To maintain constant body temperature (T_b) over a wide range of ambient temperatures (T_a) endothermic animals require large amounts of energy and water. In hot environments, the main threat to endothermic homeotherms is insufficient water to supply that necessary for thermoregulation. We investigated flexible adjustment of traits related to thermoregulation and water conservation during acclimation to hot conditions or restricted water availability, or both, in the zebra finch,Taeniopygia guttataa small arid-zone passerine. Using indirect calorimetry, we measured changes in whole animal metabolic rate (MR), evaporative heat loss (EHL) andT_bbefore and after acclimation to 23 or 40 °C, with different availability of water. Additionally, we quantified changes in partitioning of EHL into respiratory and cutaneous avenues in birds exposed to 25 and 40 °C. In response to heat and water restriction zebra finches decreased MR, which together with unchanged EHL resulted in increased efficiency of evaporative heat loss. This facilitated more preciseT_bregulation in heat-acclimated birds. Acclimation temperature and water availability had no effect on the partitioning of EHL into cutaneous or respiratory avenues. At 25 °C, cutaneous EHL accounted for ~ 60% of total EHL, while at 40 °C, its contribution decreased to ~ 20%. Consistent among-individual differences in MR and EHL suggest that these traits, provided that they are heritable, may be a subject to natural selection. We conclude that phenotypic flexibility in metabolic heat production associated with acclimation to hot, water-scarce conditions is crucial in response to changing environmental conditions, especially in the face of current and predicted climate change.

SUMMARY: Social thermoregulation and huddling bring about energy benefits to animals sharing a nest because of the smaller surface-to-volume ratio of a huddle and the higher local temperature in the nest. We tested whether living in groups and huddling affect daily torpor, metabolic rate and seasonal changes in the body mass of a small heterothermic rodent, the Siberian hamster (Phodopus sungorus), housed under semi-natural conditions both singly and in groups of four litter-mates. We predicted that in hamsters housed in groups: (1) synchronized torpor bouts would be longer and deeper than non-synchronized ones but shallower than in solitary hamsters, (2) seasonal variations in metabolic rate would be lower than in solitary hamsters, and (3) the winter decrease in body mass would be smaller in grouped than in singly housed hamsters. We found that group housing led to a smaller decrease in body mass in winter, and affected the length and depth of daily torpor. In group-living hamsters more than 50% of all torpor episodes were synchronized and torpid animals were often found in huddles formed of all cage-mates. The longest and deepest torpor bouts in groups were recorded when all animals in a group entered torpor simultaneously. Although the minimum body temperature during torpor was higher, torpor duration was slightly longer than in solitary hamsters. We did not record significant differences in the body mass-adjusted rate of oxygen consumption between solitary and grouped animals, either in the cold or at the lower critical temperature. We conclude that social thermoregulation enables maintenance of a larger body mass, and thus a larger body fat content, which can ensure better body condition at the beginning of the reproductive season.

As photoperiod shortens with the approach of winter, small mammals should reduce their energy expenditure to survive periods of food limitation. However, within seasons, animals should balance their energy budgets as abiotic conditions change, sometimes unpredictably; cold spells should increase heat production, while warm spells should do the opposite. Therefore, we addressed specific questions about the possible interactions between seasonal acclimatization and the intra-seasonal phenotypic flexibility of metabolic rate. We hypothesized that phenotypic flexibility in small mammals differs seasonally and is greater in summer than in winter, and predicted that seasonal adjustments in energetics, which are driven by photoperiod, overwhelm the influence of variations in the thermal environment. We measured body mass, basal metabolic rate (BMR), facultative non-shivering thermogenesis (fNST), body temperature, and calculated minimum thermal conductance in Siberian hamsters Phodopus sungorus. Animals were acclimated to winter-like, and then to summer-like conditions and, within each season, were exposed twice, for 3 weeks to 10, 20 or 28 °C. We used differences between values measured after these short acclimation periods as a measure of the scope of phenotypic flexibility. After winter acclimation, hamsters were lighter, had lower whole animal BMR, higher fNST than in summer, and developed heterothermy. After these short acclimations to the above-mentioned temperatures, hamsters showed reversible changes in BMR and fNST; however, these traits were less flexible in winter than in summer. We conclude that seasonal acclimation affects hamster responses to intra-seasonal variations in the thermal environment. We argue that understanding seasonal changes in phenotypic flexibility is crucial for predicting the biological consequences of global climate changes.

The success of migration of small passerine birds depends largely on effective refueling at stopover sites. In our previous studies, we found that hypothermia facilitates accumulation of fuel at the beginning of a stopover. Later we found that blackcaps, Sylvia atricapilla, might further reduce their energy expenditure by huddling while at rest. Here, we report experimental results supporting our hypothesis that huddling is beneficial to small migrating passerines both from energetic and thermoregulatory points of view. To test this hypothesis we measured metabolic rates and body temperatures of seven blackcaps placed in respirometry chambers overnight, either solitarily or in groups of three or four at ambient temperatures of 5, 10, and 15°C. Concurring with our predictions, huddling blackcaps maintained higher body temperatures than did solitary birds, but had mass-specific metabolic rates lower by ∼30% than those of solitary individuals. Based on our previous studies, we estimated energy savings through huddling to be comparable to energy savings through hypothermia in solitary blackcaps and suggest that huddling may be an important way of saving energy for small passerine birds resting at migratory stopovers. At the same time it might offer the additional benefit of lower risk of predation. In this light, we predict that huddling occurs frequently in nature, leading to significant savings of energy, faster accumulation of fuel, presumably lower risk of becoming a prey, more successful migration, and eventually increased fitness.