Fluctuating temperatures and ectotherm growth: distinguishing non-linear and time-dependent effects.

Most terrestrial ectotherms experience diurnal and seasonal variation in temperature. Because thermal performance curves are non-linear, mean performance can differ in fluctuating and constant thermal environments. However, time-dependent effects--effects of the order and duration of exposure to temperature--can also influence mean performance. We quantified the non-linear and time-dependent effects of diurnally fluctuating temperatures for larval growth rates in the tobacco hornworm, Manduca sexta L., with four main results. First, the shape of the thermal performance curve for growth rate depended on the duration of exposure: for example, optimal temperature and thermal breadth were greater for growth rates measured over short (24 h during the last instar) compared with long (the entire period of larval growth) time periods. Second, larvae reared in diurnally fluctuating temperatures had significantly higher optimal temperatures and maximal growth rates than larvae reared in constant temperatures. Third, for larvae maintained at three mean temperatures (20, 25 and 30°C) and three diurnal temperature ranges (±0, ±5 and ±10°C), diurnal fluctuations had opposite effects on mean growth rates at low versus high mean temperature. Fourth, both short- and long-term thermal performance curves yielded poor predictions of the non-linear effects of fluctuating temperature on mean growth rates (compared with our experimental results) at higher mean temperatures. Our results suggest caution in using constant temperature studies to model the consequences of variable thermal environments.