Regional Heterothermy Research Articles

Regional heterothermies recorded in the oxygen isotope composition of harbour seal skeletal elements

Regional heterothermy is a strategy used by marine mammals to maintain a high and stable core body temperature, but its identification needs in situ measurements difficult to set up in extant wild organisms and inapplicable to extinct ones. We have analysed the oxygen isotope composition of bioapatite phosphate (δ18Op) from one permanent tooth and from thirty-six skeletal elements of one adult male harbour seal (Phoca vitulina) from the Baie de Somme (Hauts-de-France, France). We propose that the observed intra-skeletal δ18Op variability reflects tissue temperature heterogeneities typical of the pinniped regional heterothermy strategy. Our δ18Op data indicate that bone hydroxylapatite from harbour seal autopod skeletal elements (metacarpals, metatarsals, and phalanxes) mineralises at a lower temperature than that of the bone from the axial skeleton (e.g. vertebrae, ribs, and girdle bones). The results suggest that it is possible to locate a history of regional heterothermies in amphibious marine vertebrates using the δ18Op values of their mineralised tissues. This enables direct evaluation of the thermophysiology of both modern and fossil Pinnipedia from their skeletons opening perspectives on understanding their thermal adaptation to the marine environment in the fossil record. In addition to thermophysiology, oxygen isotope data from the permanent teeth of Pinnipedia, which are formed during the in utero phase from body fluid of the mother and at a stable temperature, could be valuable for locating the geographical areas inhabited by existing Pinnipedia females during their gestation period.

A diminutive snake species can maintain regional heterothermy in both homogeneous and heterogeneous thermal environments.

Regional heterothermy is a pattern whereby different body regions are maintained at different temperatures, often to prioritize the function of certain body parts over others, or to maximize the function of organs and tissues that vary in thermal sensitivity. Regional heterothermy is relatively well understood in endotherms, where physiological mechanisms maintain heterogeneity. However, less is known about regional heterothermy in ectotherms, where behavioral mechanisms are more important for generating thermal variation. In particular, whether small and elongate ectotherms with high surface area to volume ratios such as diminutive snakes can maintain regional heterothermy, despite rapid thermal equilibration, is not yet known. We measured regional variation in body temperature and tested whether environmental heterogeneity is used to generate regional heterothermy in the ring-necked snake (Diadophis punctatus) using both field and laboratory studies. We found that ring-necked snakes have robust regional heterothermy in a variety of contexts, despite their small body size and elongate body shape. Temperature variation along the length of their bodies was not detectable when measured externally. However, snakes had higher mouth than cloacal temperatures both in the field and in laboratory thermal gradients. Further, this regional heterothermy was maintained even in ambient laboratory conditions, where the thermal environment was relatively homogeneous. Our results indicate that regional heterothermy in ring-necked snakes is not solely driven by environmental variation but is instead linked to physiological or morphological mechanisms that maintain regional variation in body temperature irrespective of environmental context.

Temperature independence of haemoglobin-oxygen affinity in smalleye Pacific opah (Lampris incognitus) and swordfish (Xiphias gladius).

Smalleye Pacific opah and swordfish can conserve metabolic heat and maintain specific body regions warmer than ambient water temperature (i.e. regional heterothermy). Consequently, blood O2 uptake at the gills occurs at the environmental temperature at which the individual is found, but O2 offloading will occur at different temperatures in different tissues. While several regionally heterothermic fishes (e.g. billfishes, tunas and sharks) show a reduced temperature effect on haemoglobin (Hb)-O2 affinity, the temperature dependence of Hb-O2 affinity in opah and swordfish is unknown. We hypothesized that the Hb of opah and swordfish would also show a reduced temperature dependence. Opah whole-blood-O2 affinity exhibited a reverse temperature dependence above 50% Hb-O2 saturation (10-20°C, pH7.2-8.0), while the temperature dependence of swordfish blood-O2 affinity (10-25°C) was saturation and pH dependent, becoming temperature independent below 50% Hb-O2 saturation and pH7.4. Experiments on stripped haemolysates showed that adding ATP ([ATP]/[Hb]=30) decreased the temperature sensitivity of Hb-O2 affinity, changing the overall oxygenation enthalpy (ΔH') values of opah (10-20°C) and swordfish (10-25°C) Hbs at pH7.4 from -15 and -42 kJ mol-1 O2, respectively, to +84 and -9 kJ mol-1 O2. Swordfish blood-O2 affinity was high compared with that of other large, pelagic, marine teleosts, which may be the result of unusually low ATP/Hb levels, but might also enable swordfish to forage in the potentially low-oxygenated water of the upper reaches of the oxygen minimum layer. The existence of Hbs with reduced temperature sensitivity in regionally heterothermic fishes may prevent marked changes in Hb-O2 affinity between the cold and warm tissues.

A comparison of thermal sensitivities of wing muscle contractile properties from a temperate and tropical bat species.

Endotherms experience temperature variation among body regions, or regional heterothermy, despite maintaining high core body temperatures. Bat forelimbs are elongated to function as wings, which makes them vulnerable to heat loss and exaggerates regional heterothermy. A tropical bat species, Carollia perspicillata, flies with distal wing muscles that are substantially (>10°C) cooler than proximal wing muscles and significantly less temperature sensitive. We hypothesized that the difference between proximal and distal wing muscles would be even more extreme in a temperate bat species that is capable of flight at variable environmental temperatures. We measured the contractile properties of the proximal pectoralis muscle and distal extensor carpi radialis muscle at a range of temperatures in the big brown bat, Eptesicus fuscus, and compared their thermal dependence with that of the same muscles in C. perspicillata. We found that, overall, temperature sensitivities between species were remarkably similar. The sole exception was the shortening velocity of the pectoralis muscle in E. fuscus, which was less temperature sensitive than in C. perspicillata. This decreased temperature sensitivity in a proximal muscle runs counter to our prediction. We suggest that the relative lability of body temperature in E. fuscus may make better pectoralis function at low temperatures advantageous.

Stress-induced changes in body surface temperature are repeatable, but do not differ between urban and rural birds.

Urbanisation can alter local microclimates, thus creating new thermal challenges for resident species. However, urban environments also present residents with frequent, novel stressors (e.g., noise, human interaction) which may demand investment in costly, self-preserving responses (e.g., the fight-or-flight response). One way that urban residents might cope with this combination of demands is by using regional heterothermy to reduce costs of thermoregulation during the stress response. In this study, we used black-capped chickadees (nurban = 9; nrural = 10) to test whether known heterothermic responses to stress exposure (here, at the bare skin around the eye): (1) varied consistently among individuals (i.e., were repeatable), and (2) were most pronounced among urban individuals compared with rural individuals. Further, to gather evidence for selection on stress-induced heterothermic responses in urban settings, we tested: (3) whether repeatability of this response was lower among birds sampled from urban environments compared with those sampled from rural environments. For the first time, we show that heterothermic responses to stress exposures (i.e. changes in body surface temperature) were highly repeatable across chronic time periods (R = 0.58) but not acute time periods (R = 0.13). However, we also show that these responses did not differ between urban and rural birds, nor were our repeatability estimates any lower in our urban sample. Thus, while regional heterothermy during stress exposure may provide energetic benefits to some, but not all, individuals, enhanced use of this response to cope with urban pressures appears unlikely in our study species.

Evidence from Tarentola mauritanica (Gekkota: Phyllodactylidae) helps validate thermography as a tool to infer internal body temperatures of lizards.

Infrared (IR) thermal imaging has become an increasingly popular tool to measure body temperature of animals. The high-resolution data it provides with short lag and minimum disturbance makes it an appealing tool when studying reptile thermal ecology. However, due to the common phenomenon of regional heterothermy and surface-to-core temperature gradients, it is essential to select the appropriate body part to measure and provide calibrations to accurately infer internal body temperatures. This work follows from a previous study on lacertid lizards to assess the reliability of thermography-measured body temperatures, from several body locations, as a proxy for internal body temperature in lizards. This study focuses on the Moorish gecko, Tarentola mauritanica, due to its distant phylogenetic relationship and its different ecology and morphology from the previously tested species. A total of 60 adult geckos of both sexes and of a range of sizes were tested in thermal gradients and subjected to a sequence of randomly assorted treatments of heating and cooling. The temperatures of the animals were periodically measured with a thermal camera at six different body parts and, immediately after, the cloacal temperature was then measured with a thermocouple probe. Body parts' temperatures, obtained thermographically, were regressed against cloacal temperature using OLS regression and the pairwise correlations were tested using Spearman coefficients. Relationships among all body parts and between all body parts and the cloaca were strong in all cases (R2>0.87, Spearman Correlation>0.95). The observed pattern was very similar to those previously obtained from lacertid lizards. Ultimately, the eye proved to provide the best overall proxy for internal temperature, when accounting for both the slope and intercept of the regression. Hence, this study provides further support for the establishment of the eye as the standard location to infer internal body temperatures of lizards through thermography.

Warm bodies, cool wings: regional heterothermy in flying bats.

Many endothermic animals experience variable limb temperatures, even as they tightly regulate core temperature. The limbs are often cooler than the core at rest, but because the large locomotor muscles of the limbs produce heat during exercise, they are thought to operate at or above core temperature during activity. Bats, small-bodied flying mammals with greatly elongated forelimbs, possess wings with large surfaces lacking any insulating fur. We hypothesized that during flight the relatively small muscles that move the elbow and wrist operate below core body temperature because of elevated heat loss. We measured muscle temperature continuously in the small fruit bat Carollia perspicillata before and during wind tunnel flights, and discretely in diverse bats at rest in Belize. We found that bats maintained high rectal temperatures, but that there was a steep proximal-to-distal gradient in wing muscle temperature. Forearm muscles were 4-6°C cooler than rectal temperature at rest and approximately 12°C cooler during flights at an air temperature of 22°C. These findings invite further study into how bats and other endotherms maintain locomotor performance in variable environments, when some muscles may be operating at low temperatures that are expected to slow contractile properties.

Assessing the reliability of thermography to infer internal body temperatures of lizards

For many years lizard thermal ecology studies have relied on the use of contact thermometry to obtain internal body temperature (Tb) of the animals. However, with progressing technology, an interest grew in using new, less invasive methods, such as InfraRed (IR) pyrometry and thermography, to infer Tb of reptiles. Nonetheless few studies have tested the reliability of these new tools. The present study tested the use of IR cameras as a non-invasive tool to infer Tb of lizards, using three differently body-sized lacertid species (Podarcis virescens, Lacerta schreiberi and Timon lepidus). Given the occurrence of regional heterothermy, we pairwise compared thermography readings of six body parts (snout, eye, head, dorsal, hind limb, tail base) to cloacal temperature (measured by a thermometer-associated thermocouple probe) commonly employed to measure Tb in field and lab studies. The results showed moderate to strong correlations (R2=0.84–0.99) between all body parts and cloacal temperature. However, despite the readings on the tail base showed the strongest correlation in all three species, it was the eye where the absolute values and pattern of temperature change most consistently followed the cloacal measurements. Hence, we concluded that the eye would be the body location whose IR camera readings more closely approximate that of the animal's internal environment. Alternatively, other body parts can be used, provided that a careful calibration is carried out. We provide guidelines for future research using thermography to infer Tb of lizards.

Measuring subcutaneous temperature and differential rates of rewarming from hibernation and daily torpor in two species of bats

Prolonged and remote measurement of body temperature (Tb) in undisturbed small hibernators was not possible in the past because of technological limitations. Although passive integrated transponders (PITs) have been used previously to measure subcutaneous temperature (Tsub) during daily torpor in a small marsupial, no study has attempted to use these devices at Tbs below 10°C. Therefore, we investigated whether subcutaneous interscapular PITs can be used as a viable tool for measuring Tb in a small hibernating bat (Nyctophilus gouldi; Ng) and compared it with measurements of Tb during daily torpor in a heterothermic bat (Syconycteris australis; Sa). The precision of transponders was investigated as a function of ambient temperature (Ta) and remote Tsub readings enabled us to quantify Tsub–Tb differentials during steady-state torpor and arousal. Transponders functioned well outside the manufacturer's recommended range, down to ~5°C. At rest, Tsub and rectal Tb (Trec) were strongly correlated for both bat species (Ng r2=0.88; Sa r2=0.95) and this was also true for N. gouldi in steady-state torpor (r2=0.93). During induced rewarming Tsub increased faster than Trec in both species. Our results demonstrate that transponders can be used to provide accurate remote measurement of Tb in two species of bats during different physiological states, both during steady-state conditions and throughout dynamic phases such as rewarming from torpor. We show that, at least during rewarming, regional heterothermy common to larger hibernators and other hibernating bats is also present in bats capable of daily torpor.

Thermal effect of blood feeding in the telmophagous fly Glossina morsitans morsitans

During feeding on warm-blooded hosts, haematophagous insects are exposed to thermal stress due to the ingestion of a meal which temperature may highly exceed their own body temperature. In order to avoid overheating and its subsequent deleterious effects, these insects respond by setting up molecular protective mechanisms such as heat shock proteins synthesis or by using thermoregulative strategies. Moreover, the duration of contact with the host depends on the way of feeding displayed by the different species (either telmophagous or solenophagous) and thus also impacts their exposure to heat. Solenophagous insects feed directly on blood vessels and are relatively slow feeders while telmophagous insects by lacerating capillaries, facilitate their access to blood and thus feed more quickly. The aim of this work was to investigate to what extent strictly telmophagous insects such as tsetse flies are exposed to thermal stress during feeding and consequently to evaluate the impact of the feeding strategy on the exposition to overheating in haematophagous insects in general. Real time thermographic analysis during feeding revealed that the flies’ body significantly heat up quite homogeneously. At the end of feeding, however, a marked regional heterothermy occurs as a consequence of the alary muscles warm up that precedes take-off. Feeding strategies, either solenophagy or telmophagy, thus appear to have a great impact on both exposition to predation risks and to thermal stress.

Survival and arm abscission are linked to regional heterothermy in an intertidal sea star

Body temperature is a more pertinent variable to physiological stress than ambient air temperature. Modeling and empirical studies on the impacts of climate change on ectotherms usually assume that body temperature within organisms is uniform. However, many ectotherms show significant within-body temperature heterogeneity. The relationship between regional heterothermy and the response of ectotherms to sublethal and lethal conditions remains underexplored. We quantified within-body thermal heterogeneity in an intertidal sea star (Pisaster ochraceus) during aerial exposure at low tide to examine the lethal and sublethal effects of temperatures of different body regions. In manipulative experiments, we measured the temperature of the arms and central disc, as well as survival and arm abscission under extreme aerial conditions. Survival was related strongly to central disc temperature. Arms were generally warmer than the central disc in individuals that survived aerial heating, but we found the reverse in those that died. When the central disc reached sublethal temperatures of 31-35°C, arms reached temperatures of 33-39°C, inducing arm abscission. The absolute temperature of individual arms was a poor predictor of arm abscission, but the arms lost were consistently the hottest at the within-individual scale. Therefore, the vital region of this sea star may remain below the lethal threshold under extreme conditions, possibly through water movement from the arms to the central disc and/or evaporative cooling, but at the cost of increased risk of arm abscission. Initiation of arm abscission seems to reflect a whole-organism response while death occurs as a result of stress acting directly on central disc tissues.

Reduced and reversed temperature dependence of blood oxygenation in an ectothermic scombrid fish: implications for the evolution of regional heterothermy?

Tunas (family Scombridae) are exceptional among most teleost fishes in that they possess vascular heat exchangers which allow heat retention in specific regions of the body (termed 'regional heterothermy'). Seemingly exclusive to heterothermic fishes is a markedly reduced temperature dependence of blood-oxygen (blood-O(2)) binding, or even a reversed temperature dependence where increasing temperature increases blood-O(2) affinity. These unusual binding properties have been documented in whole blood and in haemoglobin (Hb) solutions, and they are hypothesised to prevent oxygen loss from arteries to veins within the vascular heat exchangers and/or to prevent excessive oxygen unloading to the warm tissues and ensure an adequate supply of oxygen to tissues positioned efferent to the heat exchangers. The temperature sensitivity of blood-O(2) binding has not been characterised in an ectothermic scombrid (mackerels and bonitos), but the existence of the unusual binding properties in these fishes would have clear implications for their proposed association with regional heterothermy. Accordingly, the present study examined oxygenation of whole blood of the chub mackerel (Scomber japonicus) at 10, 20 and 30 degrees C and at 0.5, 1 and 2% CO(2). Oxygen affinity was generally highest at 20 degrees C for all levels of CO(2). Temperature-independent binding was observed at low (0.5%) CO(2), where the PO(2) at 50% blood-O(2) saturation (P (50)) was not statistically different at 10 and 30 degrees C (2.58 vs. 2.78 kPa, respectively) with an apparent heat of oxygenation (H degrees ) close to zero (-6 kJ mol(-1)). The most significant temperature-mediated difference occurred at high (2%) CO(2), where the P (50) at 10 degrees C was twofold higher than that at 20 degrees C with a corresponding H degrees of +43 kJ mol(-1). These results provide clear evidence of independent and reversed open-system temperature effects on blood oxygenation in S. japonicus, and it is therefore speculated that these unusual blood-O(2) binding characteristics may have preceded the evolution of vascular heat exchangers and regional heterothermy in fishes.

The heterothermic loophole exploited by penguins

Penguins are generally smaller than the predicted minimum body size for marine endotherms. Anatomical observations of the little blue penguin (Eudyptula minor) suggest that extant penguins actively defeat the lower size threshold using regional heterothermy. The wing arteries in the little blue penguin form a humeral plexus, a series of three parallel arteries (including the marginal artery) that stem from the axillary artery, replacing a single brachial artery. Each artery of the plexus is associated with at least two veins to form a counter current heat-retention system. The humeral plexus thus restricts heat from entering distal areas of the wing and dissipating into seawater, ultimately reducing the bulk mass required for heat production and insulation. Humeral plexi are confirmed as a synapomorphic character of the Spheniscinae, with the humeral plexus of E. minor most similar to that of the African penguin Spheniscus demersus. The humeral plexus represents a loophole in body size minima of marine endotherms and offers insight into the ecology of living penguins.

Sympathetic α-adrenergic regulation of blood flow and volume in hamsters arousing from hibernation

Mammals arousing from hibernation display pronounced regional heterothermy, where the thoracic and head regions warm faster than the abdominal and hindlimb regions. We used laser-Doppler flowmetry to measure peripheral hind foot blood flow during hibernation and arousal and gamma imaging of technetium-labeled albumin to measure whole blood volume distribution in hamsters arousing from hibernation. It was discovered that the hibernating hamster responds to physical but not to sound or hypercapnic stimulation with rapid, 73% reduction of hind foot blood flow. Hind foot blood flow vasoconstriction was maintained from the onset of arousal until late in arousal when rectal temperature was rapidly increased. alpha-Adrenergic blockade early in arousal increased hind foot blood flow by 700%, suggesting that vasoconstriction was mediated by activation of sympathetic tone. Gamma imaging revealed that, by the early phase of arousal from hibernation, the blood volume of the body below the liver is greatly reduced, whereas blood volumes of the thorax and head are much greater than corresponding volumes in anesthetized hamsters. As arousal progresses and cardiac activity increases and regional heterothermy develops, this regional blood volume distribution is largely maintained; however, blood volume slowly decreases in the thoracic region and slowly increases in the shoulder and head regions. The rapid increase in rectal temperature, characteristic of mid- to late- arousal phases, is probably mediated, in part, by reduction of adrenergic tone on abdominal and hindlimb vasculature. Warm blood then moves into the hind body, produces an increase in temperature, blood flow, and blood volume in the hind body and compensatory reductions of blood volume in the neck, head, and thoracic regions.

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice

Temperatures were recorded at several body sites in emperor penguins ( Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (±S.E.) in central body sites during dives were: stomach: 37.1±0.2 °C ( n=101 dives in five birds), pectoral muscle: 37.8±0.1 °C ( n=71 dives in three birds) and axillary/brachial veins: 37.9±0.1 °C ( n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=−0.59, P<0.05; range <1 °C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6±0.7 °C ( n=157 dives in three birds), 20.2±1.2 °C ( n=69 in three birds) and 35.2±0.2 °C ( n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds ( r=−0.29 to −0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration ( r=−0.49 and −0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 °C during rest periods to a grand mean of 15.0±0.7 °C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird ( r=−0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37–39 °C more closely than the anterior abdominal temperatures (19–30 °C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.

Temperature regulation in emperor penguins foraging under sea ice

Inferior vena caval (IVC) and anterior abdominal (AA) temperatures were recorded in seven emperor penguins ( Aptenodytes forsteri) foraging under sea ice in order to evaluate the hypothesis that hypothermia-induced metabolic suppression might extend aerobic diving time. Diving durations ranged from 1 to 12.5 min, with 39% of dives greater than the measured aerobic dive limit of 5.6 min. Anterior abdominal temperature decreased progressively throughout dives, and partially returned to pre-dive values during surface intervals. The lowest AA temperature was 19°C. However, mean AA temperatures during dives did not correlate with diving durations. In six of seven penguins, only minor fluctuations in IVC temperatures occurred during diving. These changes were often elevations in temperature. In the one exception, although IVC temperatures decreased, the reductions were less than those in the anterior abdomen and did not correlate with diving durations. Because of these findings, we consider it unlikely that regional hypothermia in emperor penguins leads to a significant reduction in oxygen consumption of the major organs within the abdominal core. Rather, temperature profiles during dives are consistent with a model of regional heterothermy with conservation of core temperature, peripheral vasoconstriction, and cooling of an outer body shell.

The thermoregulatory functions of the Triceratops frill and horns: heat flow measured with oxygen isotopes

ABSTRACT Variability of body temperature within fossil vertebrates can be measured by the oxygen isotopie composition of bone phosphate (δp). Intrabone and interbone δp variations indicate heat flow within an individual and can be used to establish thermoregulatory strategies such as homeothermy and regional heterothermy. This method is applied to an individual Triceratops skeleton where the results suggest very high and uniform heat flow through the parietal frill, maintaining mean frill temperatures between 0–4°C below the body core. Heat flow to the postorbital horn-core is much more variable. The frill and horn cores of Triceratops were used as thermoregulatory structures with the horn cores interpreted as being especially important in the stabilization of brain temperatures at extreme ambient temperatures and the frill serving a more consistent role in body temperature regulation.

Comparison of Thermoregulation of Four Ornithischian Dinosaurs and a Varanid Lizard from the Cretaceous Two Medicine Formation: Evidence from Oxygen Isotopes

The 180 values of vertebrate bone phosphate are dependent upon the 180O values of their body water and the temperature of bone deposition. This allows the measurement of the temperature fluctuation during bone deposition for individual skeletal elements within any individual. 8180 results for bones from a varanid lizard and four Late Cretaceous ornithischian dinosaurs (Orodromeus makelai, Hypacrosaurus stebingeri, Montanaceratops sp., and a juvenile Achelousaurus horneri) from the Campanian Two Medicine Formation in Montana are compared. Results from the varanid indicate that bone deposition occurred over a range of 10-15?C, as expected for an ectotherm. Results from the dinosaurs indicate that bone deposition occurred over a range 2-7?C within the body core and 3-8 C in the extremities. The intrabone and interbone pattern of 6180 values suggests that these dinosaurs were primarily homeotherms with all but Orodromeus exhibiting regional heterothermy. Homeothermy over the size range of the individual dinosaurs suggests that they had metabolic rates elevated above those of modern ectotherms and the range of isotopic variability suggests intermediate endothermy.

Thermoregulation of the North American Mink during Rest and Activity in the Aquatic Environment

Thermoregulation during swimming and resting in water was investigated for the North American mink (Mustela vision, Schreber) in summer and winter pelage by measuring total heat production, heat storage, and minimum thermal conductance. Swimming activity over the range of 0.13-0.60 m·s⁻¹ resulted in heat losses that exceeded metabolic heat production within 5 min of immersion. This imbalance, manifested as a decline in core body temperature (core Tb), resulted in negative levels of heat storage. Negative heat storage for mink was greatest at intermediate swimming speeds. Maximum levels were -14.28 W·kg⁻¹ at 0.29 m·s⁻¹ and -8.72 W·kg⁻¹ at 0.36 m·s⁻¹ for mink in summer and winter pelage, respectively. Thermal conductance determined from cooling curves of carcasses and a taxidermic mount increased linearly with water speed and was comparatively greater at all speeds for animals in summer pelage. Patterns of cooling for carcasses, the taxidermic mount, and swimming mink were similar. In combination with regional heterothermy and elevated heat production, insulation provided by an air layer pervading the fur will permit short bouts of swimming in which Tb is maintained by foraging mink.

Regional heterothermy in hibernated and non-hibernated woodchucks, Marmota Monax, at various ambient temperatures

Regional heterothermy in hibernated and non-hibernated woodchucks, Marmota Monax, at various ambient temperatures