Anthropogenic sodium influences butterfly responses to nitrogen-enriched resources: implications for the nitrogen limitation hypothesis

Humans are increasing the environmental availability of historically limited nutrients, which may significantly influence organismal performance and behavior. Beneficial or stimulatory responses to increases in nitrogen availability (i.e., nitrogen limitation) are generally observed in plants but less consistently in animals. One possible explanation is that animal responses to nitrogen enrichment depend on how nitrogen intake is balanced with sodium, a micronutrient crucial for animals but not plants. We tested this idea in the cabbage white butterfly (Pieris rapae), a species that frequently inhabits nutrient-enriched plants in agricultural settings and roadside verges. We asked (1) whether anthropogenic increases in sodium influence how nitrogen enrichment affects butterfly performance and (2) whether individuals can adaptively adjust their foraging behavior to such effects. Larval nitrogen enrichment enhanced growth of cabbage white larvae under conditions of low but not high sodium availability. In contrast, larval nitrogen enrichment increased egg production of adult females only when individuals developed with high sodium availability. Ovipositing females preferred nitrogen-enriched leaves regardless of sodium availability, while larvae avoided feeding on nitrogen-enriched leaves elevated in sodium. Our results show that anthropogenic increases in sodium influence whether individuals benefit from and forage on nitrogen-enriched resources. Yet, different nitrogen-to-sodium ratios are required to optimize larval and adult performance. Whether increases in sodium catalyze or inhibit benefits of nitrogen enrichment may depend on how evolved nutrient requirements vary across stages of animal development.