Ecological interactions within a community shape the structure of ecosystems and influence ecosystem function. Plant-pollinator interactions exist as mutualistic exchange networks that may collapse as habitat loss occurs, thereby threatening the overall health of an ecosystem. Understanding the impacts of human-mediated habitat disturbance on ecological interactions is therefore crucial for conservation efforts. Archbold Biological Station (ABS) in Venus, Florida contains over 2000 hectares of protected Florida scrub habitat nested within a human-dominated environment that is threatened by anthropogenic habitat disturbance. In past studies, over 113 bee species and 157 associated host plants, many endemic to the Lake Wales Ridge, have been found on ABS property, providing an understanding of this system's plant-bee network. Using those data as a baseline, this study investigated the effects of varying levels of mechanical habitat disturbance intensity on the diversity and network structure of plant-bee interaction networks. Flowering plant abundance, richness, diversity, and composition as well as bee abundance and composition were significantly different across mechanical habitat disturbance levels. Interactions between bees and flowering plants also differed with varying disturbance intensity. From these results, it is clear that plants, bees and interactions between them are impacted by mechanical habitat disturbance in this system. This project informs management efforts not only for natural systems with the threat of alteration, but also for agricultural systems, many of which heavily rely on flower visitation by bee pollinators. This research also contributes to the growing field of interaction ecology by increasing understanding of habitat alteration effects on a valuable ecological interaction and ultimately ecosystem function.

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Mutualism is an understudied interaction in ecosystems throughout the world. Within the eastern United States, one fish-fish mutualism is the nest association between Nocomis and other cyprinids. I assessed the role of host parental care while testing for the selfish-herd effect. Additionally, I examined multiple nest associates in order to elucidate potential cheaters. I utilized gonadosomatic index (GSI) to compare reproductive condition among the bluehead chub Nocomis leptocephalus and its putative nest associates in Catawba Creek, Virginia. GSI of potentially obligate associates tracked host GSI more closely than weak associates, while weak associates spawned prior to Nocomis spawning. Given their GSI patterns and behavior, central stonerollers Campostoma anomalum may be cheaters in the interaction. I used multiple experiments to test for the selfish-herd effect, the role of parental care, and how relative risk influences reproductive decisions of associates. Most eggs were located in the bottom upstream quarter of nests, and a molecular analysis revealed that stonerollers and chubs constituted the majority of identified individuals. A comparison of host-associate ratios from four nest sections failed to identify the selfish herd effect. Another experiment found that host egg covering significantly reduced egg predation. Lastly, I assessed relative egg predation risk at four potential spawning locations; predation levels did not differ significantly at any location. While GSI patterns suggest that stonerollers may be cheaters, genetic evidence indicates that stonerollers spawn on Nocomis nests. Because GSI does not completely assess reproduction, secondary stoneroller reproduction on Nocomis nests may have been overlooked. / Master of Science

Many species have evolved to live in groups. Gregarious behavior was believed to be adaptive for whole-population survival and predator evasion until selfish-herd theory was introduced. It proposed that individuals congregate not as a method of protection for the entire population, but instead to better the individual's chance of survival, thereby using the group to benefit itself (i.e., "selfishness"). Selfish behavior is a common part of mutualisms, which are complex, dynamic interactions that often change with biotic or abiotic circumstance. Here, I investigate potential selfish behavior within the mixed-species spawning aggregations hosted by bluehead chub (Nocomis leptocephalus). I hypothesized that the host benefits more directly from the aggregation it supports than previous research suggests – specifically, by using nest associates to decrease its own risk of predation (creating a selfish-herd) and simultaneously forcing associate embryos into marginal nest locations while locating its own embryo in the safer locations within the nest (creating an "embryonic selfish-herd"). In Chapter 1, I investigated the adult spawning aggregation for possible selfishness by monitoring the chub's spatial location within the aggregation and vigilance behavior under varying associate abundances. I found that the host occupied the central location within the aggregation and was less vigilant when associate abundance was high. In Chapter 2, I examined the "embryonic herd" contained within Nocomis nests for possible embryonic selfishness orchestrated by the chub host, leading to increased embryonic survival for chub young and elevated fitness for chub parents. I found that deeper nest sections support higher embryonic survival than shallower sections, and that chub embryo make up a disproportionate percentage of embryos found in those deeper sections. I also conducted a preliminary study investigating embryo-predation by host on associate embryo which produced evidence for embryo-predation by both host and two associates on each other's offspring. Cohesively, my results support the identification of the host as a selfish participant and confirm the presence of both an adult and embryonic selfish-herd in this mutualism. This is the first study to demonstrate selfish behavior on the part of the host in this system and the first to prove the existence of an embryonic selfish-herd. / Master of Science / A fish has three goals: to survive, to grow, and to reproduce. Mutualisms – when all participants experience a net benefit from interacting – arise when two or more species rely on each other to meet these objectives. In many mutualisms, however, not all participants benefit equally. Selfish-herd theory identifies those that maximize their own benefit while minimizing or negating any cost as "selfish" individuals. I examined the mutualistic mixed-species spawning aggregations hosted by a freshwater minnow, bluehead chub (Nocomis leptocephalus), for possible selfish behavior. I found that bluehead chubs disproportionately benefit from the aggregations they host and use the aggregation to improve their own survival, improve their offspring's survival, and obtain food (by eating others' offspring). Thus, I identify bluehead chub as a selfish participant and provide evidence for the existence of two selfish herds in this mutualism – one among the adult fishes and one among their young – both orchestrated by and benefiting the bluehead chub. This study is the first to identify the bluehead chub as such and rewrites our previous understanding of this mutualism, which largely denied bluehead chub as a direct beneficiary. This work contributes to the global discussion of mutualisms by attesting the complexity of these relationships and offers support for re-examination of the classification of many known interspecific interactions, such as those cursorily termed nest parasitisms.

Crayfish throughout North America, Europe, and Asia host assemblages of obligate ectosymbiotic annelid worms called branchiobdellidans. The work presented here is a detailed experimental and observational study of the ecological interactions between crayfish and their worms. In a comprehensive literature review, I show that branchiobdellidans have complex and context-dependent effects on their hosts, serving as both beneficial cleaners and tissue-consuming parasites. Using a field survey and laboratory experiments, I provide novel evidence for age-specific resistance as an adaptation to maximize life-long benefits of a mutualism. Specifically, I show that Cambarus crayfish display a consistent ontogenetic shift in resistance to the colonization of branchiobdellidans and this shift likely reflects underlying changes in the costs and benefits of symbiosis. I then show that this change in host resistance creates predictable patterns of symbiont diversity and composition throughout host ontogeny. Host resistance limits within-host symbiont communities to a few weakly interacting species, whereas relaxed resistance leads to more diverse symbiont communities that have strong interactions among symbiont taxa. Thus, host resistance has direct effects on within-host symbiont community structure by selectively filtering colonizing species, and indirect effects by moderating the strength of interactions among symbionts. Lastly, in a detailed study of the worm Cambarincola ingens, I depict a symbiont dispersal strategy that yields highly predictable transmission dynamics during pairwise host-host encounters and shows that variation in transmission dynamics can be explained by the fitness outcomes for dispersing symbionts. Field observations revealed that worm reproduction is contingent on host size and intraspecific competition for preferred microhabitats. Using a predictive model that assumes transmission of symbionts only when current conditions yield fitness below a minimum threshold, I was able to predict individual transmission events much more accurately than a comparable null model that assumed a fixed probability of transmission. My work provides empirical support for the emerging trend among researchers that advocates the adaptation of general ecological frameworks to understand symbiont population structure and diversity, but my work also emphasizes the value of detailed natural history studies to uncover system-specific ecological and co-evolutionary processes such as partner control mechanisms, symbiont microhabitat selections, and symbiont dispersal strategies. / Ph. D.

This thesis provides a contextualised exegesis and re-evaluation of the anarchist Pierre- Joseph Proudhon's writings on war and peace. The thesis has two claims to originality. The first lies in shedding new light on Proudhon's voluminous writings on international politics. These texts have been relatively marginalised in the broader secondary literature on Proudhon's thinking, and the thesis seeks to correct this important lacuna. In International Relations (IR), the academic discipline to which this thesis will make its most obvious original contribution, Proudhon's writings on war and peace have been almost completely ignored. By providing an anarchist approach to world politics, the thesis will also contribute to IR's historiographical and critical theoretical literature. The second claim to originality lies in using these writings and the context from which they emerged to tell a story about the evolution of the nineteenth century, the origins of the twentieth century and provide possible ways of thinking beyond the twenty first. The thesis employs a contextualist methodology that works in four ways. First, I have contextualised Proudhon's thought geo-politically, in relation to the dynamics of the balance of power in nineteenth-century Europe. Secondly, I have sought to understand Proudhon's ideas against the backdrop of the evolution of the French nation state in the mid to late nineteenth century. Third, I have shown how Proudhon's thought emerges out of the dominant intellectual currents of his day – ideas that range from the inspiration for the activism of Fourierist and Saint-Simonian feminists, to the epochal influence of Rousseau and Kant. Finally, I argue that Proudhon's thinking on world politics needs to be understood in relation to the evolution of his own thinking after Napoleon III's coup d'état of the 2nd of December 1851. I will show that Proudhon's mature anarchism, his mutualist federalism, was an engaged response to each of these social and intellectual contexts. I will argue that his critiques of these processes, and their intellectual champions, have been given an added poignancy given that he campaigned in large part against those very processes that culminated in two world wars.

Seed dispersal by caching rodents is a context-dependent mutualism in many systems. Plants benefit when seed remaining in shallow caches germinates before being eaten, often gaining protection from beetles and a favorable microsite in the process. Caching in highly unfavorable microsites, conversely, could undermine the dispersal benefit for the plant. Plant invasions could disrupt dispersal benefits of seed caching by attracting rodents to the protection of a dense invasive canopy which inhibits the establishment of native seedlings beneath it. To determine whether rodents disproportionately cache seed under the dense canopy of an invasive grass in southeastern Arizona, we used nontoxic fluorescent powder and ultraviolet light to locate caches of seed offered to rodents in the field. We fitted a general habitat-use model, which showed that disproportionate use of plant cover by caching rodents (principally Chaetodipus spp.) increased with moonlight. Across all moon phases, when rodents cached under plants, they cached under the invasive grass disproportionately to its relative cover. A greenhouse experiment showed that proximity to the invasive grass reduced the growth and survival of seedlings of a common native tree (Parkinsonia microphylla) whose seeds are dispersed by caching rodents. Biased dispersal of native seed to the base of an invasive grass could magnify the competitive effect of this grass on native plants, further reducing their recruitment and magnifying the effect of the invasion.

Insects are often infected with facultative endosymbiotic bacteria, which can have a range of important ecological effects. The grain aphid, Sitobion avenae, harbours diverse facultative symbionts, which suggests their importance in grain aphid biology. This thesis attempts to explain the ecological roles of the facultative endosymbionts in S. avenae. It also examines the question of whether the horizontal transmission of symbionts between aphid clones and species can be important for shaping the ecology and evolution of multi-species aphid communities. Novel techniques developed for research with the grain aphid study system are presented. Grain aphid clones vary in their tolerance to low temperatures, but this trait is not affected by their facultative endosymbionts. Strains of a symbiont Hamiltonella defensa do not protect grain aphids from hymenopterous parasitoids, regardless of the host genotype. However, experienced parasitoid females preferentially oviposit in aphids which do not harbour symbionts. Comparison of the fitness consequences of infection with the same Hamiltonella strains in their original and in novel grain aphid host clones reveal no consistent differences. Symbiont strains establish easily following artificial transfer between clones of the grain aphid, but the symbionts transferred from other aphid species form less stable infections. Hamiltonella strains do not affect the fecundity of their grain aphid host clones regardless of their host species of origin, but also do not generally confer protection against parasitoids. There are no clear patterns in the distribution of parasitoid-resistant phenotypes across phylogenetic trees of Hamiltonella and its bacteriophage APSE. Strains of four unrelated species of endosymbionts, Rickettsia, Spiroplasma, Rickettsiella and Regiella, confer the same pathogen-resistant phenotype to a single pea aphid clone. The same symbiont strains can confer resistance to clones of two different aphid species. Some strains in multiple infections may compensate for the costs of infections with other symbionts. The importance of these results for understanding the ecological and evolutionary role of facultative endosymbionts in aphids and other insects are discussed, and directions for further research are proposed.

Whether species interactions are static or change over time has wide-reaching ecological and evolutionary consequences. However, species interaction networks are typically constructed from temporally aggregated interaction data, thereby implicitly assuming that interactions are fixed. This approach has advanced our understanding of communities, but it obscures the timescale at which interactions form (or dissolve) and the drivers and consequences of such dynamics. We address this knowledge gap by quantifying the within-season turnover of plant-pollinator interactions from weekly censuses across 3years in a subalpine ecosystem. Week-to-week turnover of interactions (1) was high, (2) followed a consistent seasonal progression in all years of study and (3) was dominated by interaction rewiring (the reassembly of interactions among species). Simulation models revealed that species' phenologies and relative abundances constrained both total interaction turnover and rewiring. Our findings reveal the diversity of species interactions that may be missed when the temporal dynamics of networks are ignored.

This thesis explores the role mutualist pollinators and their symbionts play in the genetic structuring and speciation of their host plants along an elevational gradient in Papua New Guinea. Using the fig and fig-wasp mutualism as a model system, we employed high-throughput sequencing techniques to explore fine-scale population genomics of both fig and wasps along their elevational range. We found there to be clear lowland and highland clustering of tree populations along the gradient, often with a mid-elevation contact zone. In the case of the pollinating wasps, we retrieved the same clustering except in this case, the genetic difference between clusters was high enough as to consider them as separate species. This result supports evidence from other studies challenging the cospeciation paradigm of one wasp species per fig species. In addition, we explore ecological traits which may promote, or at least, maintain, reproductive isolation between fig (sub)species along with behavioural preference tests from pollinating wasps. In order to further investigate the mechanisms promoting wasp speciation along the gradient, we describe Wolbachia infection status as well as strain type. Wolbachia-induced cytoplasmic incompatibility (CI) is often invoked as a possible speciation agent since it can rapidly provoke and maintain reproductive isolation between otherwise freely interbreeding insect populations. Finally, we explore non-pollinating fig wasp (NPFW) diversity along the gradient for a subset of our focal species. Our study reveals that there is a tight relationship between NPFW diversity and host species, and a mid-elevation peak.