Heated debate: Is the seasonal polyphenism of Araschnia levana (Lepidoptera: Nymphalidae) driven by thermoregulation?

Abstract

The seasonal forms of the temperate butterfly Araschnia levana (Nymphalidae: Nymphalinae) differ in morphology (weight, wing area, and wing loading) and colouration. Spring individuals are predominantly orange with higher weight per wing area, (i.e. wing loading) while summer individuals are black with a white stripe and have lower wing loading. However, it remains unclear if and how these seasonal differences affect heating and cooling dynamics. We compared thermal responses of seasonal forms, focusing on the roles of morphology and colouration. Further, we assessed whether live butterflies heat and cool differently from dead individuals to detect the presence of active thermoregulation. Morphology and colouration influenced the thermal dynamics of the thorax and wings as expected from heat-transfer principles, but we found no evidence of active thermoregulation on the thorax. Based on aligned temperature curves, seasonal forms showed similar thermal dynamics. This similarity was driven by morphology and colouration, with larger wing area accelerating thermal change and higher body weight (or wing loading) reducing it, thereby masking underlying form-specific patterns. After accounting for significant morphological differences between forms, the thorax of spring individuals heated and cooled faster than that of summer ones. This trend suggests form-specific optimisation of thermal performance, likely as a response to temperate climates. Thermal responses differ between forms in ways not directly explained by the polyphenism itself, potentially reflecting a broader trait of multivoltine ectotherms to cope with seasonal temperature changes.