Biodiversity in a Dynamic World: How Environmental Variability Influences Coexistence between Introduced and Native Species

Abstract

Understanding broad patterns of biodiversity requires developing a unified and rigorously tested theory that explains how species coexist despite the risk of competitive exclusion. Species interactions are fundamentally shaped by environmental variability. Recent theoretical development has predicted a set of general mechanisms that promote species coexistence under variable environments. Nevertheless, this theoretical framework has received limited empirical tests. Biological invasions offer excellent opportunities to empirically test coexistence mechanisms in communities in which the stability of coexistence is likely affected by introduced species. I took this opportunity to directly test the theory of species coexistence in this dissertation work by investigating how environmental variability affected the invasion of introduced species and their coexistence with native species. My collaborators and I started the investigation of diversity maintenance by first examining the range expansion of introduced species. Studying range shifts can reveal drivers of diversity patterns, which are formed by overlapping ranges of different species. We used a novel spatial analysis to determine the scale-dependent expansion rate of an invasive winter annual species, Brassica tournefortii over North America as well as to infer the drivers of this scale dependency. We found that this species expanded rapidly on scales from 5 to 500 km historically but had ceased its current expansion on the 100-500 km scales due to climatic constraints. This finding left open the question why this species continued its spread on the 5-50 km scales and how it would impact native species within its invaded range. To address these questions, we examined relatively local scale interactions between B. tournefortii and its competitors. We compared key demographic rates of B. tournefortii with other invasive and native winter annuals over a Sonoran Desert landscape to check conditions necessary for their spatial and temporal niche differentiation. We found the presence of two essential requirements for their niche differentiation: species-specific germination responses that could differentiate species by their favored environments and buffered population growth in time and space that could prevent catastrophic population declines when species faced unfavorable conditions. These conditions could provide niche opportunities to promote both the establishment of B. tournefortii and the persistence of native species under its presence. Building upon this finding, we directly quantified one general mechanism of spatial niche differentiation between B. tournefortii and its native competitors. We measured the strength of this mechanism, the spatial storage effect, across a hierarchy of spatial scales (subhabitat -> habitat -> landscape). We found that this mechanism did not promote species coexistence on any of these scales over the study period. These species were not differentiated over their tested spatial niches because weak competition following dry growing conditions failed to intensify intraspecific competition relative to interspecific competition. The strength of this mechanism increased from occasionally producing negative effects on lower scales to consistently being non-negative on the highest scale. This scale-dependent pattern was in line with the expectation that coexistence potential would increase with scales as species interacted over a wider range of environmental heterogeneity. Our findings demonstrated empirically that environmental variability in time and space led to scale-dependent patterns of the coexistence potential between introduced and native species. This work showed that introduced and native species could be differentiated by their environmental responses given spatial and temporal environmental heterogeneity on higher scales. However, for those species to stably coexist on higher scales, competitive effects had to follow environmental responses to separate species by their own density-dependent feedback loops. This work is among the first few empirical tests of a body of theory that holds the promise to generalize the mechanisms of spatial and temporal niche differentiation. Its success and limitation can motivate more studies to adopt the guiding mathematical principles and to use similar yet more innovative approaches to address the grand question of biodiversity maintenance.