Seasonal changes in daily torpor patterns of free-ranging female and male Daubenton’s bats (Myotis daubentonii)

Daily torpor can provide significant energy and water savings in bats during cold ambient temperatures and food scarcity. However, it may reduce rates of foetal and juvenile development. Therefore, reproductive females should optimize development by minimizing times in torpor. To test this hypothesis, the use of torpor by female and male free-ranging Daubenton’s bats (Myotis daubentonii) during reproduction (gestation, lactation, and post-lactation period) was investigated in 1998 and 1999. Temperature-sensitive radio transmitters were attached to the bats to measure skin temperature. Simultaneously, ambient temperature was recorded. While both sexes became torpid during daytime, male bats used daily torpor (>6°C below individual active temperature) significantly more often during reproductive period (mean: 78.4 % of day time in May and 43 % in June) than females. Female bats went into daily torpor, particularly in late summer when juveniles were weaned (mean: 66.6 % of daytime). Lowest skin temperatures occurred in a female bat with 21.0°C during post-lactation. Skin temperatures of male bats fluctuated from 16.8°C in torpor to 37.2°C during times of activity. There was a significant effect of reproductive period on skin temperature in females whereas mean ambient temperature had no significant effect. However, mean ambient temperature affected mean skin temperatures in males. Our findings indicate that female Daubenton’s bats adopt their thermoregulatory behaviour in particular to optimize the juvenile development.     

The use of small subcutaneous transponders for quantifying thermal biology and torpor in small mammals

Remote measurements of body temperature (T_b) in animals require implantation of relatively large temperature-sensitive radio-transmitters or data loggers, whereas rectal temperature (T_rec) measurements require handling and therefore may bias the results. We investigated whether ∼0.1 g temperature-sensitive subcutaneously implanted transponders can be reliably used to quantify thermal biology and torpor use in small mammals. We examined (i) the precision of transponder readings as a function of temperature and (ii) whether subcutaneous transponders can be used to remotely record subcutaneous temperature (T_sub). Five adult male dunnarts (Sminthopsis macroura, body mass 24 g) were implanted with subcutaneous transponders to determine T_sub as a function of time and ambient temperature (T_a), and in comparison to thermocouple readings of T_rec. Transponder temperature was highly correlated with water bath temperature (r²=0.96–0.99) over a range of approximately 10.0–40.0 °C. Transponders provided reliable data (±0.6 °C) over the T_sub of 21.4–36.9 °C and could be read from a distance of up to 5 cm. Below 21.4 °C, accuracy was reduced to ±2.8 °C, but individual transponder accuracy varied. Consequently, small subcutaneous transponders are useful to remotely quantify thermal physiology and torpor patterns without having to disturb the animal and disrupt torpor. Even at T_sub<21.4 °C where the accuracy of the temperature readings was reduced, transponders do provide reliable data on whether and when torpor is used.     

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.     

Effects of reproductive condition,roost microclimate,and weather patterns on summer torpor use by a vespertilionid bat

A growing number of mammal species are recognized as heterothermic, capable of maintaining a high‐core body temperature or entering a state of metabolic suppression known as torpor. Small mammals can achieve large energetic savings when torpid, but they are also subject to ecological costs. Studying torpor use in an ecological and physiological context can help elucidate relative costs and benefits of torpor to different groups within a population. We measured skin temperatures of 46 adult Rafinesque's big‐eared bats (Corynorhinus rafinesquii) to evaluate thermoregulatory strategies of a heterothermic small mammal during the reproductive season. We compared daily average and minimum skin temperatures as well as the frequency, duration, and depth of torpor bouts of sex and reproductive classes of bats inhabiting day‐roosts with different thermal characteristics. We evaluated roosts with microclimates colder (caves) and warmer (buildings) than ambient air temperatures, as well as roosts with intermediate conditions (trees and rock crevices). Using Akaike's information criterion (AIC), we found that different statistical models best predicted various characteristics of torpor bouts. While the type of day‐roost best predicted the average number of torpor bouts that bats used each day, current weather variables best predicted daily average and minimum skin temperatures of bats, and reproductive condition best predicted average torpor bout depth and the average amount of time spent torpid each day by bats. Finding that different models best explain varying aspects of heterothermy illustrates the importance of torpor to both reproductive and nonreproductive small mammals and emphasizes the multifaceted nature of heterothermy and the need to collect data on numerous heterothermic response variables within an ecophysiological context.     

Effect of low-quality diet on torpor frequency and depth in the pichi Zaedyus pichiy (Xenarthra,Dasypodidae), a South American armadillo

Daily torpor is a physiological adaptation that allows mammals to cope with energetic challenges associated with unpredictable periods of food shortage. We experimentally tested whether food quality influences torpor frequency and depth in the pichi (Zaedyus pichiy), a small, opportunistically omnivorous armadillo endemic to arid and semi-arid habitats of southern South America. We recorded body temperature (T_sc) changes in 10 semi-captive, adult female pichis using dataloggers implanted subcutaneously during periods of 21 days. All individuals entered spontaneous daily torpor, but those receiving a low-quality diet had significantly lower daily mean and minimum T_sc, spent more time at T_sc below their individual lower limit of normothermia, and had a higher Heterothermy Index than controls. Five individuals entered prolonged torpor bouts lasting more than 24 h, two of them repeatedly. Nine out of ten prolonged torpor bouts occurred in individuals feeding on a low-quality diet, suggesting that pichis are able to enter prolonged periods of torpor during severe environmental stress. In combination with their ability to hibernate and to respond to a reduced insect abundance by ingesting other food items, this physiological adaptation allows pichis to better cope with food shortages and a more extreme climate than other armadillos. It may explain why Z. pichiy naturally occurs farther south than any other armadillo species.     

Summer heterothermy in Rafinesque’s big-eared bats (Corynorhinus rafinesquii) roosting in tree cavities in bottomland hardwood forests

Many small mammals are heterothermic endotherms capable of maintaining an elevated core body temperature or reducing their thermoregulatory set point to enter a state of torpor. Torpor can confer substantial energy savings, but also incurs ecological costs, such as hindering allocation of energy towards reproduction. We placed temperature-sensitive radio transmitters on 44 adult Rafinesque’s big-eared bats (Corynorhinus rafinesquii) and deployed microclimate dataloggers inside 34 day roosts to compare the use of torpor by different sex and reproductive classes of bats during the summer. We collected 324 bat-days of skin-temperature data from 36 females and 4 males. Reproductive females employed fewer torpor bouts per day than non-reproductive females and males (P < 0.0001), and pregnant and lactating females had higher average (P < 0.0001) and minimum (P < 0.0001) skin temperatures than non-reproductive females. Pregnant females spent less time torpid (P < 0.0001) than non-reproductive females, but lactating females used relatively deep, long torpor bouts. Microclimates varied inside tree species with different configurations of entrances to the roost cavity (P < 0.0001). Bats spent more time torpid when roosting in water tupelo (Nyssa aquatica) trees possessing only a basal entrance to the cavity (P = 0.001). Of the tree species used as roosts, water tupelo cavities exhibited the least variable daytime and nighttime temperatures. These data demonstrate that use of summer torpor is not uniform among sex and reproductive classes in Rafinesque’s big-eared bat, and variation in microclimate among tree roosts due to species and structural characteristics facilitates the use of different thermoregulatory strategies in these bats.     

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.     

Roost characteristics as indicators for heterothermic behavior of forest-dwelling bats

Many forest-dwelling bats spend their diurnal inactivity period in tree cavities. During this time bats can save energy through heterothermy. A heterothermic response (torpor) is characterized by a lowered body temperature, reduced metabolic rate, and reduction of other physiological processes, and can be influenced by the microclimatic conditions of roost cavities. The thermal and physical characteristics of roosts used by the sympatric, ecologically, and morphologically similar bat species Myotis bechsteinii, M. nattereri, and Plecotus auritus were compared. These three species differ in their heterothermic behavior, with the lowest skin temperatures observed for P. auritus. Therefore, we hypothesized that roosts occupied by the three species should differ in roost characteristics and microclimatic conditions, whereby P. auritus should select colder and thermally less stable roosts. The results showed that horizontal depth of the cavity, diameter of the roost tree, and microclimatic conditions within roosts differed among species. Roosts of P. auritus had the lowest horizontal depth, lowest thermal stability, and lowest mean minimum roost temperatures. Height of the roost, diameter of the roost tree, and vertical depth were also shown to influence microclimatic conditions. With increasing diameter of the tree and increasing horizontal depth, mean minimum roost temperature increased and thermal stability improved. Furthermore, with ascending height above ground insulation and mean roost temperatures increased. Our results imply that species such as P. auritus, which use pronounced torpor as a primary energy saving strategy, prefer colder cavities that support their heterothermic strategy.     

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.     

Hibernation by a free-ranging subtropical bat (Nyctophilus bifax)

Knowledge about torpor in free-ranging subtropical bats is scarce and it is widely believed that low and stable ambient temperatures are necessary for prolonged torpor. We present temperature-telemetry data from free-ranging male (n = 4) and female (n = 4) subtropical vespertilionid bats, Nyctophilus bifax (~10 g), exposed to pronounced daily fluctuations of ambient temperature. All bats used torpor on every day in winter and both males and females exhibited multi-day torpor bouts of up to 5.4 days. Although females were larger than males, patterns of torpor were similar in both sexes. Torpor use was correlated with prevailing weather conditions and, on days when bats remained torpid, maximum ambient temperature was significantly lower than on days when bats aroused. Moreover, the duration of interbout normothermic periods at night increased with increasing average nightly ambient temperature. Skin temperature of torpid bats varied by 10.2 ± 3.6°C day⁻¹ (n = 8, N = 47) and daily minimum skin temperature was positively correlated with the daily minimum ambient temperature. Our study shows that prolonged torpor is an important component of the winter ecology of a subtropical bat and that torpor and activity patterns of N. bifax predominantly reflect prevailing weather conditions.     

Roost use by long-tailed bats in South Canterbury: examining predictions of roost-site selection in a highly fragmented landscape

We studied the roosting ecology of the long-tailed bat (Chalinolobus tuberculatus) during the springautumn months from 1998–2002 at Hanging Rock in the highly fragmented landscape of South Canterbury, South Island, New Zealand. We compared the structural characteristics and microclimates of roost sites used by communally and solitary roosting bats with those of randomly available sites, and roosts of C. tuberculatus occupying unmodified Nothofagus forest in the Eglinton Valley, Fiordland. Roosting group sizes and roost residency times were also compared. We followed forty radio-tagged bats to 94 roosts (20% in limestone crevices, 80% in trees) at Hanging Rock. Roosts were occupied for an average of 1 day and 86% were only used once during the study period. Colony size averaged 9.8 ± 1.1 bats (range 2–38) and colonies were dominated by breeding females and young. Indigenous forest, shrubland remnants and riparian zones were preferred roosting habitats. Communally roosting bats selected roosts in split trunks of some of the largest trees available. Selection of the largest available trees as roost sites is similar to behaviour of bat species occupying unmodified forested habitats. Temperatures inside 12 maternity roosts measured during the lactation period were variable. Five roosts were well insulated from ambient conditions and internal temperatures were stable, whereas the temperatures inside seven roosts fluctuated in parallel with ambient temperature. Tree cavities used by bats at Hanging Rock were significantly nearer ground level, had larger entrance dimensions, were less well insulated, and were occupied by fewer bats than roosts in the Eglinton Valley. These characteristics appear to expose their occupants to unstable microclimates and to a higher risk of threats such as predation. We suggest that roosts at Hanging Rock are of a lower quality than those in the Eglinton Valley, and that roost quality may be one of the contributory factors in the differential reproductive fitness observed in the two bat populations. The value of introduced willows (especially Salix fragilis) as bat roosts should be acknowledged. We recommend six conservation measures to mitigate negative effects of deterioration of roosting habitat: protection and enhancement of the quality of existing roosts, replanting within roosting habitat, provision of high quality artificial roosts, predator control, and education of landowners and statutory bodies.     

Hibernation by tree-roosting bats

In summer, long-eared bats (Nyctophilus spp.) roost under bark and in tree cavities, where they appear to benefit from diurnal heating of roosts. In contrast, hibernation is thought to require a cool stable temperature, suggesting they should prefer thermally insulated tree cavities during winter. To test this prediction, we quantified the winter thermoregulatory physiology and ecology of hibernating tree-roosting bats, Nyctophilus geoffroyi and N. gouldi in the field. Surprisingly, bats in winter continued to roost under exfoliating bark (65%) on the northern, sunny side of trees and in shallow tree cavities (35%). Despite passive re-warming of torpid bats by 10-20 degrees C per day, torpor bouts lasted up to 15 days, although shorter bouts were also common. Arousals occurred more frequently and subsequent activity lasted longer on warmer nights, suggesting occasional winter foraging. We show that, because periodic arousals coincide with maximum roost temperatures, when costs of rewarming and normothermic thermoregulation are minimal, exposure to a daily temperature cycle could largely reduce energy expenditure during hibernation. Our study provides further evidence that models of torpor patterns and energy expenditure from hibernators in cold temperate climates are not directly applicable in milder climates, where prolonged torpor can be interspersed with more frequent arousals and occasional foraging.     

Natural use of heterothermy by a small, tree-roosting bat during summer

Little is known about the use of heterothermy by wild bats during summer, especially for tree-roosting species. Because thermal conditions within tree roosts can fluctuate widely with ambient temperature, which affects thermoregulatory energy expenditure during diurnal roosting, we measured skin temperatures of free-ranging male Nyctophilus geoffroyi (8 g) to quantify the relation between summer torpor use and roost thermal conditions. Bats roosted under bark on the northern (sunny) side of trees and entered torpor every day, usually near sunrise. Bats exhibited two bouts of torpor on most days: the first occurred in the morning, was terminated by partially passive rewarming, and was followed by a period of normothermy during the warmest part of the day; a second torpor bout occurred in the late afternoon, with arousal near sunset. On the warmest days, bats had only a single, short morning bout. On the coolest days, bats remained torpid throughout the day, and one 2-d bout was observed. Thus, presumably owing to their poorly insulated roosts and the high energetic cost of normothermy at temperatures below 30 degrees C, the extent and timing of heterothermy was closely related to the cycle of diurnal temperatures. Our study indicates that torpor use is important for energy maintenance during summer diurnal roosting of N. geoffroyi and likely of other small, tree-roosting bats.     

Potential energy expenditure by litter-roosting bats associated with temperature under leaf litter during winter

In temperate portions of North America, some bats that remain active during winter undergo short periods of hibernation below leaf litter on the forest floor during episodes of below-freezing weather. These winter roosts may provide above-freezing conditions, but the thermal conditions under leaf litter are unclear. Further, little is known of the relationship between temperatures under litter and potential energy expenditure by bats. Therefore, I characterized thermal conditions below leaf litter, compared temperatures encountered under different litter depths, and evaluated the quality of these sites as hibernacula based on potential energy use by eastern red bats (Lasiurus borealis) during winter in forests of the Ouachita Mountains, Arkansas, USA. Over an averaged 24-h period, there was no significant difference in temperature among different depths of leaf litter, but temperatures under litter remained significantly warmer than air temperatures, especially during nighttime and under snow cover. Temperatures below leaf litter were significantly warmer on south-facing slopes than north-facing slopes, but predicted metabolic rates did not differ among aspects. Predicted metabolic rates of eastern red bats were lowest under the deepest leaf litter measured (8 cm) and highest under ambient air conditions. Depending on depth of leaf litter cover, predicted energy savings based on O₂ consumption from roosting under litter were 1.9 to 3.1 times greater than remaining in ambient air during periods of freezing weather and around 5.6 times greater when roosting under leaf litter with snow cover. A model for predicted total energy consumption (estimated as the total oxygen consumption during a 24-h period) by eastern red bats indicated that when roosting below leaf litter, energy consumption would be reduced with greater ground temperatures, greater leaf litter moisture, and when located on south-facing slopes. Predicted metabolic rates and total energy consumption may provide more insight on the quality of roost sites for wintering bats than temperature of roost sites alone.     

Sex differences in the use of daily torpor and foraging time by big brown bats (Eptesicus fuscus) during the reproductive season

Temperate zone bats can use daily torpor as a means of saving energy. Some argue, however, that torpor is costly for both males and females and that individuals should only use it during times of poor foraging conditions. Others hypothesize that the costs are greater for females and that males should enter torpor more regularly. We tested these alternative hypotheses by using temperature-sensitive radiotransmitters to monitor use of torpor and foraging by free-ranging big brown bats ( Eptesicus fuscus ). During the pregnancy period, males used torpor at night more and foraged less often than did females. Males also went into deep torpor more often and remained in torpor longer than did females. When they foraged, males and females were away from the roosts for equal periods of time. During the lactation period, males and females rarely failed to forage and foraging times were again no different between the sexes, although males may roost at night away from the maternity colonies. Males again used torpor and deep torpor more often and for longer than females did. These results support the hypothesis that the fitness costs of using torpor are lower for males than for reproductive females and that males regularly use torpor as an energy-saving mechanism. Females enter torpor only when foraging conditions are poor, presumably because torpor prolongs gestation and slows neonatal growth thereby leaving less time for females and their young to prepare for hibernation.     

Thermal biology of flight in a butterfly: genotype,flight metabolism,and environmental conditions

Knowledge of the effects of thermal conditions on animal movement and dispersal is necessary for a mechanistic understanding of the consequences of climate change and habitat fragmentation. In particular, the flight of ectothermic insects such as small butterflies is greatly influenced by ambient temperature. Here, variation in body temperature during flight is investigated in an ecological model species, the Glanville fritillary butterfly (Melitaea cinxia). Attention is paid on the effects of flight metabolism, genotypes at candidate loci, and environmental conditions. Measurements were made under a natural range of conditions using infrared thermal imaging. Heating of flight muscles by flight metabolism has been presumed to be negligible in small butterflies. However, the results demonstrate that Glanville fritillary males with high flight metabolic rate maintain elevated body temperature better during flight than males with a low rate of flight metabolism. This effect is likely to have a significant influence on the dispersal performance and fitness of butterflies and demonstrates the possible importance of intraspecific physiological variation on dispersal in other similar ectothermic insects. The results also suggest that individuals having an advantage in low ambient temperatures can be susceptible to overheating at high temperatures. Further, tolerance of high temperatures may be important for flight performance, as indicated by an association of heat‐shock protein (Hsp70) genotype with flight metabolic rate and body temperature at takeoff. The dynamics of body temperature at flight and factors affecting it also differed significantly between female and male butterflies, indicating that thermal dynamics are governed by different mechanisms in the two sexes. This study contributes to knowledge about factors affecting intraspecific variation in dispersal‐related thermal performance in butterflies and other insects. Such information is needed for predictive models of the evolution of dispersal in the face of habitat fragmentation and climate change.     

Deeply torpid bats can change position without elevation of body temperature

Because body temperature is tightly coupled to physiological function, hibernating animals entering deep torpor are typically immobile. We analysed thermal behaviour and locomotory activity of hibernating greater mouse-eared bats Myotis myotis and found two types of movement behaviour related to body temperature, i.e. movement at high fur temperature and at low fur temperatures (Tflow; <5 °C). First Tflow movements appeared at the beginning of March and often occurred during long torpor bouts. In most cases, Tflow events represented slow displacements between clusters of bats. In several cases, however, departure or arrivals from and into clusters was also recorded without any elevation in body temperature. Distance travelled, flight duration and speed of locomotion during Tflow events was lower than in high fur temperature events. Such behaviour could allow bats to save energy long-term and prolong torpor bouts. Tflow movement in torpid bats significantly changes our understanding of basic hibernation principles and we strongly recommend further studies on the subject.     

Defining torpor in free-ranging bats: experimental evaluation of external temperature-sensitive radiotransmitters and the concept of active temperature

A variety of definitions involving body temperature (Tb), metabolic rate and behavior have been used to define torpor in mammals and birds. This problem is confounded in some studies of free-ranging animals that employ only skin temperature (Tsk), a measure that approximates but may not precisely reflect Tb. We assess the accuracy of Tsk in the context of a recent definition for torpor called active temperature. We compared the active temperatures of individual big brown bats (Eptesicus fuscus), which aggregate in cavities, with solitary, foliage-roosting hoary bats (Lasiurus cinereus). In captive big brown bats, we compared Tsk and core Tb at a range of ambient temperatures for clustered and solitary roosting animals, compared Tsk and Tb during arousal from torpor, and quantified the effect of flight on warming from torpor. Hoary bats had significantly lower active temperatures than big brown bats despite having the same normothermic Tsk. Tsk was significantly lower than Tb during normothermia but often greater than Tb during torpor. Flight increased the rate of warming from torpor. This effect was more pronounced for Tsk than Tb. This suggests that bats could rely on heat generated by flight muscles to complete the final stages of arousal. Using active temperature to define torpor may underestimate torpor due to ambient cooling of external transmitters or animals leaving roosts while still torpid. Conversely, active temperature may also overestimate shallow torpor use if it is recorded during active arousal when shivering and non-shivering thermogenesis warm external transmitters. Our findings illuminate the need for laboratory studies that quantify the relationship between metabolic rate and Tsk over a range of ambient temperatures.     

Does fear beget fear? Risk-mediated habitat selection triggers predator avoidance at lower trophic levels

Seasonal changes in weather and food availability differentially impact energy budgets of small mammals such as bats. While most thermal physiological research has focused on species that experience extreme seasonal temperature variations, knowledge is lacking from less variable temperate to subtropical climates. We quantified ambient temperature (T _a) and skin temperature (T _sk) responses by individuals from a population of New Zealand lesser short-tailed bats (Mystacina tuberculata) during summer and winter using temperature telemetry. During summer, communal roosts were more thermally stable than T _a. During winter, solitary roosts were warmer than T _a indicating significant thermal buffering. Communal roost trees were used on 83 % of observation days during summer, and individuals occupying them rarely entered torpor. Solitary roosts were occupied on 93 % of observation days during winter, and 100 % of individuals occupying them used torpor. During summer and winter, bats employed torpor on 11 and 95 % of observation days, respectively. Maximum torpor bout duration was 120.8 h and winter torpor bout duration correlated negatively with mean T _a. Torpor bout duration did not differ between sexes, although female minimum T _sk was significantly lower than males. The summer Heterothermy Index varied, and was also significantly affected by T _a. Mean arousal time was correlated with sunset time and arousals occurred most frequently on significantly warmer evenings, which are likely associated with an increased probability of foraging success. We provide the first evidence that torpor is used flexibly throughout the year by M. tuberculata, demonstrating that roost choice and season impact torpor patterns. Our results add to the growing knowledge that even small changes in seasonal climate can have large effects on the energy balance of small mammals.     

Torpor during the reproductive season in a free-ranging subtropical bat, Nyctophilus bifax

During times of energetic stress many small mammals reduce their body temperature and metabolic rate, a state known as torpor. Whereas torpor is effective in energy conservation it also entails costs, such as reduced foetal development in pregnant females. Because it is currently not known how subtropical bats deal with energetic challenges during the reproductive season, the thermal biology of free-ranging non-reproductive male and pregnant female Nyctophilus bifax was examined during spring. Males entered torpor much more frequently than pregnant females. However, night time activity periods were similar in both sexes. My results show that even in the subtropics torpor is used regularly during the reproductive period in spring by non-reproductive male N. bifax to conserve energy, but is used rarely by pregnant females likely to prevent slowed foetal development.