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Beyond Nector: Unravelling Puddling in Butterflies

Puddling is a specialized nutrient-acquisition behaviour in butterflies and several other insect groups, where individuals visit mineral-rich substrates such as moist soil, dung, carrion and animal secretions. Since nectar and plant sap are poor in essential minerals particularly sodium puddling enables insects to supplement their diet with sodium, nitrogen and amino acids necessary for physiological and reproductive functions. Male butterflies puddle more frequently than females, as the sodium obtained is incorporated into spermatophores, enhancing female fecundity and offspring fitness. Puddling behaviour varies across species and families, with Lycaenidae exhibiting the highest frequency, followed by Papilionidae and Nymphalidae. Aggregation patterns range from solitary individuals to mixed-species groups, influenced by substrate quality, chemical cues and social stimuli. Environmental factors such as sunlight, moisture, humidity and mineral concentration strongly shape site selection. Puddling is also documented in non-lepidopteran insects, including Orthoptera, Hymenoptera, Diptera and Blattodea, indicating its broader ecological significance. Overall, puddling represents a multifaceted behaviour integrating nutritional demand, reproductive strategies and environmental interactions and remains an important subject for understanding insect ecology and evolutionary adaptation.