Body size distributions of the pale grass blue butterfly in Japan: Size rules and the status of the Fukushima population.

The body size of the pale grass blue butterfly, Zizeeria maha, has been used as an environmental indicator of radioactive pollution caused by the Fukushima nuclear accident. However, geographical and temporal size distributions in Japan and temperature effects on size have not been established in this species. Here, we examined the geographical, temporal, and temperature-dependent changes of the forewing size of Z. maha argia in Japan. Butterflies collected in 2012 and 2013 from multiple prefectures throughout Japan demonstrated an inverse relationship of latitude and forewing size, which is the reverse of Bergmann's cline. The Fukushima population was significantly larger than the Aomori and Miyagi populations and exhibited no difference from most of the other prefectural populations. When monitored at a single geographic locality every other month, forewing sizes were the largest in April and the smallest in August. Rearing larvae at a constant temperature demonstrated that forewing size followed the temperature-size rule. Therefore, the converse Bergmann's rule and the temperature-size rule coexist in this multivoltine species. Our study establishes this species as a useful environmental indicator and supports the idea that the size reduction observed only in Fukushima Prefecture in 2011 was caused by the environmental stress of radioactive pollution.