Ecological implications of metabolic compensation at low temperatures in salamanders.

Global warming is influencing the biology of the world's biota. Temperature increases are occurring at a faster pace than that experienced by organisms in their evolutionary histories, limiting the organisms' response to new conditions. Mechanistic models that include physiological traits can help predict species' responses to warming. Changes in metabolism at high temperatures are often examined; yet many species are behaviorally shielded from high temperatures. Salamanders generally favor cold temperatures and are one of few groups of metazoans to be most species-rich in temperate regions. I examined variation in body temperature, behavioral activity, and temperature dependence of resting heart rate, used as a proxy for standard metabolic rate, in fire salamanders (Salamandra salamandra). Over 26 years, I found that salamanders are behaviorally active at temperatures as low as 1 °C, and aestivate at temperatures above 16 °C. Infrared thermography indicates limited thermoregulation opportunities for these nocturnal amphibians. Temperature affects resting heart rate, causing metabolic depression above 11 °C, and metabolic compensation below 8 °C: heart rate at 3 °C is 224% the expected heart rate. Thus, salamanders operating at low temperatures during periods of peak behavioral activity are able to maintain a higher metabolic rate than the rate expected in absence of compensation. This compensatory mechanism has important ecological implications, because it increases estimated seasonal heart rates. Increased heart rate, and thus metabolism, will require higher caloric intake for field-active salamanders. Thus, it is important to consider a species performance breadth over the entire temperature range, and particularly low temperatures that are ecologically relevant for cold tolerant species such as salamanders.