Chemically-mediated interactions in salt marshes: mechanisms that plant communities use to deter closely associated herbivores and pathogens

Herbivores and pathogens pose a consistent threat to plant productivity. In response, plants invest in structural and/or chemical defenses that minimize damage caused by these biotic stressors. In salt marshes along the Atlantic coast of the United States, a facultative mutualism between snails (Littoraria irrorata) and multiple species of fungi exert intense top-down control of the foundation grass species Spartina alterniflora. Since exposure to herbivores and pathogens are tightly coupled in this system, I investigated whether S. alterniflora utilizes chemical and/or structural defenses to deter both snails and fungi, and examined how plant defenses varied among S. alterniflora individuals and populations. I also assessed how other marsh plants prevent snails from establishing farms, and considered whether interspecific variation in plant chemical defenses influences marsh community structure. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that plant secondary chemistry was the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that differences in plant chemistry were the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability against S. alterniflora. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future.Ph