Testing multiple hypotheses on the colour change of treefrogs in response to various external conditions.

Amphibians are famous for their ability to change colours. And a considerable number of studies have investigated the internal and external factors that affect the expression of this phenotypic plasticity. Evidence to date suggests that thermoregulation and camouflage are the main pressures that influence frogs' adaptive colour change responses. However, certain gaps in our knowledge of this phenomenon remain, namely: (i) how do frogs adjust their colour in response to continuously changing external conditions?; (ii) what is the direction of change when two different functions of colour (camouflage and thermoregulation) are in conflict?; (iii) does reflectance in the near-infrared region show thermally adaptive change?; and (iv) is the colour change ability of each frog an individual trait (i.e., consistent within an individual over time)? Using Dryophytes japonicus (Hylidae, Hyla), we performed a series of experiments to answer the above questions. We first showed that frogs' responses to continuously-changing external conditions (i.e., background colour and temperature) were not linear and limited to the range they experience under natural conditions. Second, when a functional conflict existed, camouflage constrained the adaptive response for thermoregulation and vice versa. Third, though both temperature and background colour induced a change in near-infrared reflectance, this change was largely explained by the high correlation between colour (reflectance in the visible spectrum) and near-infrared reflectance. Fourth, within-individual variation in colour change capacity (i.e., the degree of colour change an individual can display) was lower than inter-individual variation, suggesting individuality of colour change capacity; however, we also found that colour change capacity could change gradually with time within individuals. Our results collectively reveal several new aspects of how evolution shapes the colour change process and highlight how variation in external conditions restricts the extent of colour change in treefrogs.