Overwinter body temperature patterns in captive jerboas (<Emphasis Type="Italic">Jaculus orientali</Emphasis>s): influence of sex and group

The jerboa (Jaculus orientalis) has been described in the past as a hibernator, but no reliable data exist on the daily and seasonal rhythmicity of body temperature (T _b). In this study, T _b patterns were determined in different groups of jerboas (isolated males and females, castrated males and grouped animals) maintained in captivity during autumn and winter, and submitted to natural variations of light and ambient temperature (T _a). T _b and T _a variations were recorded with surgically implanted iButton temperature loggers at 30-min intervals for two consecutive years. About half (6/13) of isolated female jerboas hibernated with a T _b < 33°C, with hibernation bouts interspersed with short periods of normothermy from November to February. Hibernation bout durations were longer (4–5 days) than those of normothermia phases (1–4 days). During hibernation, the minimum T _b was low (T _bmin ~10.7°C). In contrast, one of the 12 isolated males showed short hibernation bouts of ca. 2 days late in the hibernation season, February–March. The males had T _bmin values of 15.1°C. In contrast to predictions, no castrated males hibernated. When jerboas were grouped, females and males exhibited concomitant torpor bouts. In males, the longest bouts were observed during the late hibernation season. These data suggest complex regulation of hibernation in jerboas.     

Factors affecting the daily rhythm of body temperature of captive mouse lemurs (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Microcebus murinus, a small nocturnal Malagasy primate, exhibits adaptive energy-saving strategies such as daily hypothermia and gregarious patterns during diurnal rest. To determine whether ambient temperature (T_a), food restriction and nest sharing can modify the daily body temperature (T_b) rhythm, T_b was recorded by telemetry during winter in six males exposed to different ambient temperatures (T_a=25, 20, 15°C) and/or to a total food restriction for 3 days depending on social condition (isolated versus pair-grouped). At 25°C, the daily rhythm of T_b was characterized by high T_b values during the night and lower values during the day. Exposure to cold significantly decreased minimal T_b values and lengthened the daily hypothermia. Under food restriction, minimal T_b values were also markedly lowered. The combination of food restriction and cold induced further increases in duration and depth of torpor bouts, minimal T_b reaching a level just above T_a. Although it influenced daily hypothermia less than environmental factors, nest sharing modified effects of cold and food restriction previously observed by lengthening duration of torpor but without increasing its depth. In response to external conditions, mouse lemurs may thus adjust their energy expenditures through daily modifications of both the duration and the depth of torpor.     

Seasonality of torpor and thermoregulation in three dasyurid marsupials

Summary Seasonal variation in the pattern of torpor and temperature regulation was investigated in the closely related arid zone dasyurid marsupialsSminthopsis crassicaudata (17 g),S. macroura (24 g), andDasyuroides byrnei (120 g). The tendency to enter torpor was greater, torpor commenced earlier, torpor duration was longer, and body temperatures (T _b) were lower inSminthopsis spp. than inD. byrnei. The minimum mass-specific rate of oxygen consumption ( ) of torpid animals was similar among the three species despite the differences in minimumT _b. The mass-specific oxygen consumption of normothermic animals was reduced during winter when compared with the summer values in all species, but there was no seasonal variation in normothermicT _b in any species. The tendency to enter torpor was incrased during winter. TorpidSminthopsis spp. had lower values ofT _b and during winter than during summer;D. byrnei did not show seasonal changes in these variables. These results suggest that seasonal changes in the pattern of thermoregulation and torpor in small dasyurids may be more distinct than in larger species.Abbreviations RMR resting metabolic rate - BMR basal metabolic rate     

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.     

Basking and torpor in a rock-dwelling desert marsupial: survival strategies in a resource-poor environment

Australian deserts are characterized by unpredictability, low primary productivity, and high temperature fluctuations. Despite these adverse conditions the diversity of small insectivorous marsupials of the family Dasyuridae is surprisingly high. We quantified the thermal biology of the dasyurid Pseudantechinus madonnellensis (body mass ∼30 g) in the wild to gain some understanding of whether the success of dasyurids in the arid zone may be related to some extent to their use of energy conservation strategies. In winter, most free-ranging Pseudantechinus frequently (58.3% of 131 animal days) entered daily torpor after midnight (mean 0157 hours) in rock crevices when outside ambient temperatures (T _a) were low. Most animals remained torpid until the next morning when they moved while still torpid from rock crevices to sun-exposed basking sites. We visually observed basking during rewarming from torpor (mean commencement at 0943 hours) at body temperatures (T _b) as low as 19.3°C when radiant heat was high and T _a was rising. Basking continued for the rest of the day. Torpor use was not strongly correlated with T _a, but the temporal organization of daily torpor and activity were apparently linked to the thermal characteristics of basking sites. Our study suggests that by frequently employing daily torpor and basking and by appropriately coordinating their thermal biology with that of specific locations in their environment, Pseudantechinus can reduce daily energy expenditure and thus can live and reproduce in a challenging environment. It is likely that the success of other small dasyurids and perhaps many other small mammals living in deserts is linked to employment of torpor and basking for energy conservation.     

Winter body temperature patterns in free-ranging Cape ground squirrel, Xerus inauris: no evidence for torpor

The body temperature (T _b) of Cape ground squirrels (Xerus inauris, Sciuridae) living in their natural environment during winter has not yet been investigated. In this study we measured abdominal T _b of eight free-ranging Cape ground squirrels over 27 consecutive days during the austral winter. Mean daily T _b was relatively stable at 37.0 ± 0.2°C (range 33.4 to 40.2°C) despite a marked variation in globe temperature (T _g) (range −7 to 37°C). Lactating females (n = 2) consistently had a significantly higher mean T _b (0.7°C) than non-lactating females (n = 3) and males. There was a pronounced nychthemeral rhythm with a mean active phase T _b of 38.1 ± 0.1°C and a mean inactive phase T _b of 36.3 ± 0.3°C for non-lactating individuals. Mean daily amplitude of T _b rhythm was 3.8 ± 0.2°C. T _b during the active phase closely followed T _g and mean active phase T _b was significantly correlated with mean active phase T _g (r ² = 0.3–0.9; P < 0.01). There was no evidence for daily torpor or pronounced hypothermia during the inactive phase, and mean minimum inactive phase T _b was 35.7 ± 0.3°C for non-lactating individuals. Several alternatives (including nocturnal huddling, an aseasonal breeding pattern and abundant winter food resources) as to why Cape ground squirrels do not employ nocturnal hypothermia are discussed.     

The energetics of basking behaviour and torpor in a small marsupial exposed to simulated natural conditions

Limited information is available on basking behaviour in torpid mammals and its energetic consequences. We investigated the effects of physiological and behavioural strategies on the energetics of the fat-tailed dunnart (Sminthopsis crassicaudata). Metabolic rate and body temperature during torpor, basking and rest were measured over 24 h in response to simulated environmental conditions: (a) constant ambient temperature (T _a) of 15°C, (b) constant T _a of 15°C with access to a radiant heat lamp, (c) a T _a cycle (range 15–31°C), and (d) a T _a cycle with access to a radiant heat lamp. When a radiant heat source was provided, all dunnarts (n = 16) basked during all measurements, which resulted in energy savings of up to 74% during rest. Overall, torpor was used on 59% of measurements with a maximum duration of 16.2 h and reductions in metabolic rate of 90% compared to normothermic values. Torpid dunnarts actively moved from a shaded area to position themselves under the heat lamp with body temperatures as low as 17.5°C and thereby reduced rewarming costs by 66%. We demonstrated, for the first time in the laboratory, that torpid animals actively move to a heat source to bask, and that this behaviour results in considerable energy savings. Our finding supports the view that basking during normothermia and rewarming from torpor substantially reduces energetic requirements, which may be important for the survival of small dasyurids living on limited resources in the Australian arid zone.     

Daily torpor and energetics in a tropical mammal, the northern blossom-bat Macroglossus minimus (Megachiroptera)

Little is known about torpor in the tropics or torpor in megachiropteran species. We investigated thermoregulation, energetics and patterns of torpor in the northern blossom-bat Macroglossus minimus (16 g) to test whether physiological variables may explain why its range is limited to tropical regions. Normothermic bats showed a large variation in body temperature (T _b) (33 to 37 °C) over a wide range of ambient temperatures (T _as) and a relatively low basal metabolic rate (1.29 ml O₂ g⁻¹h⁻¹). Bats entered torpor frequently in the laboratory at T _as between 14 and 25 °C. Entry into torpor always occurred when lights were switched on in the morning, independent of T _a. MRs during torpor were reduced to about 20–40% of normothermic bats and T _bs were regulated at a minimum of 23.1 ± 1.4 °C. The duration of torpor bouts increased with decreasing T _a in non-thermoregulating bats, but generally terminated after 8 h in thermoregulating torpid bats. Both the mean minimum T _b and MR of torpid M. minimus were higher than that predicted for a 16-g daily heterotherm and the T _b was also about 5 °C higher than that of the common blossom-bat Syconycteris australis, which has a more subtropical distribution. These observations suggest that variables associated with torpor are affected by T _a and that the restriction to tropical areas in M. minimus to some extent may be due to their ability to enter only very shallow daily torpor. Accepted: 22 September 1997     

Daily torpor in the gray mouse lemur (Microcebus murinus) in Madagascar: energetic consequences and biological significance

Patterns and energetic consequences of spontaneous daily torpor were measured in the gray mouse lemur (Microcebus murinus) under natural conditions of ambient temperature and photoperiod in a dry deciduous forest in western Madagascar. Over a period of two consecutive dry seasons, oxygen consumption (VO₂) and body temperature (T _b) were measured on ten individuals kept in outdoor enclosures. In all animals, spontaneous daily torpor occurred on a daily basis with torpor bouts lasting from 3.6 to 17.6 h, with a mean torpor bout duration of 9.3 h. On average, body temperatures in torpor were 17.3±4.9°C with a recorded minimum value of 7.8°C. Torpor was not restricted to the mouse lemurs’ diurnal resting phase: entries occurred throughout the night and arousals mainly around midday, coinciding with the daily ambient temperature maximum. Arousal from torpor was a two-phase process with a first passive, exogenous heating where the T _b of animals increased from the torpor T _b minimum to a mean value of 27.1°C before the second, endogenous heat production commenced to further raise T _b to normothermic values. Metabolic rate during torpor (28.6±13.2 ml O₂ h^–1) was significantly reduced by about 76% compared to resting metabolic rate (132.6±50.5 ml O₂ h^–1). On average, for all M. murinus individuals measured, hypometabolism during daily torpor reduced daily energy expenditure by about 38%. In conclusion, all these energy-conserving mechanisms of the nocturnal mouse lemurs, with passive exogenous heating during arousal from torpor, low minimum torpor T _bs, and extended torpor bouts into the activity phase, comprise an important and highly adapted mechanism to minimize energetic costs in response to unfavorable environmental conditions and may play a crucial role for individual fitness. Received: 8 July 1999 / Accepted: 3 December 1999     

Thermoregulation in free‐ranging ground woodpeckers Geocolaptes olivaceus: no evidence of torpor

Heterothermic responses characterised by pronounced hypometabolism and reductions in body temperature (T_b) are one of the most effective ways in which small endotherms can offset the energetic cost of endothermic homeothermy. It remains unclear, therefore, why daily torpor and hibernation are restricted to only a subset of avian lineages. To further our understanding of the phylogenetic distribution of avian torpor, we investigated winter thermoregulation in the southern African ground woodpecker Geocolaptes olivaceus. We considered this species a good candidate for heterothermy, because it is resident year‐round in mountainous regions with cold winters and reliant on small ectothermic prey. We recorded T_b patterns in free‐ranging individuals and measured T_b and metabolic rates in captive individuals. Neither free‐ranging nor captive woodpeckers showed any indication of daily torpor or even shallow rest‐phase hypothermia. All birds maintained bimodally distributed T_b characteristic of classic endothemic homeothermy, with a mean rest‐phase T_b of 37.9 ± 0.2°C and no data below 37.0°C. The mean circadian amplitude of T_b was 4.2°C, equivalent to approximately twice the expected value. There was some evidence of seasonal acclimatisation in T_b, with a small decrease in rest‐phase T_b with the onset of the austral winter. Captive birds showed patterns of resting metabolic rate and T_b consistent with the classic model of endothermic homeothermy. The apparent absence of torpor in G. olivaceus supports the notion that, unlike the case in mammals, many avian taxa that may a priori be expected to benefit from deep heterothermy do not use it.     

How to budget metabolic energy: torpor in a small Neotropical mammal

Neotropical nectar-feeding bats (Glossophaginae) are highly specialized in the exploitation of floral nectar and have one of the highest mass-specific metabolic rates among mammals. Nevertheless, they are distributed throughout the tropics and subtropics over a wide elevational range, and thus encounter many extreme and energetically challenging environmental conditions. Depressing their otherwise high metabolic rate, e.g., in situations of food restriction, might be an important adaptive physiological strategy in these dietary specialists. We investigated the thermoregulatory behavior of captive 10-g nectar feeding bats (Glossophaga soricina; Chiroptera, Phyllostomidae) under variable ambient temperatures (T _a) and feeding regimes and predicted that bats would use torpor as an energy-conserving behavior under energetic constraints. All tested animals entered torpor in response to energetic restrictions and the depth of torpor was dependent on the body condition of the animals and hence on their degree of physiological constraints. Periods of torpor with body temperatures (T _b) below 34°C were precisely adjusted to the photoperiod. The median length of diurnal torpor was 11.43 h. The lowest T _b measured was 21°C at a T _a of 19°C. Estimated energy savings due to torpor were considerable, with reductions in metabolic rate to as low as 5% of the metabolic rate of normothermic bats at the same T _a. However, contrary to temperate zone bats that also employ diurnal torpor, G. soricina regulated their T _b to the highest possible levels given the present energetic supplies. To summarize, G. soricina is a precise thermoregulator, which strategically employs thermoregulatory behavior in order to decrease its energy expenditure when under energetic restrictions. This adaptation may play a crucial role in the distribution and the assembly of communities of nectar-feeding bats and may point to a general capacity for torpor in tropical bats.     

THE TEMPORAL ORGANIZATION OF DAILY TORPOR AND HIBERNATION: CIRCADIAN AND CIRCANNUAL RHYTHMS

Mammals and birds have evolved the ability to maintain a high and constant body temperature T_b over a wide range of ambient temperatures T_a using endogenous heat production. In many, especially small endotherms, cost for thermoregulatory heat production can exceed available energy; to overcome these energetic bottlenecks, they enter a state of torpor (a regulated reduction of T_b and metabolic rate). Since the occurrence of torpor in many species is a seasonal event and occurs at certain times of the day, we review whether circadian and circannual rhythms, important in the timing of biological events in active animals, also play an important role during torpor when T_b is reduced substantially and may even fall below 0°C. The two distinct patterns of torpor, hibernation (prolonged torpor) and daily torpor, differ substantially in their interaction with the circadian system. Daily torpor appears to be integrated into the normal circadian rhythm of activity and rest, although torpor is not restricted only to the normal rest phase of an animal. In contrast, hibernation can last for several days or even weeks, although torpor never spans the entire hibernation season, but is interrupted by periodic arousals and brief normothermic periods. Clearly, a day is no longer divided in activity and rest, and at first glance the role of the circadian system appears negligible. However, in several hibernators, arousals not only follow a regular pattern consistent with a circadian rhythm, but also are entrainable by external stimuli such as photoperiod and T_a. The extent of the interaction between the circadian and circannual system and hibernation varies among species. Biological rhythms of hibernators for which food availability appears to be predictable seasonally and that hibernate in deep and sealed burrows show little sensitivity to external stimuli during hibernation and hence little entrainability of arousal events. In contrast, opportunistic hibernators, which some times use arousals for foraging and hibernate in open and accessible hibernacula, are susceptible to external zeitgebers. In opportunistic hibernators, the circadian system plays a major role in maintaining synchrony between the normal day-night cycle and occasional foraging. Although the daily routine of activity and rest is abandoned during hibernation, the circadian system appears to remain functional, and there is little evidence it is significantly affected by low T_b. (Chronobiology International, 17(2), 103–128, 2000)     

Hibernation and daily torpor in an armadillo, the pichi (Zaedyus pichiy)

Hibernation and daily torpor are physiological strategies to cope with energetic challenges that occur in many mammalian and avian taxa, but no reliable information exists about daily torpor or hibernation for any xenarthran. Our objective was to determine whether the pichi (Zaedyus pichiy), a small armadillo (Xenarthra, Dasypodidae) that inhabits arid and semi-arid habitats in central and southern Argentina and Chile, enters shallow daily torpor or prolonged deep hibernation during winter when environmental temperature and food availability are low. We studied body temperature changes during winter in semi-captive pichis by means of temperature dataloggers implanted subcutaneously. All individuals entered hibernation, characterized by torpor events of 75 ± 20 h during which the subcutaneous temperature (T_sc) decreased to 14.6 ± 2.1 °C. These events were interrupted by periods of euthermia of 44 ± 38 h with a T_sc of 29.1 ± 0.7 °C. After the hibernation season, daily torpor bouts of 4 to 6 h occurred irregularly, with T_sc dropping to as low as 24.5 °C. We conclude that the pichi is a true hibernator and can enter daily torpor outside of the hibernation season.     

Heterothermy in an Australian passerine, the Dusky Woodswallow (Artamus cyanopterus)

Information regarding passerine heterothermy and torpor is scant, although many species are small and must cope with a fluctuating food supply and presumably would benefit from energy savings afforded by torpor. We studied whether insectivorous Dusky Woodswallows (Artamus cyanopterus; ∼35 g) enter spontaneous torpor (food ad libitum) when held outdoors as a pair in autumn/winter. Woodswallows displayed pronounced and regular daily fluctuations in body temperature (T _b) over the entire study period. The mean T _b ranged from ∼39°C to 40°C (photophase, day time) and ∼33°C to 36°C (scotophase, night time). However, on 88% of bird nights, nocturnal T _b minima fell to < 35°C. The lowest T _b observed in air was 29.2°C. However, when a bird fell into water its T _b dropped further to ∼22°C; this T _b was regulated for several hours and the bird survived. Our observations suggest that heterothermy is a normal part of the daily thermal regime for woodswallows to minimise energy expenditure. Spontaneous nocturnal torpor in captive woodswallows suggests that torpor in the wild may be more pronounced than recorded here because free-living birds are likely challenged by both low food availability and adverse weather.     

Warming up for dinner: torpor and arousal in hibernating Natterer’s bats (Myotis nattereri) studied by radio telemetry

The frequency and function of arousals during hibernation in free-living mammals are little known. We used temperature-sensitive radio transmitters to measure patterns of torpor, arousal and activity in wild Natterer’s bats Myotis nattereri during hibernation. Duration of torpor bouts ranged from 0.06 to 20.4 days with individual means ranging from 0.9 to 8.9 days. Arousals from torpor occurred most commonly coincident with the time (relative to sunset) typical for bats emerging from summer roosts to forage. Bats with lower body condition indices had a shorter average duration of their torpor bouts. We found a non-linear relationship between duration of torpor bout and ambient temperature: the longest average torpor bouts were at temperatures between 2 and 4°C with shorter bouts at lower and higher ambient temperatures. One individual was radio-tracked for ten nights, remained active for an average of 297 min each night and was active for longer on warmer nights. Our results suggest that vespertilionid bats use relatively short torpor bouts during hibernation in a location with a maritime climate. We hypothesise that Natterer’s bats time arousals to maximise opportunities for potential foraging during winter although winter feeding is not the sole determinant of arousal as bats still arouse at times when foraging is unlikely.     

Evidence of facultative daytime hypothermia in a small passerine wintering at northern latitudes

The use of hypothermia as a means to save energy is well documented in birds. This energy‐saving strategy is widely considered to occur exclusively at night in diurnally active species. However, recent studies suggest that facultative hypothermia may also occur during the day. Here, we document the use of daytime hypothermia in foraging Black‐capped Chickadees Poecile atricapillus wintering in eastern Canada. We measured the body temperature (T_b) of 126 individuals (plus 48 repeated measures) during a single winter and related values to ambient temperature (T_a) at the time of capture. We also tested whether daytime hypothermia was correlated with the size of body reserves (residuals of mass on structural size and fat score) and levels of metabolic performance (basal metabolic rate and maximum thermogenic capacity). We found that T_b of individual birds was lower when captured at low T_a, reaching values as low as 35.5 °C in actively foraging individuals. T_b was unrelated to metabolic performance or measures of body reserves. Therefore, daytime hypothermia does not result from individuals being unable to maintain T_b during cold spells or to a lack of body reserves. Our data also demonstrated a high level of individual variation in the depth of hypothermia, the causes of which remain to be explored.     

Data logging of body temperatures provides precise information on phenology of reproductive events in a free-living arctic hibernator

Precise measures of phenology are critical to understanding how animals organize their annual cycles and how individuals and populations respond to climate-induced changes in physical and ecological stressors. We show that patterns of core body temperature (T _b) can be used to precisely determine the timing of key seasonal events including hibernation, mating and parturition, and immergence and emergence from the hibernacula in free-living arctic ground squirrels (Urocitellus parryii). Using temperature loggers that recorded T _b every 20 min for up to 18 months, we monitored core T _b from three females that subsequently gave birth in captivity and from 66 female and 57 male ground squirrels free-living in the northern foothills of the Brooks Range Alaska. In addition, dates of emergence from hibernation were visually confirmed for four free-living male squirrels. Average T _b in captive females decreased by 0.5–1.0°C during gestation and abruptly increased by 1–1.5°C on the day of parturition. In free-living females, similar shifts in T _b were observed in 78% (n = 9) of yearlings and 94% (n = 31) of adults; females without the shift are assumed not to have given birth. Three of four ground squirrels for which dates of emergence from hibernation were visually confirmed did not exhibit obvious diurnal rhythms in T _b until they first emerged onto the surface when T _b patterns became diurnal. In free-living males undergoing reproductive maturation, this pre-emergence euthermic interval averaged 20.4 days (n = 56). T _b-loggers represent a cost-effective and logistically feasible method to precisely investigate the phenology of reproduction and hibernation in ground squirrels.     

Comparison of hibernation, estivation and daily torpor in the edible dormouse, Glis glis

Three major forms of dormancy in mammals have been classified: hibernation in endotherms is characterised by reduced metabolic rate (MR) and body temperature (T _b) near ambient temperature (T _a) over prolonged times in the winter. Estivation is a similar form of dormancy in a dry and hot environment during summertime. Daily torpor is defined as reduced MR and T _b lower than 32 °C, limited to a duration of less than 24 h. The edible dormouse (Glis glis) is capable for all three distinct forms of dormancy. During periods of food restriction and/or low T _a, daily torpor is displayed throughout the year, alternating with hibernation and estivation in winter and summer respectively. We recorded T _b, O₂-consumption and CO₂-production in unrestrained dormice at different T _a's for periods of up to several months. Cooling rate and rate of metabolic depression during entrance into the torpid state was identical in all three forms of dormancy. The same was true for thermal conductance, maximum heat production, duration of arousal and cost of an arousal. The only difference between hibernation and daily torpor was found in the bout duration. A daily torpor bout lasted 3–21 h, a hibernation bout 39–768 h. As a consequence of prolonged duration, MR, T _b and also the T _b − T _a gradient decreased to lower values during hibernation bouts when compared to daily torpor bouts. Our findings suggest that all three forms of dormancy are based on the same physiological mechanism of thermal and metabolic regulation. Accepted: 27 June 2000     

Heterothermy in the southern African hedgehog, <Emphasis Type="Italic">Atelerix frontalis</Emphasis>

Most research on mammalian heterothermic responses in southern Africa tends to be laboratory based and biased towards rodents and smaller members of the Afrotheria. In this study, we continuously measured body temperature of southern African hedgehogs (Atelerix frontalis) between April and August 2009 (−10°C < T _a < 43°C), kept under semi-captive conditions. A. frontalis showed a high propensity for torpor with animals spending up to 84% of the measurement period torpid. During this study, A. frontalis displayed the lowest T _b min (ca 1°C) yet recorded in an Afrotropical placental heterotherm. Bout lengths of between 0.7 h (40 min) and 116.3 h (4.8 days) were recorded. Differences in bout length were observed between lighter individuals compared with an individual exhibiting a higher body mass at the onset of winter, with low M _b individuals exhibiting daily torpor whereas a heavier individual exhibited torpor bouts that were indicative of hibernation. Our results suggest that heterothermic responses are an important feature in the energy balance equation of this species and that body mass at the onset of winter may determine the patterns of heterothermy utilised in this species.     

Absence of heterothermy in the European red squirrel (Sciurus vulgaris)

The European red squirrel Sciurus vulgaris inhabits areas which undergo profound seasonal declines in food availability and ambient temperature. We measured the body temperature (T_b) of free-ranging S. vulgaris over the course of one year to examine its thermoregulatory strategies and found no evidence of heterothermy, with T_b never dropping below 36.7 °C. A lower average T_b and a reduced active phase are likely to have resulted in some energetic savings, sufficient for survival during the particularly mild winter with unhindered access to food stores. We cannot exclude that a different T_b pattern may be seen in energetically more demanding years, but we can confirm that heterothermy is not an obligatory behaviour in this species to counter energetic bottlenecks. Either S. vulgaris is indeed a strict homeotherm, or the need for torpor is flexibly adjusted.