Multiple cues influence multiple traits in the phenotypically plastic melanization of the cabbage white butterfly

Phenotypic plasticity, or the ability of organisms to produce different phenotypes depending upon environmental factors, may be adaptive in varying environments. However, because environments differ in many ways and organisms consist of many traits perfect phenotype-environment matches are unlikely. Studies that investigate multiple interacting environmental factors and the plastic responses of multiple traits should increase our understanding of the limits of adaptive plasticity. We experimentally examined the effects of variation in temperature and photoperiod on the seasonally plastic, and likely adaptive, melanization of a temperate butterfly, Pieris rapae. Although several melanin-based traits changed in response to temperature and photoperiodic variation, these traits tended to fall into two 'trait groups' consisting of traits covarying positively. However, these two trait groups responded to environmental factors, particularly temperature, in independent and sometimes opposing ways, with one increasing and the other decreasing in melanization with increased temperature. In some cases, plastic responses were complex and non-linear. Furthermore, when temperature and photoperiod were manipulated orthogonally, we sometimes detected interactive effects on melanization. These complex responses to two environmental cues may reflect sub-optimal responses or may occur if the two cues together provide more reliable information about future conditions than would either cue alone. Our results highlight the limits of studies of phenotypic plasticity that consider only single environmental factors and limit treatments to just two levels.