Understanding what drives variation in species diversity in space and time and limits coexistence in local communities is a main focus of community ecology and biogeography. My doctoral work aims to document patterns of ant diversity and explore the possible ecological mechanisms leading to these patterns. Elucidating the processes by which communities assemble and species coexist might help explain spatial variation in species diversity. Using a combination of manipulative experiments, broad-scale surveys, behavioral assays and phylogenetic analyses, I examine which ecological processes account for the number of species coexisting in ant communities. Ants are found in most terrestrial habitats, where they are abundant, diverse and easy to sample (Agosti et al. 2000). Hölldobler and Wilson (1990) noted that competition was the hallmark of ant ecology, and we know that ant diversity varies along environmental gradients (Kusnezov 1957). Thus ants are an ideal taxon to examine the factors shaping the structure of ecological communities and how the determinants of community structure vary in space.

Thelohania solenopsae are intracellular pathogens found in the red imported fire ant, Solenopsis invicta. These pathogens cause detrimental effects to their fire ant hosts. The present study revealed that the midgut and the meconium materials from pupating fourth instar larvae were possible vehicles for the horizontal transmission of the disease. The pathogen was further found to cause a reduction of humeral proteins. In SDS-PAGE stained with silver, several proteins were observed only in controls but not in infected fire ant queens. Different queens were found to have variable proteins reduced due to infection of this pathogen. Furthermore, vitellogenin titers were found to be significantly reduced in infected fire ant queens, although the infection rates of the fat body cells was found to be less than 20%. Finally, although the pathogens did not directly induce apoptosis in infected cells, there were more infected cells undergoing apoptosis than uninfected cells. There was no evidence to support the idea that infected fat body cells became more resistant to apoptosis inducers.

The aim of this thesis is to study phylogeny, species-wide phylogeography and genetic diversity in Formica ants across Eurasia in connection with the history of biotic responses to Quaternary environmental changes. The mitochondrial DNA phylogeny of Palaearctic Formica species supported the subgeneric grouping based on morphological similarity. The exception was that F. uralensis formed a separate phylogenetic group. The mitochondrial DNA phylogeny of the F. rufa group showed the division into three major phylogenetic groups: one with the species F. polyctena and F. rufa, one with F. aquilonia, F. lugubris and F. paralugubris, and the third one with F. pratensis. West-east phylogeographic divisions were found in F. pratensis suggesting post-glacial colonization of western Europe and a wide area from Sweden to the Baikal Lake from separate forest refugia. In contrast, no phylogeographic divisions were detected in either F. lugubris or F. exsecta. Contraction of the distribution range to a single refugial area during the late Pleistocene and the following population expansion could offer a general explanation for the lack of phylogeographic structure across most of Eurasia in these species. Sympatrically distributed and ecologically similar species F. uralensis and F. candida showed clear difference in the phylogeographic structure that reflected difference in their vicariant history. Whereas no phylogeographic divisions were detected in F. uralensis across Europe, F. candida showed a well-supported phylogeographic division between the western, the central and the southern group. In socially polymorphic F. cinerea, the overall level of intrapopulation microsatellite diversity was relatively high and differentiation among populations was low, indicating recent historical connections. The lack of correspondence between genetic affinities and geographic locations of studied populations did not provide any evidence for differentiating between alternative hypotheses concerning the directions and sources of postglacial colonization of Fennoscandia.

Social insects are used as models for understanding the evolution of sociality because they show seemingly complex behavioral and physiological traits that enforce group cohesion, collective organization, and group level reproduction. Social organization in insect societies requires workers to share information. Information sharing allows workers to efficiently perform and switch among tasks to meet colony needs. For many species that nest in preformed cavities, colony growth results in crowding inside the nest which can affect colony productivity and fitness. How does colony size and social density affect individual and collective behavior? Using a combination of laboratory and field experiments, I have begun to answer this question. In Temnothorax rugatulus ants, high social density resulted in greater colony energy use. In addition, larger colonies used proportionally less energy compared to smaller colonies, but showed reduced brood production. These results indicate that the way colonies use energy changes with social density and group size. In analyzing the effects of colony size and density on worker behavior, I found that high density increased worker connectivity and information sharing. Workers in larger colonies showed less connectivity compared to workers in smaller colonies. Interestingly, workers with more interactions spent less time in brood care. This study shows that workers' access to information and the overall pattern of information flow are affected by social density and colony size, and changes in worker connectivity can influence task behavior. The next study shows that field colonies maintained a relatively constant level of intranidal density irrespective of colony size; this suggests that Temnothorax ants actively regulate social density. When colonies were established in high density nests, they showed greater foraging and scouting activities, and this led to a higher probability for becoming polydomous, i.e., occupying multiple nests. When polydomy occurred, colonies divided evenly between two nests, but distributed fewer, heavier workers and brood to the supplemental nests. Taken together, the first four studies indicate that social density is an important colony phenotype that affects individual and collective behavior and energetics in ants, and the collective management of social density may be a group adaptation in ants and other social insects. Lastly, because crowding affects polydomy behavior, the final two experiments tested whether colony emigration and nest construction and dispersion, two strategies for reducing intranidal crowding, are influenced by food distribution. Temnothorax colonies preferred to emigrate to nests positioned closer to food, and weaver ants (Oecophylla smaragdina) positioned newly constructed nests in food-rich areas. Furthermore, weaver ants used the newly constructed nests to more rapidly retrieve and safeguard valuable food items. Thus, strategic emigrations and adaptive nest dispersion can remedy intranidal crowding and at the same time allow growing colonies to acquire adequate food to meet colony needs.

Granular materials are abundant in the natural and industrial environment. Typical granular materials are collections of inert, passive particles in which the constituent grains of the material are macroscopic; thus they fill space, are athermal, and interact through only local contact forces. This definition can be broadened to include non-inert particles as well-active particles-in which the grains of an active granular material possess an internal energy source which drives motion. Active granular materials are found in many areas of the biological world, from cattle stampedes and pedestrian traffic flow, to the subterranean world of ant colonies and their collective motion within the nest. We study the rheology and dynamics of inert granular material, and an active granular system of collections of fire-ants, which together we call matter with granularity. In both of these systems we observe bifurcations in the force and flow dynamics which results from confinement effects of the effectively rigid granular materials. In inert granular systems, the onset of flow among particles that are closely packed together causes them to dilate as particles must separate away from each other to accommodate flow. Dilation is a property unique to matter with granularity and other complex fluids in which particles interact locally and occupy space. We explore how dilation influences the inert granular system in situations of local and global forcing: drag of an immersed intruder and avalanche flow respectively. We next study collections of fire ants which also interact with each other locally through contact forces and exclude volume. We study the construction of, and locomotion within subterranean tunnels by groups of fire ants. We find that the traffic dynamics of ants within confined tunnels are significantly affected by tunnel diameter. Reducing tunnel diameter increases the formation of traffic jams due to the inability of ants to pass each other easily. However, we show that jamming within tunnels may have beneficial effects on subterranean locomotion. Individual ants jam there bodies against the walls of vertical tunnels to resist falling. From physics studies of fire ant mobility in confined spaces, we show that subterranean tunnel size has a significant effect on the stability and mobility of ants within these environments.

Higher densities of S. invicta in the United States relative to South America are explained mainly by the absence of natural enemies and low interspecific competition (IC). Despite advances in S. invicta management, broadcast insecticide baits remain as the primary tool for effective control. I studied interspecific interactions of ants and the use of baits on the management of S. invicta to test the following hypotheses: 1) relative abundance of native ants increases ~25% for bait treated sites compared to untreated, 2) behavioral dominance by S. invicta decreases ~10% in bait treated sites compared to untreated, and, 3) foraging by S. invicta on insecticide baits is higher ~10% in low native ant densities areas compared to high densities. Experiments were conducted on three sites with different densities of native ants (low, medium, high), but with similar densities of S. invicta. An enhanced BACI (Before/After-Control/Impact) design was used. Experimental units consisted of 0.4 ha plots. Three treatments were randomly assigned to units and replicated four times; 1) Slow acting bait, 2) Fast acting bait, 3) Untreated Control. Samples and observations were collected for several weeks before/after the treatments to account for temporal variation and to determine rates of reinvasion. Ants were monitored using pitfalls and food lures. Interspecific competition was determined by applying placebo bait. Results indicate that different management methods did not impact resident ants when they are at low and medium densities and that S. invicta is greatly affected; at high native ant density, competition for these baits is observed affecting both natives and S. invicta. Reinvasion of S. invicta was reduced in areas of low and medium native ant densities previously treated with baits and native ant abundance increased by ~25%. At high native ant densities, reinvasion of S. invicta was similar to Control sites. I conclude: 1) properly used, baits can help in recovery of native ant species that then compete with S. invicta; however, rote re-treatments may have a negative impact on restored populations. 2) “bait failures” may be due to interspecific competition when initial native ant densities are high. Preliminary management considerations and recommendations are presented.

Thelohania solenopsae are intracellular pathogens found in the red imported fire ant, Solenopsis invicta. These pathogens cause detrimental effects to their fire ant hosts. The present study revealed that the midgut and the meconium materials from pupating fourth instar larvae were possible vehicles for the horizontal transmission of the disease. The pathogen was further found to cause a reduction of humeral proteins. In SDS-PAGE stained with silver, several proteins were observed only in controls but not in infected fire ant queens. Different queens were found to have variable proteins reduced due to infection of this pathogen. Furthermore, vitellogenin titers were found to be significantly reduced in infected fire ant queens, although the infection rates of the fat body cells was found to be less than 20%. Finally, although the pathogens did not directly induce apoptosis in infected cells, there were more infected cells undergoing apoptosis than uninfected cells. There was no evidence to support the idea that infected fat body cells became more resistant to apoptosis inducers.

I examined the impacts of the invasive red imported fire ant (RIFA, Solenopsis invicta) on native ants (Monomorium minimum, Paratrechina sp., S. krockowi, Pheidole metallescens, Forelius pruinosus, and Camponotus americanus) and litter removal in a post oak savannah community in central Texas. The study site was divided into 3 adjacent areas, and ant-toxic bait was used, along with additional colonies of RIFA, to establish 3 different densities of RIFA (naturally occurring, low, and high). I surveyed the ants in the 3 density areas and calculated the catch per unit effort for each species. Litter baits were placed in the 3 density areas for 14 12-hour trials. The masses of the litter removed were measured, and means were calculated for each speciesdensity/ trial/date/period/bait combination. The average amounts of litter removed by RIFA and native ant were different in the 3 density areas (0.42 g, 0.0 g, and 0.75 g for RIFA in the natural RIFA density area, low RIFA density area, and high RIFA density area, respectively; 0.0 g, 0.16 g, and 0.15 g for native ants in the natural RIFA density area, low RIFA density area, and high RIFA density area, respectively), indicating that RIFA does have an effect on native ant habitat use.

Biodiversity is distributed unevenly across geographic space and the tree of life. A key task of biology is to understand the ecological and evolutionary processes that generate these patterns. I investigate how the structure and geometry of a landscape, for example the sizes and arrangements of islands in an archipelago, affects processes contributing to the generation and conservation of biodiversity patterns. In the first chapter, I integrate two disparate bodies of theory, ecological neutral theory and network theory into a powerful new framework for investigating patterns of biodiversity in a complex landscape. I examine the consequences of network structure, such as size, topology, and connectivity, for diversity patterning across the metacommunity. The second chapter focuses on how the position of a node within a network controls local community (node) diversity. Network statistics, such as node centrality, are found to predict diversity patterns with more central nodes accumulating the most diversity. In the third chapter, I use the theory to evaluate how well fundamental concepts in conservation biology perform when neutral metacommunity processes generate diversity patterns. I find that contemporary diversity patterns are poor predictors of the long-term capacity of a network to support diversity, challenging a host of conservation concepts and applications. In the fourth chapter, I consider biodiversity dynamics in a network with a different model of speciation, where spatial structure is needed for divergence. In this case, speciation hotspots form where the dispersal properties of an organism and the spatial structure of the landscape coincide. In the final chapter I study the biodiversity of a natural structured metacommunity, the ants of the Fijian archipelago. I used a variety of collecting techniques to inventory the ant species occurring across a system of islands in the southwest Pacific. Approximately 50 new species were discovered, and the distributions of the ant species across the islands are firmly established. Radiations are observed in the genera Pheidole, Camponotus, Lordomyrma, Leptogenys, Cerapachys, Strumigenys, Poecilomyrma, and Hypoponera. / text

Social parasitism, the exploitation of a society by other social organisms, has evolved independently numerous times within social animals. In this thesis, I integrate behavioral, evolutionary and chemical analyses to elucidate the evolution of social parasitism in Megalomyrmex ants. I examine host-parasite interactions in two Megalomyrmex species, identify venom alkaloids, and reconstruct the phylogenetic relationships between species. In Chapter 1, I analyze nest architecture and behavioral interactions between the ant host Cyphomyrmex cornutus and its parasite Megalomyrmex mondabora. This is the first detailed account of the natural history of this host and its social parasite. In Chapter 2, I report a one-year-long fitness experiment that tests whether Trachymyrmex cf. zeteki colonies suffer reduced fitness from an association with the social parasite Megalomyrmex symmetochus. I show that M. symmetochus parasites negatively impact host fitness though several mechanisms, including a) manipulation of the host worker grooming behavior; b) castration of host queens produced by the host colony, which then become workers; and c) reduction of garden size, host worker number, and host reproductive output. In Chapter 3, I determine that five venom alkaloids of Megalomyrmex are taxonomically informative to help differentiate cryptic species within the M. mondabora complex; new species in this complex need to be described in a future taxonomic revision. In Chapter 4, I reconstruct phylogenetic relationships of the genus Megalomyrmex with DNA sequence information. I conclude that the genus is monophyletic and corroborate two of the four species groups proposed by Brandão (1990) in a previous morphological revision. I also find evidence in support of Darwin’s Predation Hypothesis on the origin of social parasitism, which postulates that socialparasitic behaviors evolve from predatory behaviors. Lastly, I discuss promising future research directions on the evolution of social parasitism in the ant genus Megalomyrmex, which could serve as a model for the study of social parasitism in other lineages of social insects. / text