umbilicatus, parasitoid-mediated associational resistance, stemborers


Indirect ecological effects such as associational resistance and resource competition have the potential to affect ecological interactions and influence the structure of ecological communities. Although resource competition is commonly studied, the effects of associational resistance are not as evident if studies are not designed to detect them. Additionally, the relative strengths of different ecological mechanisms ought to be measured in studies, rather than the strength of singular mechanisms. This permits proper attribution of causes and effects in community structure and detection of higher order interactions in a way that naïve reductionism will not.

In a series of experiments, I looked at the effects of large-scale addition and removal of Borrichia frutescens on associational resistance of Iva frutescens to the gallformer Asphondylia borrichiae in order to test the mechanism and strength of associational resistance in the system. Additionally, I measured the effects of relative host abundance and interpatch distance of hosts on associational resistance. Finally, I looked at the effect of the presence of stemborers competing with the gall former for host plant resources on parasitism rate and parasitoid guild composition. I found evidence for a strong effect of associational resistance by natural enemies over short distances, although the phenomenon is likely of minor importance in comparison to other factors known to affect gall former population density, such as environmental effects and host plant genotype. Competitors also had a weak effect, reducing mean gall diameter, but not significantly altering total parasitism rate. However, the presence of stemborer competitors did slightly alter the composition of the parasitoid guild.

In sum, both associational resistance and competition from stemborers have detectable effects on A. borrichiae, albeit weak ones. Environmental factors, such as soil nitrogen content, are likely much stronger determinants of gall former population size.