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Evidence accumulates about the role of arbuscular mycorrhizal (AM) fungi in shaping plant communities, but little is known about the factors determining the biomass and coexistence of several types of AM fungi in a plant community. Here, using a consumer-resource framework that treats the relationship between plants and fungi as simultaneous, reciprocal exploitation, we investigated what patterns of dynamic preferential plant carbon allocation to empirically-defined fungal types (on-going partner choice) would be optimal for plants, and how these patterns depend on successional dynamics. We found that ruderal AM fungi can dominate under low steady-state nutrient availability, and competitor AM fungi can dominate at higher steady-state nutrient availability; these are conditions characteristic of early and late succession, respectively. We also found that dynamic preferential allocation alone can maintain a diversity of mutualists, suggesting that on-going partner choice is a new coexistence mechanism for mutualists. Our model can therefore explain both mutualist coexistence and successional strategy, providing a powerful tool to derive testable predictions.

Recent studies of mutualistic networks show that interactions between partners change across years. Both biological mechanisms and chance could drive these patterns, but the relative importance of these factors has not been separated. We established a field experiment consisting of 102 monospecific plots of 17 native plant species, from which we collected 6713 specimens of 52 bee species over four years. We used these data and a null model to determine whether bee species' foraging choices varied more or less over time beyond the variation expected by chance. Thus we provide the first quantitative definition of rewiring and fidelity as these terms are used in the literature on interaction networks. All 52 bee species varied in plant partner choice across years, but for 27 species this variation was indistinguishable from random partner choice. Another 11 species showed rewiring, varying more across years than expected by chance, while 14 species showed fidelity, indicating that they both prefer certain plant species and are consistent in those preferences across years. Our study shows that rewiring and fidelity both exist in mutualist networks, but that once sampling effects have been accounted for, they are less common than has been reported in the ecological literature.

Introduced species inevitably experience novel selection pressures in their new environments as a result of changes in mutualist and antagonist relationships. While most previous work has examined how escape from specialist enemies has influenced herbivore or pathogen resistance of exotic species, post-introduction shifts in exotic dependence on mutualists have not been considered. In a common environment, we compared dependence on AM fungi of North American and European populations of Hypericum perforatum (St. John's Wort), a forb native to Europe. Introduced North American populations responded less to inoculation with AM fungi than did European populations. Root architecture was strongly correlated with mycorrhizal response, and introduced populations had finer root architecture than native populations. Finally, introduced populations exhibited decreased root and increased reproductive allocation relative to European populations, consistent with a transition to a weedier life history; however, biomass allocation patterns were uncorrelated with mycorrhizal response. These findings are the first demonstration of a genetically based reduction of mycorrhizal dependence and shift in root architecture in an introduced species.

Multiplayer symbioses are common in nature, but our understanding of the ecological dynamics occurring in complex symbioses is limited. The tripartite mutualism between fungus-growing ants, their fungal cultivars, and antibiotic-producing bacteria exemplifies symbiotic complexity. Here we reveal how black yeasts, newly described symbionts of the ant-microbe system, compromise the efficiency of bacteria-derived antibiotic defense in fungus-growing ants. We found that symbiotic black yeasts acquire nutrients from the ants' bacterial mutualist, and suppress bacterial growth. Experimental manipulation of ant colonies and their symbionts shows that ants infected with black yeasts are significantly less effective at defending their fungus garden from Escovopsis, a prevalent and specialized pathogen. The reduction of mutualistic bacterial biomass on ants, likely caused by black yeast symbionts, apparently reduces the quantity of antibiotics available to inhibit the garden pathogen. Success of the ant-fungal mutualism is directly dependent on fungus garden health. Thus our finding that black yeasts compromise the ants' ability to deal with the garden parasite indicates that it is an integral component of the symbiosis. This is further evidence that a full understanding of symbiotic associations requires examining the direct and indirect interactions of symbionts in their ecological community context.

Endophytes are fungi that form inapparent infections within leaves and stems of healthy plants. Closely related to virulent pathogens but with limited, if any, pathogenic effects themselves, many endophytes protect host plants from natural enemies. Animal herbivores and, in some cases, pathogenic microbes are poisoned by the mycotoxins produced by endophytes. "Constitutive mutualism" is the relatively faithful association, usually with grasses, of endophytes that infect host ovules and are propagated in host seed; substantial fungal biomass with probable high metabolic cost develops throughout the aerial parts of the host plant. "Inducible mutualist" endophytes are not involved with host seed and disseminate independently through air or in water. Infecting only vegetative parts of the host and remaining metabolically inactive for long periods with relatively little fungal biomass, inducible mutualists grow rapidly and produce toxins against herbivores when damaged host tissues provide new sites for infection. I surmise that endophytes may be as common among plants as are mycorrhyzae.