Cutaneous and respiratory water loss in larks from arid and mesic environments

Birds from deserts generally have lower total evaporative water loss (TEWL), the sum of cutaneous (CWL) and respiratory water loss (RWL), than species from mesic areas. We investigated the role of CWL and RWL as a function of air temperature (T(a)) in hoopoe larks (Alaemon alaudipes) and Dunn's larks (Eremalauda dunni) from the Arabian Desert and skylarks (Alauda arvensis) and woodlarks (Lullula arborea) from temperate mesic grasslands. The proportional contribution of CWL to TEWL in all larks at moderate T(a) ranged from 50% to 70%. At high T(a) (40 degrees -45 degrees C), larks enhanced CWL by only 45%-78% and relied on an increase in RWL by 676%-2,733% for evaporative cooling. Surface-specific CWL at 25 degrees C was 29% lower in the arid-zone species than in the mesic larks. When acclimated to constant T(a), 15 degrees C-acclimated hoopoe larks increased CWL by 22% compared with 35 degrees C-acclimated birds, but the other species did not change CWL. This study is consistent with the hypothesis that larks from deserts have a reduced CWL at moderate and low T(a) but provided no support for the hypothesis that at high T(a) larks from arid regions rely more on CWL than larks from mesic environments. Interspecific differences in CWL cannot be attributed to acclimation to environmental temperature and are possibly the result of genetic differences due to natural selection or of phenotypically plastic responses to divergent environments during ontogeny.     

Energy metabolism and evaporative water loss in the European free-tailed bat and Hemprich's long-eared bat (Microchiroptera): species sympatric in the Negev Desert

We compared the thermoregulatory abilities of two insectivorous bat species, Tadarida teniotis (mean body mass 32 g) and Otonycteris hemprichii (mean body mass 25 g), that are of different phylogenetic origins and zoogeographic distributions but are sympatric in the Negev Desert. At night, both were normothermic. By day, both were torpid when exposed to ambient temperatures (T(a)) below 25 degrees Celsius, with concomitant adjustments in metabolic rate (MR). Otonycteris hemprichii entered torpor at higher T(a) than T. teniotis, and, when torpid, their body temperatures (T(b)) were 1 degrees -2 degrees Celsius and 5 degrees -8 degrees Celsius above T(a), respectively; MR was correspondingly reduced. At night, the lower critical temperature of T. teniotis was 31.5 degrees Celsius, and that of O. hemprichii was 33 degrees Celsius. Mean nocturnal thermoneutral MR of T. teniotis was 37% greater than that of O. hemprichii. At high T(a), evaporative water loss (EWL) increased markedly in both species, but it was significantly higher in T. teniotis above 38 degrees Celsius. In both species, the dry heat transfer coefficient (thermal conductance) followed the expected pattern for small mammals, by day and by night. Total EWL was notably low in normothermic and torpid animals of both species, much lower than values reported for other bats, indicating efficient water conservation mechanisms in the study species. Comparing thermoregulatory abilities suggests that O. hemprichii is better adapted to hot, arid environments than T. teniotis, which may explain its wider desert distribution. By both standard and phylogenetically informed ANCOVA, we found no differences in basal metabolic rate (BMR) between desert and nondesert species of insectivorous bats, substantiating previous studies suggesting that low BMR is a characteristic common to insectivorous bats in general.     

Bias, precision and accuracy in the estimation of cuticular and respiratory water loss: a case study from a highly variable cockroach, Perisphaeria sp

We compared the precision, bias and accuracy of two techniques that were recently proposed to estimate the contributions of cuticular and respiratory water loss to total water loss in insects. We performed measurements of VCO2 and VH2O in normoxia, hyperoxia and anoxia using flow through respirometry on single individuals of the highly variable cockroach Perisphaeria sp. to compare estimates of cuticular and respiratory water loss (CWL and RWL) obtained by the VH2O-VCO2 y-intercept method with those obtained by the hyperoxic switch method. Precision was determined by assessing the repeatability of values obtained whereas bias was assessed by comparing the methods' results to each other and to values for other species found in the literature. We found that CWL was highly repeatable by both methods (R0.88) and resulted in similar values to measures of CWL determined during the closed-phase of discontinuous gas exchange (DGE). Repeatability of RWL was much lower (R=0.40) and significant only in the case of the hyperoxic method. RWL derived from the hyperoxic method is higher (by 0.044 micromol min(-1)) than that obtained from the method traditionally used for measuring water loss during the closed-phase of DGE, suggesting that in the past RWL may have been underestimated. The very low cuticular permeability of this species (3.88 microg cm(-2) h(-1) Torr(-1)) is reasonable given the seasonally hot and dry habitat where it lives. We also tested the hygric hypothesis proposed to account for the evolution of discontinuous gas exchange cycles and found no effect of respiratory pattern on RWL, although the ratio of mean VH2O to VCO2 was higher for continuous patterns compared with discontinuous ones.     

Cocoon and epidermis of Australian Cyclorana frogs differ in composition of lipid classes that affect water loss

For amphibians to survive in environments that experience annual droughts, they must minimize evaporative water loss. One genus of Australian hylid frogs, Cyclorana, prevents desiccation by burrowing in the soil and forming cocoons composed of alternating layers of shed epidermis and glandular secretions. Previous data are inconclusive about the role that lipids play in reducing evaporative water loss through skin (cutaneous water loss [CWL]) when Cyclorana spp. are within cocoons. In this study, we measured CWL and lipids in the epidermis and in cocoons of five species of Cyclorana. CWL was significantly lower in frogs within cocoons than in frogs without cocoons. Surface-area-specific CWL for the three small species was significantly higher than that of the two larger species of Cyclorana, but this difference was not apparent in frogs within cocoons. Although lipids were responsible for more of the dry mass of the epidermis (approximately 20%) than of the cocoons (approximately 7%) we found that cerebrosides and ceramides, two polar lipid classes, were almost exclusively found in cocoons. This suggests that these lipid classes are in the glandular secretions rather than in the epidermis. Because these polar lipids are the types that reduce water loss in birds (cerebrosides and ceramides) and mammals (ceramides), we conclude that they are important not only for holding together the shed layers of skin but also for contributing to the barrier against water loss.     

The role of polyunsaturated fatty acids in the expression of torpor by mammals: a review

Heterothermic mammals increase the proportion of polyunsaturated fatty acids (PUFA) in their body fats prior to entering torpor. Because PUFA have low melting points, it is thought that they play an important role in maintaining the fluidity of depot fats and membrane phospholipids at low body temperatures. However, PUFA are more prone to autoxidation when exposed to reactive oxygen species (ROS) during torpor and during the periodic arousals that characterize hibernation. A lack of PUFA or an excess of PUFA may constrain the use of torpor by heterothermic mammals. We performed a mixed model meta-analysis of 17 controlled-feeding studies to test the effect of dietary PUFA on the depth and expression of torpor by daily heterotherms and hibernators. We also reviewed the literature on the PUFA content of the diet and depot fats of heterothermic mammals to address two principal topics: (1) Do low dietary levels of PUFA reduce the expression of torpor under laboratory conditions and, if so, are free-ranging animals constrained by a lack of PUFA? (2) Do high dietary levels of PUFA result in a reduction in the use, depth, and duration of torpor and, if so, do free-ranging animals seek to optimize rather than maximize PUFA intake? Low-PUFA diets consistently increase the lower setpoint for body temperature and minimum metabolic rate for both hibernators and daily heterotherms. Above the lower setpoint, low-PUFA diets usually increase body temperature and metabolic rate and decrease the duration of torpor bouts and this effect is similar for hibernators and daily heterotherms. Free-ranging rodent hibernators have dietary PUFA intakes that are far higher than those of the low-PUFA diets offered in controlled-feeding experiments, so these hibernators may never experience the constraints associated with a lack of PUFA. Diets of free-ranging insectivorous bats and echidnas have PUFA levels that are less than half as high as those offered in experimental low-PUFA diets, yet they exhibit deep and extended bouts of torpor. We argue that alternate mechanisms exist for maintaining the fluidity of body fats and that high-PUFA intake may not be a prerequisite for deep and extended bouts of torpor. Four studies indicate that animals that were fed high-PUFA diets are reluctant to enter torpor and show shallower and shorter torpor bouts. Although authors attribute this response to autoxidation, these animals did not have a higher PUFA content in their depot fats than animals where PUFA was shown to enhance torpor. We suggest that these contradictory results indicate inter-specific or inter-individual variation in the ability to control ROS and limit autoxidation of PUFA. High dietary levels of PUFA will constrain the expression of torpor only when the oxidative challenge exceeds the capacity of the antioxidant defence system. Studies of diet selection indicate that insectivorous species with low dietary PUFA levels seek to maximize PUFA intake. However, herbivorous species that have access to plants and plant parts of high-PUFA content do not appear to maximize PUFA intake. These data suggest that animals attempt to optimize rather than maximize PUFA intake. The effect of PUFA should be viewed in the light of a cost-benefit trade-off, where the benefit of high-PUFA intake is an easier access to low body temperatures and the cost is increased risk of autoxidation.     

Cutaneous water loss and lipids of the stratum corneum in two syntopic species of bats

The lipid matrix of the stratum corneum (SC), the outer layer of the epidermis of mammals and birds, constitutes the barrier to diffusion of water vapor through the skin. The lipids of the SC are structured in the intercellular spaces of the mammalian epidermis in ordered layers, called lamellae, which have been postulated to prevent water loss. Lipids in the mammalian SC are mainly cholesterol, free fatty acids and ceramides, the latter forming the structural support for the lamellae. However, knowledge on how the lipid composition of the SC alters cutaneous water loss (CWL) in mammals is rudimentary, and is largely derived from studies on laboratory animals and humans. We measured CWL of individuals of two species of syntopic bats, Tadarida brasiliensis and Myotis velifer. In the first study of its kind on wild mammals, we correlated CWL with the lipid composition of the SC, measured using thin layer chromatography and high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. Surface-specific CWL was 20.6% higher in M. velifer than in T. brasiliensis, although differences were not significant. Compared with individuals of M. velifer, individuals of T. brasiliensis had more classes, and a higher proportion, of polar ceramides in the SC, a feature associated with lower CWL. Individuals of T. brasiliensis also had a class of non-polar ceramides that presumably spans the lamellae and gives more cohesiveness to the lipid matrix of the SC. We conclude that qualitative and quantitative modifications of the lipid composition of the SC contribute to regulate CWL of these two species of bats.     

Hung out to dry? Intraspecific variation in water loss in a hibernating bat

Hibernation is a period of water deficit for some small mammals, and humidity strongly influences hibernation patterns. Dry conditions reduce length of torpor bouts, stimulate arousals, and decrease overwinter survival. To mitigate these effects, many small mammals hibernate in near saturated (100% RH) conditions. However, big brown bats (Eptesicus fuscus) hibernate in a wider variety of conditions and tolerate lower humidity than most other bats. To assess arid tolerance in this species, we compared torpid metabolic rates (TMR) and rates of total evaporative water loss (TEWL) between two populations of E. fuscus with differing winter ecologies: one that hibernates in humid karst caves and one that hibernates in relatively dry rock crevices. We used flow-through respirometry to measure TMR and TEWL of bats in humid and dry conditions. Torpid metabolic rates did not differ between populations or with humidity treatments. Rates of TEWL were similar between populations in humid conditions, but higher for cave-hibernating bats than crevice-hibernating bats in dry conditions. Our results suggest that E. fuscus hibernating in arid environments have mechanisms to decrease evaporative water loss that are not evident at more humid sites. Drought tolerance may facilitate the sedentary nature of the species, allowing them to tolerate more variable microclimates during hibernation and thus increasing the availability of overwintering habitat. The ability to survive arid conditions may also lessen the susceptibility of E. fuscus to diseases that affect water balance.     

Evaporative water loss in two sympatric species of vespertilionid bat, Plecotus auritus and Myotis daubentoni: relation to foraging mode and implications for roost site selection

Simultaneous measures of oxygen consumption and evaporative water loss (EWL) were made in two species of temperate-zone vespertilionid bat ( Plecotus auritus and Myotis daubentoni ; mean body mass 9.12 and 10.12g, respectively) at ambient temperatures (T_a) of 5, 15 and 25°C and variable vapour pressure deficit. EWL was directly dependent on vapour pressure deficit and oxygen consumption and inversely dependent on T_a. EWL was significantly greater in P. auritus than in M. daubentoni. A model for EWL in P. auritus under a variety of environmental conditions (5–25°C and 20–80% relative humidity) suggested that EWL from bats in shallow summer torpor will be lowest at low T_a, and that, except at low (> 50%) relative humidity, EWL from euthermic bats will be lowest at high T_a. At low relative humidity (< 20%), resting bats could lose over 30% of body mass per day (24 h) through evaporation. At high T_a (> 25°C), EWL from euthermic bats could be over 65% lower at high (> 80%) compared to low (< 20%) relative humidity. In bats in shallow summer torpor at low (5°C) T_a the equivalent saving was > 96%. At low relative humidity predicted EWL from bats in shallow summer torpor was 34 to 81% of that from euthermic bats, and at low T_a and high relative humidity was only 2%. In the wild, M. daubentoni has freer access to drinking water than does P. auritus and yet EWL at rest was higher in the latter species. We suggest that post-prandial dumping of urinary water by M. daubentoni leads to a limit in the amount of body water available to this species to cover evaporative losses once within the day roost, which in turn has led to an adaptation of physiology towards the minimization of EWL when at rest.     

Hibernation and torpor in tropical and subtropical bats in relation to energetics, extinctions, and the evolution of endothermy

Torpor, the most effective means of energy conservation available to endotherms, is still widely viewed as a specific adaptation in a few high-latitude, cold-climate endotherms with no adaptive function in warm regions. Nevertheless, a growing number of diverse terrestrial mammals and birds from low latitudes (0-30°), including species from tropical and subtropical regions, are heterothermic and employ torpor. Use of torpor is especially important for bats because they are small, expend large amounts of energy when active, rely on a fluctuating food supply, and have only a limited capacity for storage of fat. Patterns of torpor in tropical/subtropical bats are highly variable, but short bouts of torpor with relatively high body temperatures (T(b)) are most common. Hibernation (a sequence of multiday bouts of torpor) has been reported for free-ranging subtropical tree-dwelling vespertilionids, cave-dwelling hipposiderids, and house-dwelling molossids. The observed range of minimum T(b) is ～6-30 °C, and the reduction of energy expenditure through the use of torpor, in comparison to normothermic values, ranges from 50 to 99%. Overall, torpor in the tropics/subtropics has been reported for 10 out of the currently recognized 18 bat families, which contain 1079 species, or 96.7% of all bats. Although it is unlikely that all of these are heterothermic, the large majority probably will be. Frequent use of torpor, including hibernation in diverse groups of tropical/subtropical bats, suggests that heterothermy is an ancestral chiropteran trait. Although data especially from the field are still scarce, it is likely that torpor, highly effective in reducing requirements for energy and water even under warm conditions, plays a crucial role in the long-term survival of the majority of small tropical and subtropical bats. Discovering how bats achieve this provides numerous opportunities for exiting new research.     

Tropical bats counter heat by combining torpor with adaptive hyperthermia

Many tropical mammals are vulnerable to heat because their water budget limits the use of evaporative cooling for heat compensation. Further increasing temperatures and aridity might consequently exceed their thermoregulatory capacities. Here, we describe two novel modes of torpor, a response usually associated with cold or resource bottlenecks, as efficient mechanisms to counter heat. We conducted a field study on the Malagasy bat Macronycteris commersoni resting in foliage during the hot season, unprotected from environmental extremes. On warm days, the bats alternated between remarkably short micro-torpor bouts and normal resting metabolism within a few minutes. On hot days, the bats extended their torpor bouts over the hottest time of the day while tolerating body temperatures up to 42.9°C. Adaptive hyperthermia combined with lowered metabolic heat production from torpor allows higher heat storage from the environment, negates the need for evaporative cooling and thus increases heat tolerance. However, it is a high-risk response as the torpid bats cannot defend body temperature if ambient temperature increases above a critical/lethal threshold. Torpor coupled with hyperthermia and micro-torpor bouts broaden our understanding of the basic principles of thermal physiology and demonstrate how mammals can perform near their upper thermal limits in an increasingly warmer world.     

Subtropical mouse-tailed bats use geothermally heated caves for winter hibernation

We report that two species of mouse-tailed bats (Rhinopoma microphyllum and R. cystops) hibernate for five months during winter in geothermally heated caves with stable high temperature (20°C). While hibernating, these bats do not feed or drink, even on warm nights when other bat species are active. We used thermo-sensitive transmitters to measure the bats’ skin temperature in the natural hibernacula and open flow respirometry to measure torpid metabolic rate at different ambient temperatures (T_a, 16–35°C) and evaporative water loss (EWL) in the laboratory. Bats average skin temperature at the natural hibernacula was 21.7 ± 0.8°C, and no arousals were recorded. Both species reached the lowest metabolic rates around natural hibernacula temperatures (20°C, average of 0.14 ± 0.01 and 0.16 ± 0.04 ml O₂ g⁻¹ h⁻¹ for R. microphyllum and R. cystops, respectively) and aroused from torpor when T_a fell below 16°C. During torpor the bats performed long apnoeas (14 ± 1.6 and 16 ± 1.5 min, respectively) and had a very low EWL. We hypothesize that the particular diet of these bats is an adaptation to hibernation at high temperatures and that caves featuring high temperature and humidity during winter enable these species to survive this season on the northern edge of their world distribution.     

Environmental conditions, rather than season, determine torpor use and temperature selection in large mouse-eared bats (Myotis myotis)

We tested whether food availability, thermal environment and time of year affect torpor use and temperature selection in the large mouse-eared bat (Myotis myotis) in summer and winter. Food-deprived bats were torpid longer than bats offered food ad libitum. Bats placed in a gradient of low (0 degrees C-25 degrees C) ambient temperatures (T(a)) spent more time in torpor than bats in a gradient of high (7 degrees C-43 degrees C) T(a)'s. However, we did not observe seasonal variations in the use of torpor. Moreover, even when food deprived in winter, bats never entered prolonged torpor at T(a)'s characteristic of their natural hibernation. Instead, bats preferred shallow torpor at relatively high T(a), but they always maintained a difference between body and ambient temperatures of less than 2 degrees C. Calculations based on respirometric measurements of metabolic rate showed that food deprived bats spent less energy per unit of time in torpor than fed individuals, even when they entered torpor at higher T(a)'s. We conclude that T(a) likely serves as a signal of food availability and daily torpor is apparently an adaptation to unpredictable changes in food availability, such as its decrease in summer or its increase in winter. Thus, we interpret hibernation to be a second step in the evolution of heterothermy in bats, which allows survival in seasonal environments.     

Evaporative water loss is a plausible explanation for mortality of bats from white-nose syndrome

White-nose syndrome (WNS) has caused alarming declines of North American bat populations in the 5 years since its discovery. Affected bats appear to starve during hibernation, possibly because of disruption of normal cycles of torpor and arousal. The importance of hydration state and evaporative water loss (EWL) for influencing the duration of torpor bouts in hibernating mammals recently led to "the dehydration hypothesis," that cutaneous infection of the wing membranes of bats with the fungus Geomyces destructans causes dehydration which in turn, increases arousal frequency during hibernation. This hypothesis predicts that uninfected individuals of species most susceptible to WNS, like little brown bats (Myotis lucifugus), exhibit high rates of EWL compared to less susceptible species. We tested the feasibility of this prediction using data from the literature and new data quantifying EWL in Natterer's bats (Myotis nattereri), a species that is, like other European bats, sympatric with G. destructans but does not appear to suffer significant mortality from WNS. We found that little brown bats exhibited significantly higher rates of normothermic EWL than did other bat species for which comparable EWL data are available. We also found that Natterer's bats exhibited significantly lower rates of EWL, in both wet and dry air, compared with values predicted for little brown bats exposed to identical relative humidity (RH). We used a population model to show that the increase in EWL required to cause the pattern of mortality observed for WNS-affected little brown bats was small, equivalent to a solitary bat hibernating exposed to RH of ～95%, or clusters hibernating in ～87% RH, as opposed to typical near-saturation conditions. Both of these results suggest the dehydration hypothesis is plausible and worth pursuing as a possible explanation for mortality of bats from WNS.     

Standard energetics of leaf-nosed bats (Hipposideridae): its relationship to intermittent- and protracted-foraging tactics in bats and birds

Basal rates of metabolism within the insectivorous genera Hipposideros and Ascelliscus, Old World leaf-nosed bats (Hipposideridae), ranged from 58% to 77% of the mammalian standard. The larger species, Hipposideros diadema and Hipposideros maggietaylori, effectively thermoregulated at ambient temperatures down to 9 degrees C, whereas two smaller species, Hipposideros galeritus and Hipposideros cervinus, occasionally permitted body temperatures to fall below 32 degrees C. The low basal rates of metabolism in hipposiderids correlated with a predatory life-style characterized by intermittent flight from a perch to capture insects, a correlation similar to that found in nonpasserine birds. Intermittent-foraging bats and nonpasserines collectively had basal rates of metabolism that averaged 75% of those that pursue insects during protracted flight. However, no difference in basal rate was found between protracted- and intermittent-foraging passerines, which had basal rates 1.8- and 2.4-times those of protracted-foraging and intermittent-foraging bats and nonpasserines, respectively. Bats, swifts, and caprimulgids that enter torpor have basal rates that are 85% of those of similar species that do not enter torpor. Body mass, order affiliation, foraging mode, and propensity to enter into torpor collectively account for 97% of the variation in basal rate of metabolism in insectivorous bats and birds. Foraging style therefore appears to be a factor contributing to the diversity in endotherm energetics. Minimal thermal conductance in the genus Hipposideros ranged from 75% to 102% of the mammalian standard. Birds have minimal thermal conductances that are 75% of mammals and intermittent foragers have minimal conductances that are 78% of protracted foragers.     

Torpor in free-ranging tawny frogmouths (Podargus strigoides).

Several small caprimulgiform birds (<80 g) are known to enter torpor, apparently to cope with a fluctuating supply of insect prey. Since the large Australian tawny frogmouth (Podargus strigoides; 381-556 g) is also insectivorous, we investigated its thermoregulatory behaviour and thermal biology to determine whether this species is also heterothermic. In an open woodland at approximately 1,000 m altitude, we equipped eight free-ranging birds with external temperature-sensitive radio transmitters attached to an elastic harness to measure skin temperature (T(skin)). Core body temperature (T(b)) was measured in three of these birds fitted with an additional intraperitoneal transmitter. T(skin) was closely correlated with T(b), although T(skin) was usually several degrees below T(b). During the three coldest months of the year (June-August), shallow torpor with T(b) as low as 29.1 degrees C occurred frequently, whereas during spring and summer, torpor was not recorded. Torpor occurred either during the night and/or during the first half of the day. Night torpor bouts were initiated after a short activity period around dusk and lasted on average for about 7 h. Torpid birds always aroused before sunrise to either commence a second short foraging period or to fly directly to a day roost tree. After birds roosted, T(b) fell again around sunrise, and birds occasionally entered a second dawn torpor bout; however, in most cases, T(b) increased rapidly not long after entry, most likely due to passive heating by the sun. We conclude that despite their large body size and energetically conservative hunting strategy, tawny frogmouths, like several related caprimulgiform species, frequently enter shallow torpor when low T(a) demands high energetic costs for normothermic thermoregulation and likely reduces insect availability.     

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.     

Do season and distribution affect thermal energetics of a hibernating bat endemic to the tropics and subtropics?

Although many tropical and subtropical areas experience pronounced seasonal changes in weather and food availability, few studies have examined and none have compared the thermal physiology and energetics of a hibernating mammal that is restricted to these regions. We quantified thermal energetics of northern long-eared bats (Nyctophilus bifax; body mass ～10 g) during summer, winter, and spring from a subtropical habitat, and also during winter from a tropical habitat, to determine how N. bifax cope with climate and seasonal changes in weather. We captured bats in the wild and measured metabolic rates via open-flow respirometry. The basal metabolic rate of subtropical bats at an ambient temperature (T(a)) of 32.6 ± 0.7°C was 1.28 ± 0.06 ml O(2)·g(-1)·h(-1) during both summer and winter, similar to other species of Nyctophilus. Resting metabolic rates below the thermoneutral zone increased similarly with decreasing T(a) during all seasons and in both regions. All individuals showed a high proclivity to enter torpor at T(a) values below the thermoneutral zone. Metabolic rates in torpid thermoconforming bats fell with T(a) and body temperature, and mean minimum metabolic rates during torpor were similar during all seasons and in both regions and as predicted from body mass in temperate zone hibernators. At very low T(a), torpid N. bifax thermoregulated, and this threshold T(a) differed significantly between subtropical (T(a) = 3.5 ± 0.3°C) and tropical (T(a) = 6.7 ± 0.7°C) individuals, but not between seasons. Our data show that thermal energetics of N. bifax do not vary seasonally and in many aspects are similar in tropical and subtropical bats; however, torpid individuals from the subtropics allow body temperature to fall to significantly lower values than those from the tropics.     

Frequent Arousals from Winter Torpor in Rafinesque’s Big-Eared Bat (Corynorhinus rafinesquii)

Extensive use of torpor is a common winter survival strategy among bats; however, data comparing various torpor behaviors among species are scarce. Winter torpor behaviors are likely to vary among species with different physiologies and species inhabiting different regional climates. Understanding these differences may be important in identifying differing susceptibilities of species to white-nose syndrome (WNS) in North America. We fitted 24 Rafinesque’s big-eared bats (Corynorhinus rafinesquii) with temperature-sensitive radio-transmitters, and monitored 128 PIT-tagged big-eared bats, during the winter months of 2010 to 2012. We tested the hypothesis that Rafinesque’s big-eared bats use torpor less often than values reported for other North American cave-hibernators. Additionally, we tested the hypothesis that Rafinesque’s big-eared bats arouse on winter nights more suitable for nocturnal foraging. Radio-tagged bats used short (2.4 d ± 0.3 (SE)), shallow (13.9°C ± 0.6) torpor bouts and switched roosts every 4.1 d ± 0.6. Probability of arousal from torpor increased linearly with ambient temperature at sunset (P<0.0001), and 83% (n = 86) of arousals occurred within 1 hr of sunset. Activity of PIT-tagged bats at an artificial maternity/hibernaculum roost between November and March was positively correlated with ambient temperature at sunset (P<0.0001), with males more active at the roost than females. These data show Rafinesque’s big-eared bat is a shallow hibernator and is relatively active during winter. We hypothesize that winter activity patterns provide Corynorhinus species with an ecological and physiological defense against the fungus causing WNS, and that these bats may be better suited to withstand fungal infection than other cave-hibernating bat species in eastern North America.     

Increased heat loss affects hibernation in golden-mantled ground squirrels

During hibernation at ambient temperatures (T(a)) above 0 degrees C, rodents typically maintain body temperature (T(b)) approximately 1 degrees C above T(a), reduce metabolic rate, and suspend or substantially reduce many physiological functions. We tested the extent to which the presence of an insulative pelage affects hibernation. T(b) was recorded telemetrically in golden-mantled ground squirrels (Spermophilus lateralis) housed at a T(a) of 5 degrees C; food intake and body mass were measured at regular intervals throughout the hibernation season and after the terminal arousal. Animals were subjected to complete removal of the dorsal fur or a control procedure after they had been in hibernation for 3-4 wk. Shaved squirrels continued to hibernate with little or no change in minimum T(b), bout duration, duration of periodic normothermic bouts, and food intake during normothermia. Rates of rewarming from torpor were, however, significantly slower in shaved squirrels, and rates of body mass loss were significantly higher, indicating increased depletion of white adipose energy stores. An insulative pelage evidently conserves energy over the course of the hibernation season by decreasing body heat loss and reducing energy expenditure during periodic arousals from torpor and subsequent intervals of normothermia. This prolongs the hibernation season by several weeks, thereby eliminating the debilitating consequences associated with premature emergence from hibernation.     

Thermoregulation and roost selection by reproductive female big brown bats (Eptesicus fuscus) roosting in rock crevices

A free-ranging maternity colony of big brown bats Eptesicus fuscus roosting in rock crevices along the South Saskatchewan River in south-eastern Alberta, Canada, was studied to understand better the discrepancy that exists in the literature regarding torpor use by reproductive female bats. Using radio-telemetry, thermoregulatory patterns and roost microclimate were recorded for pregnant, lactating and post-lactating females. Relative torpor use is described in several ways: the proportion of days on which torpor was used, depth, minimum body temperature, time spent in torpor, and a comprehensive torpor unit (degree-min). Pregnant and lactating female E. fuscus used torpor to the same extent overall (degree-min), but pregnant bats used torpor less frequently and with more time in deep torpor. Torpor was used to the greatest extent after weaning (post-lactation). Evidence is presented that the cost:benefit ratio for deep and prolonged periods of torpor may be highest during lactation. Microclimates of rock-crevice roosts mirrored the use of torpor throughout reproduction by bats. Lactation roosts (deeper, larger opening size) were more thermally stable and remained warmer at night compared to the shallow roosts used by pregnant and post-lactating females. It is shown that conclusions about relative use of torpor can differ depending on the units of comparison, necessitating measurement of all aspects of torpor (depth, duration and frequency). Comprehensive measurements, individual-based normothermic temperatures, and a definition of torpor that accounts for all energy savings, allow a more accurate depiction of patterns and facilitates inter-study comparisons.