The extraordinary abundance of ants in tropical rainforest canopies has led to speculation that numerous arboreal ant taxa feed principally as "herbivores" of plant and insect exudates. Based on nitrogen (N) isotope ratios of plants, known herbivores, arthropod predators, and ants from Amazonia and Borneo, we find that many arboreal ant species obtain little N through predation and scavenging. Microsymbionts of ants and their hemipteran trophobionts might play key roles in the nutrition of taxa specializing on N-poor exudates. For plants, the combined costs of biotic defenses and herbivory by ants and tended Hemiptera are substantial, and forest losses to insect herbivores vastly exceed current estimates.
Feeding extensively on plant exudates and honeydews, many tropical arboreal ant species exhibit δ 15 N values characteristic of herbivores. Consistent with hypothesized herbivory, these taxa behave in feeding assays as though more N-deprived than are strictly carnivorous ants. However, to an as yet uncertain degree, relationships with N-upgrading and/or recycling microsymbionts may lower isotopic ratios, making ants appear to be more herbivorous than they actually are. Nutritional (N) contributions from microsymbionts have been inferred for a variety of ant taxa based on intracellular or extracellular associations between ants and bacteria. However, stronger and more specific inferences are possible when variability in microsymbiont locations within the digestive system is considered in the context of taxonomic variability in ant diets and digestive anatomy. Diets of exudate feeders may vary predictably in ratios of usable carbohydrates (CHOs) to N, depending on the extent to which they tend melezitose-producing Homoptera. Status of the peritrophic membrane, proventricular structure, and number and placement of Malpighian tubules can be interpreted as traits contributing to supply of N and/or CHOs to microsymbionts. In general, a more integrative understanding of ant diets, digestive anatomy, and associated microsymbionts helps to set out specific hypotheses to be tested experimentally and (where possible) in a phylogenetic context.
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