Spatial variation in the evolutionary potential and constraints of basal metabolic rate and body mass in a wild bird.

An organism's energy budget is strongly related to resource consumption, performance, and fitness. Hence, understanding the evolution of key energetic traits, such as basal metabolic rate (BMR), in natural populations is central for understanding life-history evolution and ecological processes. Here we used quantitative genetic analyses to study evolutionary potential of BMR in two insular populations of the house sparrow (Passer domesticus). We obtained measurements of BMR and body mass (Mb ) from 911 house sparrows on the islands of Leka and Vega along the coast of Norway. These two populations were the source populations for translocations to create an additional third, admixed 'common garden' population in 2012. With the use of a novel genetic group animal model concomitant with a genetically determined pedigree, we differentiate genetic and environmental sources of variation, thereby providing insight into the effects of spatial population structure on evolutionary potential. We found that the evolutionary potential of BMR was similar in the two source populations, whereas the Vega population had a somewhat higher evolutionary potential of Mb than the Leka population. BMR was genetically correlated with Mb in both populations, and the conditional evolutionary potential of BMR (independent of body mass) was 41% (Leka) and 53% (Vega) lower than unconditional estimates. Overall, our results show that there is potential for BMR to evolve independently of Mb , but that selection on BMR and/or Mb may have different evolutionary consequences in different populations of the same species.

High latitude northern bats (Eptesicus nilssonii) reveal adaptations to both high and low ambient temperatures.

Insectivorous bats at northern latitudes need to cope with long periods of no food for large parts of the year. Hence, bats which are resident at northern latitudes throughout the year will need to undergo a long hibernation season and a short reproductive season where foraging time is limited by extended daylight periods. Eptesicus nilssonii is the northernmost occurring bat species worldwide and hibernates locally when ambient temperatures (Ta) limit prey availability. Therefore, we investigated the energy spent maintaining normothermy at different Ta, as well as how much bats limit energy expenditure while in torpor. We found that, despite being exposed to Ta as low as 1.1°C, bats did not increase torpid metabolic rate, thus indicating that E. nilssonii can survive and hibernate at low ambient temperatures. Furthermore, we found a lower critical temperature (Tlc) of 27.8°C, which is lower than in most other vespertilionid bats, and we found no indication of any metabolic response to Ta up to 37.1°C. Interestingly, carbon dioxide production increased with increasing Ta above the Tlc, presumably caused by a release of retained CO2 in bats that remained in torpor for longer and aroused at Ta above the Tlc. Our results indicate that E. nilssonii can thermoconform at near-freezing Ta, and hence maintain longer torpor bouts with limited energy expenditure, yet also cope with high Ta when sun exposed in roosts during long summer days. These physiological traits are likely to enable the species to cope with ongoing and predicted climate change.

The small-bat-in-summer paradigm: Energetics and adaptive behavioural routines of bats investigated through a stochastic dynamic model.

Strong seasonality at high latitudes represents a major challenge for many endotherms as they must balance survival and reproduction in an environment that varies widely in food availability and temperature. To avoid energetic mismatches caused by limited foraging time and stochastic weather conditions, bats employ the energy-saving state of torpor during summer to save accumulated energy reserves. However, at high-latitude small-bats-in-summer face a particular challenge: as nocturnal foragers, they rely on the darkness at night to avoid predators and/or interspecific competition, but live in an environment with short, light summer nights, and even a lack of true night at the northernmost distributions of some bat species. To predict optimal behaviour in relation to latitudinal variation in diurnal cycles, we constructed a stochastic dynamic programming model of bats living at high latitudes. Using a stochastic dynamic programming framework with values that are representative for our study system, we show that individual energetic reserves are a strong driver of daytime use of torpor and night-time foraging behaviour alike, with these linked effects being both temperature- and photoperiod-dependent. We further used the model to predict survival probabilities at five locations across a latitudinal gradient (60.1° N to 70.9° N), finding that combinations of photoperiod and temperature conditions limited population distributions in the model. To verify our model results, we compared predictions for optimal decisions with our own empirical data collected on northern bats (Eptesicus nilssonii) from two latitudes in Norway. The similarities between our predictions and observations provide strong evidence that this model framework incorporates the most important drivers of diurnal decision-making in bat physiology and behaviour. Comparing empirical data and model predictions also revealed that bats facing lighter night conditions further north restrict their mass gain, which strengthens the hypothesis that predation threat is a main driver of bat nocturnality. Our model findings regarding state-dependent decisions in bats should contribute to the understanding of how bats cope with the summer challenges at high latitudes.

Determining the different phases of torpor from skin- or body temperature data in heterotherms.

Technological innovations have made heat-sensitive data-loggers smaller, more efficient and less expensive, which has led to a growing body of literature that measures the skin- or body temperatures of small animals in their natural environments. Studies of this type on heterothermic endotherms have prompted much debate regarding how to best define 'torpor' expressions from skin- or body temperature data alone. We propose a new quantitative method for defining torpor 'entries', 'arousals' and 'stable torpor periods' whilst comparing the results to 'torpor bout' durations identified using only the torpor cut-off method. By decomposing a torpor bout into 'entries', 'stable torpor periods', and 'active arousals', we avoid biases introduced by using strict threshold temperature values for the onset of torpor, thereby allowing better insight into individual use of torpor. We present our method as an easy-to-use function written in R-code, offering an un-biased and consistent methodology to be applied on skin- or body temperature measurements across datasets and research groups. When testing the function on a large dataset of skin temperature data collected on three bat species in Norway (Plecotus auritus: Nind = 39; Eptesicus nilssonii: Nind = 11; Myotis brandtii: Nind = 10), we identified 461 complete torpor bouts across species. More than 40% of the torpor bouts (Nbouts = 192) did not contain stable torpor periods, because the bats aroused before they had reached a stable skin temperature level. Furthermore, only considering 'torpid' and 'euthermic' temperature values by applying strict cut-off thresholds led to potentially large underestimations of torpor bout durations compared to our quantitative determination of the onset and termination of each torpor bout. We highlight the importance of differentiating between torpor phases, especially for active arousals that can be very energetically expensive and may alter our evaluation of the actual energetic savings gained by an individual employing torpor.

Nightly torpor use in response to weather conditions and individual state in an insectivorous bat.

Torpor is a well-known energy conservation strategy in many mammal and bird species. It is often employed when environmental conditions are unfavourable to maximize survival probabilities. However, torpor often carries with it the physiological costs of a low body temperature and of rewarming in addition to potential missed opportunities for foraging. Therefore, we hypothesised that decision making regarding when to use torpor should reflect the most important environmental conditions for species distributions, and thus how they may be impacted by ongoing climate change. We investigated how weather conditions affect nightly torpor patterns in the nocturnal insectivorous Australian eastern long-eared bat (Nyctophilus bifax). By measuring the skin temperature of 37 free-ranging individuals, we confirmed that torpor was used more frequently during the winter and at subtropical compared to tropical locations. Using mixed-effect models we show that lower ambient temperatures were the main driver of individual torpor use, probably due to lower roost temperatures and prey availability. However, increased rain, wind and humidity, and decreasing barometric pressure, as well as brighter moonlight, also led to more time spent torpid per night. We suggest that bats evaluate multiple environmental cues to make decisions regarding torpor use versus active foraging based upon their expectations of the energetic benefits, prey availability and relative predation risk. Interactions between some of these effects and body mass (whilst controlling for forearm length) indicate that individual variation in body size and/or state-dependent effects of energy reserves also partly determined the use of nightly torpor in these bats.

How to quantify thermal acclimation capacity?

The capacity of organisms to acclimate will influence their ability to cope with ongoing global changes in thermal regimes. Here we highlight methodological issues associated with recent attempts to quantify variation in acclimation capacity among taxa and environments, and describe how these may introduce bias to conclusions. We then propose a measure of thermal acclimation capacity that more directly quantifies the process of acclimation. Future studies of variation in acclimation capacity should critically evaluate whether their chosen empirical metric accurately reflects the theoretical concept of acclimation.

Cold-hearted bats: uncoupling of heart rate and metabolism during torpor at sub-zero temperatures.

Many hibernating animals thermoregulate during torpor and defend their body temperature (Tb) near 0°C by an increase in metabolic rate. Above a critical temperature (Tcrit), animals usually thermoconform. We investigated the physiological responses above and below Tcrit for a small tree-dwelling bat (Chalinolobus gouldii, ∼14 g) that is often exposed to sub-zero temperatures during winter. Through simultaneous measurement of heart rate (fH) and oxygen consumption (V̇O2 ), we show that the relationship between oxygen transport and cardiac function is substantially altered in thermoregulating torpid bats between 1 and -2°C, compared with thermoconforming torpid bats at mild ambient temperatures (Ta 5-20°C). Tcrit for this species was at a Ta of 0.7±0.4°C, with a corresponding Tb of 1.8±1.2°C. Below Tcrit, animals began to thermoregulate, as indicated by a considerable but disproportionate increase in both fH and V̇O2 The maximum increase in fH was only 4-fold greater than the average thermoconforming minimum, compared with a 46-fold increase in V̇O2 The differential response of fH and V̇O2  to low Ta was reflected in a 15-fold increase in oxygen delivery per heart beat (cardiac oxygen pulse). During torpor at low Ta, thermoregulating bats maintained a relatively slow fH and compensated for increased metabolic demands by significantly increasing stroke volume and tissue oxygen extraction. Our study provides new information on the relationship between metabolism and fH in an unstudied physiological state that may occur frequently in the wild and can be extremely costly for heterothermic animals.

A new cue for torpor induction: charcoal, ash and smoke.

Recent work has shown that the use of torpor for energy conservation increases after forest fires in heterothermic mammals, probably in response to the reduction of food. However, the specific environmental cues for this increased torpor expression remain unknown. It is possible that smoke and the novel substrate of charcoal and ash act as signals for an impending period of starvation requiring torpor. We therefore tested the hypothesis that the combined cues of smoke, a charcoal/ash substrate and food shortage will enhance torpor expression in a small forest-dwelling marsupial, the yellow-footed antechinus (Antechinus flavipes), because like other animals that live in fire-prone habitats they must effectively respond to fires to ensure survival. Activity and body temperature patterns of individuals in outdoor aviaries were measured under natural environmental conditions. All individuals were strictly nocturnal, but diurnal activity was observed shortly after smoke exposure. Overall, torpor in females was longer and deeper than that in males. Interestingly, while both males and females increased daily torpor duration during food restriction by >2-fold as anticipated, a combination of food restriction and smoke exposure on a charcoal/ash substrate further increased daily torpor duration by ∼2-fold in both sexes. These data show that this combination of cues for torpor induction is stronger than food shortage on its own. Our study provides significant new information on how a small forest-dwelling mammal responds to fire cues during and immediately after a fire and identifies a new, not previously recognised, regulatory mechanism for thermal biology in mammals.

Post-fire recovery of torpor and activity patterns of a small mammal.

To cope with the post-fire challenges of decreased availability of food and shelter, brown antechinus (Antechinus stuartii), a small marsupial mammal, increase the use of energy-conserving torpor and reduce activity. However, it is not known how long it takes for animals to resume pre-fire torpor and activity patterns during the recovery of burnt habitat. Therefore, we tested the hypothesis that antechinus will adjust torpor use and activity after a fire depending on vegetation recovery. We simultaneously quantified torpor and activity patterns for female antechinus from three adjacent areas: (i) the area of a management burn 1 year post-fire, (ii) an area that was burned 2 years prior, and (iii) a control area. In comparison to shortly after the management burn, antechinus in all three groups displayed less frequent and less pronounced torpor while being more active. We provide the first evidence that only 1 year post-fire antechinus resume pre-fire torpor and activity patterns, probably in response to the return of herbaceous ground cover and foraging opportunities.

Can hibernators sense and evade fires? Olfactory acuity and locomotor performance during deep torpor.

Increased habitat fragmentation, global warming and other human activities have caused a rise in the frequency of wildfires worldwide. To reduce the risks of uncontrollable fires, prescribed burns are generally conducted during the colder months of the year, a time when in many mammals torpor is expressed regularly. Torpor is crucial for energy conservation, but the low body temperatures (T b) are associated with a decreased responsiveness and torpid animals might therefore face an increased mortality risk during fires. We tested whether hibernators in deep torpor (a) can respond to the smell of smoke and (b) can climb to avoid fires at T bs below normothermic levels. Our data show that torpid eastern pygmy-possums (Cercartetus nanus) are able to detect smoke and also can climb. All males aroused from torpor when the smoke stimulus was presented at an ambient temperature (T a) of 15 °C (T b ∼18 °C), whereas females only raised their heads. The responses were less pronounced at T a 10 °C. The first coordinated movement of possums along a branch was observed at a mean T b of 15.6 °C, and animals were even able to climb their prehensile tail when they reached a mean T b of 24.4 °C. Our study shows that hibernators can sense smoke and move at low T b. However, our data also illustrate that at T b ≤13 °C, C. nanus show decreased responsiveness and locomotor performance and highlight that prescribed burns during winter should be avoided on very cold days to allow torpid animals enough time to respond.

Black or white? Physiological implications of roost colour and choice in a microbat.

Although roost choice in bats has been studied previously, little is known about how opposing roost colours affect the expression of torpor quantitatively. We quantified roost selection and thermoregulation in a captive Australian insectivorous bat, Nyctophilus gouldi (n=12) in winter when roosting in black and white coloured boxes using temperature-telemetry. We quantified how roost choice influences torpor expression when food was provided ad libitum or restricted in bats housed together in an outdoor aviary exposed to natural fluctuations of ambient temperature. Black box temperatures averaged 5.1°C (maximum 7.5°C) warmer than white boxes at their maximum daytime temperature. Bats fed ad libitum chose black boxes on most nights (92.9%) and on 100% of nights when food-restricted. All bats used torpor on all study days. However, bats fed ad libitum and roosting in black boxes used shorter torpor and spent more time normothermic/active at night than food-restricted bats and bats roosting in white boxes. Bats roosting in black boxes also rewarmed passively more often and to a higher skin temperature than those in white boxes. Our study suggests that N. gouldi fed ad libitum select warmer roosts in order to passively rewarm to a higher skin temperature and thus save energy required for active midday rewarming as well as to maintain a normothermic body temperature for longer periods at night. This study shows that colour should be considered when deploying bat boxes; black boxes are preferable for those bats that use passive rewarming, even in winter when food availability is reduced.

Evolution of basal metabolic rate in bank voles from a multidirectional selection experiment.

A major theme in evolutionary and ecological physiology of terrestrial vertebrates encompasses the factors underlying the evolution of endothermy in birds and mammals and interspecific variation of basal metabolic rate (BMR). Here, we applied the experimental evolution approach and compared BMR in lines of a wild rodent, the bank vole (Myodes glareolus), selected for 11 generations for: high swim-induced aerobic metabolism (A), ability to maintain body mass on a low-quality herbivorous diet (H) and intensity of predatory behaviour towards crickets (P). Four replicate lines were maintained for each of the selection directions and an unselected control (C). In comparison to C lines, A lines achieved a 49% higher maximum rate of oxygen consumption during swimming, H lines lost 1.3 g less mass in the test with low-quality diet and P lines attacked crickets five times more frequently. BMR was significantly higher in A lines than in C or H lines (60.8, 56.6 and 54.4 ml O2 h(-1), respectively), and the values were intermediate in P lines (59.0 ml O2 h(-1)). Results of the selection experiment provide support for the hypothesis of a positive association between BMR and aerobic exercise performance, but not for the association of adaptation to herbivorous diet with either a high or low BMR.

The importance of mammalian torpor for survival in a post-fire landscape.

Wildfires have increased in frequency and intensity worldwide with climate change as a main driving factor. While a number of studies have focused on population changes in regard to fires, there are essentially no quantitative data on behavioural and physiological adjustments that are vital for the persistence of individuals during and after fires. Here we show that brown antechinus, a small insectivorous marsupial mammal, (i) endured a prescribed fire in situ, (ii) remained in their scorched home range despite unburned areas nearby, and (iii) substantially increased post-fire torpor use and thus reduced foraging requirements and exposure to predators. Hence, torpor is a physiological adaptation that, although not quantified in this context previously, appears to play a key role in post-fire survival for this and other heterothermic species.

Selection for high activity-related aerobic metabolism does not alter the capacity of non-shivering thermogenesis in bank voles.

An intriguing question is how the capacity of non-shivering thermogenesis (NST)-a special mechanism supporting endothermic thermoregulation in mammals-is affected by selection for high exercise metabolism. It has been proposed that high NST could be a mechanism to compensate for a low basal production of heat. On the other hand, high basal or activity metabolism is associated with physiological characteristics such as high performance of the circulatory system, which are also required for achieving a high NST. Here we tested whether selection for high aerobic exercise performance, which correlates with an increased basal metabolic rate, led to a correlated evolution of maximum and facultative NST. Therefore, we measured the NST of bank voles, Myodes (= Clethrionomys) glareolus, from lines selected for 13-14 generations (n=46) for high aerobic metabolism achieved during swimming and from unselected, control lines (n=46). Open-flow respirometry was used to measure the rate of oxygen consumption (V(·)O2) in anesthetized bank voles injected with noradrenaline (NA). After adjusting for body mass, maximum NST (maximum V(·)O2 recorded after injection of NA) did not differ between the selected (2.38±0.08 mLO2min(-1)) and control lines (2.36±0.08 mLO2min(-1); P=0.891). Facultative NST (= maximum NST minus resting metabolic rate of anesthetized animals) did not differ between the selected (1.49±0.07 mLO2min(-1)) and control lines (1.50±0.07 mLO2min(-1); P=0.985), either. Therefore, our results suggest that NST capacity is not strongly linked to maximum activity-related aerobic metabolic rate.

Rare and Opportunistic Use of Torpor in Mammals-An Echo from the Past?

Torpor was traditionally seen as a winter survival mechanism employed by animals living in cold and highly seasonal habitats. Although we now know that torpor is also used by tropical and subtropical species, and in response to a variety of triggers, torpor is still largely viewed as a highly controlled, seasonal mechanism shown by Northern hemisphere species. To scrutinize this view, we report data from a macroanalysis in which we characterized the type and seasonality of torpor use from mammal species currently known to use torpor. Our findings suggest that predictable, seasonal torpor patterns reported for Northern temperate and polar species are highly derived forms of torpor expression, whereas the more opportunistic and variable forms of torpor that we see in tropical and subtropical species are likely closer to the patterns expressed by ancestral mammals. Our data emphasize that the torpor patterns observed in the tropics and subtropics should be considered the norm and not the exception.

Species and reproductive status influence element concentrations in bat fur.

To assess the impact of increasing pollutant levels on wildlife, we measured chemical contaminant loads in bats with different habitat and dietary preferences. Samples were taken from the fur of bats (Eptesicus nilssonii, Myotis brandtii, Myotis mystacinus and Plecotus auritus) to measure concentrations of 55 elements by inductively coupled plasma mass spectrometry (ICP-MS). Variations in element concentrations between different bat groups (species, sex, reproductive status) were analysed with a focus on arsenic (As), mercury (Hg) and lead (Pb) as these are known to cause specific health concerns in wildlife. For M. brandtii we found the highest As concentrations, especially in lactating bats, with a maximum exceeding those from other studies where bats had compromised health. Whereas for M. mystacinus there was a negative correlation between body condition index (BCI) and As concentration, indicating a potential danger for bats in the study area. In M. mystacinus and M. brandtii Hg concentrations were higher for sixteen individuals than in other studies where bats suffered genotoxic effects, although median levels were still below this threshold. Lactating bats from P. auritus and M. brandtii had higher Hg concentrations than bats of other reproductive status, which could endanger offspring as Hg can be transferred through lactation. In females from M. mystacinus Pb concentrations were more than three times higher compared to males. There was also a negative correlation between Pb concentration and BCI, which could mean that Pb has an adverse effect on health. Although many other biotic and abiotic factors should be considered, some of the variations in element concentrations could be due to different behaviours (foraging, roosting, etc.) in the studied species. The high levels of chemical contamination in some of the bats in our study, particularly reproductive individuals, is of conservation concern as bats are important agents for insect control.

A species-level trait dataset of bats in Europe and beyond.

Knowledge of species' functional traits is essential for understanding biodiversity patterns, predicting the impacts of global environmental changes, and assessing the efficiency of conservation measures. Bats are major components of mammalian diversity and occupy a variety of ecological niches and geographic distributions. However, an extensive compilation of their functional traits and ecological attributes is still missing. Here we present EuroBaTrait 1.0, the most comprehensive and up-to-date trait dataset covering 47 European bat species. The dataset includes data on 118 traits including genetic composition, physiology, morphology, acoustic signature, climatic associations, foraging habitat, roost type, diet, spatial behaviour, life history, pathogens, phenology, and distribution. We compiled the bat trait data obtained from three main sources: (i) a systematic literature and dataset search, (ii) unpublished data from European bat experts, and (iii) observations from large-scale monitoring programs. EuroBaTrait is designed to provide an important data source for comparative and trait-based analyses at the species or community level. The dataset also exposes knowledge gaps in species, geographic and trait coverage, highlighting priorities for future data collection.

Universality of Torpor Expression in Bats.

AbstractAlthough heterothermy is employed by species at a global level within the order of Chiroptera (bats), the possibility of torpor being expressed in bat species inhabiting warmer climate zones has been explored only in the past couple decades. Recent studies suggest that the benefit of expressing torpor is not limited to saving energy during cold exposure or food shortage but may be just as important for saving water during heat waves. Thus, even if the physiological challenges faced by bats may depend on the habitat they live in, species expressing torpor should be found in any climate zone where employing torpor may yield benefits and increase their survival probability. Here, we summarize available data on torpor metabolic rates and daily skin temperature patterns of bats across climate zones, emphasizing similarities found in the data. We also present data that we have collected from a southern subtropical species (Nyctophilus bifax) and a northern subarctic species (Plecotus auritus) to illustrate specific examples of torpor expressions in two bat species living in highly different environments. Our findings highlight that torpor metabolic rates and skin temperature patterns of bats outside of the hibernation season can be universal across vastly different habitats, although arid environments indicate potential divergence in mean minimum torpor metabolic rates compared with measurements of populations inhabiting other climate zones.

Thermal biology and roost selection of free-ranging male little forest bats, Vespadelus vulturnus, during winter.

Insectivorous bats are particularly susceptible to heat loss due to their relatively large surface area to volume ratio. Therefore, to maintain a high normothermic body temperature, bats require large amounts of energy for thermoregulation. This can be energetically challenging for small bats during cold periods as heat loss is augmented and insect prey is reduced. To conserve energy many bats enter a state of torpor characterized by a controlled reduction of metabolism and body temperature in combination with selecting roosts based upon thermal properties. Our study aimed to quantify torpor patterns and roost preferences of free-ranging little forest bats (Vespadelus vulturnus) during winter to identify physiological and behavioral mechanisms used by this species for survival of the cold season. All bats captured were male (body mass 4.9 ± 0.7 g, n = 6) and used torpor on every day monitored, with bouts lasting up to 187.58 h (mean = 35.5 ± 36.7 h, n = 6, total number of samples [N] = 61). Torpor bout duration was significantly correlated with daily minimum and maximum ambient temperature, mean skin temperature, insect mass, and body mass of individuals and the multiday torpor bouts recorded in the cold qualify as hibernation. The lowest skin temperature recorded was 5.2°C, which corresponded to the lowest ambient temperature measurement of -5.8°C. Most bats chose tall, large, live Eucalyptus trees for roosting and to leave their roost for foraging on warmer days. Many individuals often switched roosts (every 3-5 days) and movements increased as spring approached (every 1-2 days). Our data suggest that V. vulturnus are capable of using the environmental temperature to gauge potential foraging opportunities and as a cue to reenter torpor when conditions are unsuitable. Importantly, frequent use of torpor and appropriate roost selection form key roles in the winter survival of these tiny bats.

State dependence of arousal from torpor in brown long-eared bats (Plecotus auritus).

To cope with periods of low food availability and unsuitable environmental conditions (e.g., short photoperiod or challenging weather), many heterothermic mammals can readily go into torpor to save energy. However, torpor also entails several potential costs, and quantitative energetics can, therefore, be influenced by the individual state, such as available energy reserves. We studied the thermal energetics of brown long-eared bats (Plecotus auritus) in the northern part of its distributional range, including torpor entry, thermoregulatory ability during torpor and how they responded metabolically to an increasing ambient temperature (Ta) during arousal from torpor. Torpor entry occurred later in bats with higher body mass (Mb). During torpor, only 10 out of 21 bats increased oxygen consumption (V̇O2) to a greater extent above the mean torpor metabolic rates (TMR) when exposed to low Ta. The slope of the torpid thermoregulatory curve was shallower than that of resting metabolic rate (RMR) during normothermic conditions, indicating a higher thermal insulation during torpor. During exposure to an increasing Ta, all bats increased metabolic rate exponentially, but the bats with higher Mb aroused at a lower Ta than those with lower Mb. In bats with low Mb, arousal was postponed to an Ta above the lower critical temperature of the thermoneutral zone. Our results demonstrate that physiological traits, which are often considered fixed, can be more flexible than previously assumed and vary with individual state. Thus, future studies of thermal physiology should to a greater extent take individual state-dependent effects into account.