Cutaneous Evaporative Water Loss Research Articles

Comparing the Contributions of Cutaneous and Respiratory Evaporative Water Loss to Cooling in Anna’s Hummingbirds (Calypte anna) Under Fasted and Fed Conditions

During high-temperature periods, endothermic animals evaporate water from the body surface to cool down. The two primary routes of evaporative water loss are: (1) evaporating water from the skin; and (2) evaporation from the respiratory surfaces during breathing and panting. In smaller avian species like Anna's hummingbirds ( Calypte anna), the higher cutaneous surface area to volume ratio presents an opportunity for the skin to become an important avenue for evaporative cooling, possibly becoming a major site. Additionally, hummingbirds feed on water-rich nectar. As a result, they eliminate surplus water through urination, but significant evaporation can also occur through their skin and respiratory system. Thus, total evaporative water loss could rise significantly for these animals when in a hydrated state. Here, this study investigates the relative importance of the two evaporative water loss routes (cutaneous and respiratory) of Anna's hummingbirds in fed and fasted states. We hypothesize that cutaneous evaporation has a high contribution to total evaporation. Moreover, we predict that evaporative water loss rates are higher under the fed (hydrated) condition relative to fasted states. Metabolic and evaporation rates from the head and the body were measured separately in a partitioned chamber at 30°C. Birds were fed immediately prior to entry into respirometry, and the transition from fed to fasted states was recorded over a two-hour period. Total evaporative water loss nearly doubled in the fed versus fasted condition (1.75 ± 0.85 mg H2O/min and 0.97 ± 0.28 mg H2O/min, respectively), with respiratory water loss being significantly higher in the fed compared to the fasted condition (0.87 ± 0.38 mg H2O/min versus 0.42 ± 0.12 mg H2O/min, respectively). On the other hand, the body had a large, but non-significant, difference in evaporative water loss rates between the fed and fasted conditions (0.88 ± 0.55 mg H2O/min and 0.57 ± 0.17 mg H2O/min, respectively). Importantly, cutaneous evaporative water loss made up half of the total evaporation in both the fed and fasted conditions. These results demonstrate that evaporative water loss is modulated by hydration status at both the respiratory and cutaneous surfaces. Furthermore, cutaneous evaporation makes up approximately the same proportion of total evaporation to other birds at thermoneutral temperatures. Future work manipulating ambient temperature in conjunction with hydration status will further our understanding of the relative importance of cutaneous evaporation for these small-bodied birds. Funded by the Genentech Foundations Scholarship, CSUPERB New Investigator Grant, and San Francisco State University. This is the full abstract presented at the American Physiology Summit 2024 meeting and is only available in HTML format. There are no additional versions or additional content available for this abstract. Physiology was not involved in the peer review process.

Cutaneous Evaporative Water Loss in Lizards Changes Immediately with Temperature.

AbstractCutaneous evaporative water loss (CEWL) makes up a significant portion of total evaporative water loss in many terrestrial vertebrates. CEWL changes on evolutionary and acclimatory timescales in response to temperature and humidity. However, the lability of CEWL on acute timescales is unknown. To examine this, we increased or decreased body temperatures of western fence lizards (Sceloporus occidentalis) over a 15-min period while continuously recording CEWL with a handheld evaporimeter. CEWL increased in response to heating and decreased in response to cooling on the order of seconds. However, CEWL was different between heating and cooling groups at a common body temperature. We observed the same positive relationship between CEWL and body temperature, as well as the difference in CEWL between treatments, for deceased lizards that we opportunistically measured. However, deceased lizards had more extreme CEWL values for any given body temperature and treatment. Overall, our results suggest that both structural traits and active physiological processes likely influence the rates and plasticity of CEWL.

Hydric physiology and ecology of a federally endangered desert lizard.

Animals can respond to extreme climates by behaviourally avoiding it or by physiologically coping with it. We understand behavioural and physiological thermoregulation, but water balance has largely been neglected. Climate change includes both global warming and changes in precipitation regimes, so improving our understanding of organismal water balance is increasingly urgent. We assessed the hydric physiology of US federally endangered blunt-nosed leopard lizards (Gambelia sila) by measuring cutaneous evaporative water loss (CEWL), plasma osmolality and body condition. Measurements were taken throughout their active season, the short period of year when these lizards can be found aboveground. Compared to a more mesic species, G. sila had low CEWL which is potentially desert-adaptive, and high plasma osmolality that could be indicative of dehydration. We hypothesized that throughout the G. sila active season, as their habitat got hotter and drier, G. sila would become more dehydrated and watertight. Instead, CEWL and plasma osmolality showed minimal change for females and non-linear change for males, which we hypothesize is connected to sex-specific reproductive behaviours and changes in food availability. We also measured thermoregulation and microhabitat use, expecting that more dehydrated lizards would have lower body temperature, poorer thermoregulatory accuracy and spend less time aboveground. However, we found no effect of CEWL, plasma osmolality or body condition on these thermal and behavioural metrics. Finally, G. sila spends considerable time belowground in burrows, and burrows may serve not only as essential thermal refugia but also hydric refugia.

Hydration and evaporative water loss of lizards change in response to temperature and humidity acclimation.

Testing acclimation plasticity informs our understanding of organismal physiology and applies to conservation management amidst our rapidly changing climate. While there is a wealth of research on the plasticity of thermal and hydric physiology in response to temperature acclimation, there is a comparative gap for research on acclimation to different hydric regimes, as well as the interaction between water and temperature. We sought to fill this gap by acclimating Western Fence Lizards (Sceloporus occidentalis) to experimental climate conditions (crossed design of Hot or Cool, Dry or Humid) for eight days, and measuring cutaneous evaporative water loss (CEWL), plasma osmolality, hematocrit, and body mass before and after acclimation. CEWL changed plastically in response to the different climates, with lizards acclimated to Hot Humid conditions experiencing the greatest increase in CEWL. Change in CEWL among individuals was negatively related to treatment vapor pressure deficit and positively related to treatment water vapor pressure. Plasma osmolality, hematocrit, and body mass all showed greater changes in response to temperature than to humidity or vapor pressure deficit. CEWL and plasma osmolality were positively related across treatment groups before acclimation and within treatment groups after acclimation, but the two variables showed different responses to acclimation, suggesting that they are interrelated but governed by different mechanisms. This study is among few that assess more than one metric of hydric physiology and that test the interactive effects of temperature and humidity. Such measurements will be essential for predictive models of activity and survival for animals under climate change.

Evaporative water loss from dairy cows in climate-controlled respiration chambers

The effects of ambient temperature (AT) on total evaporative water loss from dairy cows at different relative humidity (RH) and air velocity (AV) levels were studied. Twenty Holstein dairy cows with an average parity of 2.0 ± 0.7 and body weight of 687 ± 46 kg participated in the study. Two climate-controlled respiration chambers were used. The experimental indoor climate was programmed to follow a diurnal pattern with AT at night being 9°C lower than during the day. Night AT was gradually increased from 7 to 21°C and day AT was increased from 16°C to 30°C within an 8-d period, both with an incremental change of 2°C/d. The effect of 3 RH levels with a diurnal pattern were studied as well, with low values during the day and high values during the night: low (day, 30%; night, 50%), medium (day, 45%; night, 70%), and high (day, 60%; night, 90%). The effects of AV were studied during the daytime at 3 levels: no fan (0.1 m/s), fan at medium speed (1.0 m/s), and fan at high speed (1.5 m/s). The medium and high AV levels were only combined with medium RH. In total, there were 5 treatments with 4 replicates each. The animals had free access to feed and water. Based on the water balance principle inside the respiration chambers, the total evaporative water loss from dairy cows at a daily level was quantified by measuring the mass of water in the incoming and outgoing air, condensed water, added water from a humidifier, and evaporative water from a wet floor, drinking bowl, manure reservoir, and water bucket. Water evaporation from a sample skin area was measured with a ventilated skin box, and water evaporation, through respiration with a face mask. The results show that RH/AV levels had no significant effect on total evaporative water loss, whereas the interaction effect between RH/AV with AT was significant. Cows at a high RH had a tendency for a lower increasing rate of evaporative water loss compared with cows at a low RH (0.61 vs. 0.79 kg/d per 1°C increase of AT). Cows at medium and high AV levels had a greater increasing rate than cows at low AV (0.91 and 0.95 vs. 0.71 kg/d per 1°C increase of AT, respectively). The increase of evaporative heat loss from dairy cows was mainly a result of the increase in evaporation (of sweat) from the skin. The skin water evaporation determined with the water balance method (less evaporation from respiration) and the ventilated skin box method showed no significant difference. The implication of this study is that cows at a high AT depend mainly on evaporative cooling from the skin. The ventilated skin box method, measuring only a small part of the skin during a short period during the day, can be a convenient and accurate way to determine the total cutaneous evaporative water loss from cows.

Heat tolerance, thermal equilibrium and environmental management strategies for dairy cows living in intertropical regions.

This review makes an attempt to characterize the physical attributes of heat tolerance, thermal equilibrium and thermal stress thresholds for dairy cows living in tropical environments, with a particular emphasis on pasture-based systems. Under such circumstances, the radiant heat load is the principal climatic factor that determines rates of heat and mass exchanges between cows and the environment. This fact may explain why simple mechanistic models based on air temperature and humidity are not adequately predicting thermal stress thresholds for cattle in tropical regions. To overcome this limitation, the Index of Thermal Stress for Cows (ITSC) and Index for the time spent in shade (ITS), which account for various sources of thermal radiation, were proposed to predict autonomous and behavioral thermoregulation of cows. Overall, the evolutionary adaptation of cattle in tropics favored animals that have cutaneous surface with a skin well protected against penetration of ultraviolet solar radiation (UV), covered by a coat surface with high thermal conductivity. For Holstein breed, although predominantly black animals absorb greater levels of short-wave solar radiation, they may present better protection of skin than white ones. However, dark-colored cows in tropical pastures have potential to absorb as much as 640 W m-2 of thermal radiation. This amount of heat load would require close to 1,300 g h -1 of cutaneous evaporative water loss through sweating to prevent increases to body temperature, where cows do not have access to shade. Cows are motivated to reduce time spent grazing and to seek shade when solar irradiance exceeds 550 W m-2, levels that in equatorial latitudes are likely to occur between 08:00 and 16:00h. This information may help producers improve the welfare of cows, as they can determine more comfortable hours for them to graze, for example, by employing nocturnal grazing. Over the daytime, cows should have access to areas with shade and this could include shade provided via solar panels, which has the potential to improve thermal comfort and sustainability of dairy production in tropical areas.

Climate aridity and habitat drive geographical variation in morphology and thermo-hydroregulation strategies of a widespread lizard species

Abstract Thermo-hydroregulation strategies involve concurrent changes in functional traits related to energy, water balance and thermoregulation and play a key role in determining life-history traits and population demography of terrestrial ectotherms. Local thermal and hydric conditions should be important drivers of the geographical variation of thermo-hydroregulation strategies, but we lack studies that examine these changes across climatic gradients in different habitat types. Here, we investigated intraspecific variation of morphology and thermo-hydroregulation traits in the widespread European common lizard (Zootoca vivipara louislantzi) across a multidimensional environmental gradient involving independent variation in air temperature and rainfall and differences in habitat features (access to free-standing water and forest cover). We sampled adult males for morphology, resting metabolic rate, total and cutaneous evaporative water loss and thermal preferences in 15 populations from the rear to the leading edge of the distribution across an elevational gradient ranging from sea level to 1750 m. Besides a decrease in adult body size with increasing environmental temperatures, we found little effect of thermal conditions on thermo-hydroregulation strategies. In particular, relict lowland populations from the warm rear edge showed no specific ecophysiological adaptations. Instead, body mass, body condition and resting metabolic rate were positively associated with a rainfall gradient, while forest cover and water access in the habitat throughout the season also influenced cutaneous evaporative water loss. Our study emphasizes the importance of rainfall and habitat features rather than thermal conditions for geographical variation in lizard morphology and physiology.

Cutaneous Evaporative Water Loss in Lizards is Variable across Body Regions and Plastic in Response to Humidity

As the climate crisis continues to alter temperature regimes and water availability, studying how animals regulate water balance grows in importance. We studied variation in water loss across the skin among body regions and its plasticity in response to humidity acclimation in Western Fence Lizards (Sceloporus occidentalis). We examined the effects of climatic and physiological variables on cutaneous evaporative water loss (CEWL) and compared CEWL rates among body regions (dorsum, ventrum, head, dewlap, and mite patch). The best model to explain baseline variation in log-transformed CEWL included: body region; lizard mass; ambient temperature, vapor pressure deficit (VPD), and solar radiation at the time of capture; body temperature, ambient temperature, and VPD at the time of CEWL measurement; and the interactions between body region and each of mass and VPD at the time of capture. These results demonstrate cutaneous osmoregulatory variability among lizards based on climate and body size and within individual lizards based on body region. We also tested CEWL plasticity in response to humidity acclimation. Lizards exposed to humid conditions for 8 d exhibited increased CEWL, and lizards exposed to dry conditions exhibited decreased CEWL compared to initial measurements. This is evidence for rapid and significant acclimatory responses of CEWL in response to changes in environmental humidity. Such variation and plasticity suggest that lizards possess a certain degree of ability for using osmoregulatory changes to respond to climate change.

Small Alpine Marsupials Regulate Evaporative Water Loss, Suggesting a Thermoregulatory Role Rather than a Water Conservation Role.

AbstractWe show here that evaporative water loss (EWL) is constant over a wide range of ambient relative humidity for two species of small, mesic habitat dasyurid marsupials (Antechinus agilis and Antechinus swainsonii) below thermoneutrality (20°C) and within thermoneutrality (30°C). This independence of EWL from the water vapor pressure deficit between the animal and its environment indicates that EWL is physiologically controlled by both species. The magnitude of this control of EWL was similar to that of two other small marsupials from more arid habitats, which combined with the observation that there were no effects of relative humidity on body temperature or metabolic rate, suggests that control of EWL is a consequence of precise thermoregulation to maintain heat balance rather than a water-conserving strategy at low relative humidities. The antechinus appear to manipulate cutaneous EWL rather than respiratory EWL to control their total EWL by modifying their cutaneous resistance and/or skin temperature. We propose that there is a continuum between enhanced thermoregulatory EWL at high ambient temperature and so-called insensible EWL at and below thermoneutrality.

Reliability of methods to determine cutaneous evaporative water loss rate in furred and fleeced mammals.

We used a high-precision weighing system and flow-through respirometry to quantify cutaneous evaporative water loss rates in woolly sheep (wool thickness, ca. 6.5 cm) and haired goats (coat thickness, ca. 2.5 cm), while simultaneously recording parallel data obtained from (1) a flow-through ventilated capsule, (2) a closed hand-held electronic evaporimeter chamber, and (3) a closed colorimetric paper disc chamber. In comparison to the weighing system and respirometry, used here as a "gold standard" measure of cutaneous evaporative water loss rate, we found relatively good agreement with data obtained from the flow-through ventilated capsules. However, we found poor agreement with data obtained from the closed electronic evaporimeter chambers (underestimated by 60%, on average) and the closed colorimetric paper disc chambers (overestimated by 52%, on average). This deviation was likely associated with a requirement for shaved skin in the closed chamber methods. Our results therefore cast doubt on the validity of the closed chamber methods for measurement of cutaneous evaporative water loss rates in furred and fleeced mammals, and instead show that more accurate values can be obtained using flow-through ventilated capsules.

The thermoregulatory role of relative bill and leg surface areas in a Mediterranean population of Great tit (Parus major).

There is growing evidence on the role of legs and bill as ‘thermal windows’ in birds coping with heat stress. However, there is a lack of empirical work examining the relationship between the relative bill and/or leg surface areas and key thermoregulatory traits such as the limits of the thermoneutral zone (TNZ) or the cooling efficiency at high temperatures. Here, we explored this relationship in a Mediterranean population of Great tit (Parus major) facing increasing thermal stress in its environment. The lower and upper critical limits of the TNZ were found to be 17.7 ± 1.6ºC and 34.5 ± 0.7°C, respectively, and the basal metabolic rate was 0.96 ± 0.12 ml O2 min−1 on average. The evaporative water loss (EWL) inflection point was established at 31.85 ± 0.27°C and was not significantly different from the value of the upper critical limit. No significant relationship was observed between the relative bill or tarsi size and TNZ critical limits, breadth, mass‐independent VO2, or mass‐independent EWL at any environmental temperature (from 10 to 40°C). However, Great tit males (but not females) with larger tarsi areas (a proxy of leg surface area) showed higher cooling efficiencies at 40°C. We found no support for the hypothesis that the bill surface area plays a significant role as a thermal window in Great tits, but the leg surface areas may play a role in males’ physiological responses to high temperatures. On the one hand, we argue that the studied population occupies habitats with available microclimates and fresh water for drinking during summer, so active heat dissipation by EWL might be favored instead of dry heat loss through the bill surface. Conversely, male dominance behaviors could imply a greater dependence on cutaneous EWL through the upper leg surfaces as a consequence of higher exposure to harsh environmental conditions than faced by females.

Lipid composition of the stratum corneum in different regions of the body of Kuhl's pipistrelle from the Negev Desert, Israel

The most superficial epidermal layer in endotherms is the stratum corneum (SC), which is composed of dead corneocytes embedded in a lipid matrix with free fatty acids, cholesterol, ceramides, and cerebrosides; the lipid composition of the SC determines its permeability to water vapor. Lipids that are more polar, have longer hydrocarbon chains, and are less bulky are often packed in more ordered phase states to slow cutaneous evaporative water loss (CEWL); these lipids also resist transitions to more disordered phases at high ambient temperatures (Ta). In bats, wing and tail membranes (wing patagia and tail uropatagium, respectively) allow powered flight, but increase surface area, and hence CEWL, with implications for survival in arid environments. We captured Pipistrellus kuhlii from an arid habitat and measured the lipid composition of the SC of the plagiopatagium in the wing, the uropatagium, and the non-membranous region (NMR) of the body using thin layer chromatography and reversed phase high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. The patagia contained more cholesterol and shorter-chained ceramides, and fewer cerebrosides than the NMR, indicating that the lipid phase transition temperature in the patagia is lower than in the NMR. Thus, at moderate Ta the lipids in the SC in all body regions will remain in an ordered phase state, allowing water conservation; but as Ta increases, the lipids in the SC of the patagia will more easily transition into a disordered phase, resulting in increased CEWL from the patagia facilitating efficient heat dissipation in hot environments.

A worldwide and annotated database of evaporative water loss rates in squamate reptiles

AbstractMotivationThe understanding of physiological adaptations, of evolutionary radiations and of ecological responses to global change urges for global, comprehensive databases of the functional traits of extant organisms. The ability to maintain an adequate water balance is a critical functional property influencing the resilience of animal species to climate variation. In terrestrial or semi‐terrestrial organisms, total water loss includes a significant contribution from evaporative water loss (EWL). The analysis of geographic and phylogenetic variation in EWL rates must however account for differences in methods and potential confounding factors, which influence standard measures of whole‐organism water loss. We compiled the global and standardized SquamEWL database of total, respiratory and cutaneous EWL for 325 species and subspecies of squamate reptiles (793 samples and 2,536 estimates) from across the globe. An extensive set of companion data and annotations associated with the EWL measurements of potential value for future investigation, including metabolic rate data, is provided. We present preliminary descriptive statistics for the compiled data, discuss gaps and biases, and identify promising avenues to update, expand and explore this database.Main types of variables containedStandard water loss rates, geographic data, metabolic rates.Spatial locationGlobal.Time periodData were obtained from extant species and were collected between 1945 and 2020.Major taxaReptilia, Squamata including lizards, snakes and amphisbaenians.Level of measurementsIndividual samples of animals from the same species, locality, age class and sex category.Software formatcsv.

Bearded dragons (Pogona vitticeps) with reduced scalation lose water faster but do not have substantially different thermal preferences.

Whether scales reduce cutaneous evaporative water loss in lepidosaur reptiles (Superorder Lepidosauria) such as lizards and snakes has been a contentious issue for nearly half a century. Furthermore, while many studies have looked at whether dehydration affects thermal preference in lepidosaurs, far fewer have examined whether normally hydrated lepidosaurs can assess their instantaneous rate of evaporative water loss and adjust their thermal preference to compensate in an adaptive manner. We tested both of these hypotheses using three captive-bred phenotypes of bearded dragon (Pogona vitticeps) sourced from the pet trade: 'wild-types' with normal scalation, 'leatherbacks' exhibiting scales of reduced prominence, and scaleless bearded dragons referred to as 'silkbacks'. Silkbacks on average lost water evaporatively at about twice the rate that wild-types did. Leatherbacks on average were closer in their rates of evaporative water loss to silkbacks than they were to wild-types. Additionally, very small (at most ∼1°C) differences in thermal preference existed between the three phenotypes that were not statistically significant. This suggests a lack of plasticity in thermal preference in response to an increase in the rate of evaporative water loss, and may be reflective of a thermal 'strategy' as employed by thermoregulating bearded dragons that prioritises immediate thermal benefits over the threat of future dehydration. The results of this study bolster an often-discounted hypothesis regarding the present adaptive function of scales and have implications for the applied fields of animal welfare and conservation.

A mechanistic model to scale up biophysical processes into geographical size gradients in ectotherms

AbstractAimUnderstanding the mechanisms behind broad‐scale gradients in animal body size remains challenging. In contrast to endotherms, the extent to which biophysical properties of ectotherms scale up into interspecific size gradients at macroecological scales remains elusive. Our aim here is to investigate what latitudinal body size gradients are expected to emerge from first biophysical principles regarding heat and water balances in terrestrial vertebrate ectotherms.LocationNearctic and Western Palaearctic.Time periodTheoretical model under current climatic conditions.Major taxa studiedTerrestrial vertebrate ectotherms.MethodsWe developed a biophysical model to derive heat balances and rates of water loss of lizard‐like reptiles and anuran‐like amphibians from microclimatic variables. We projected geographically model predictions for different size classes to explore how body temperature (Tb) and cutaneous evaporative water loss (EWL) are influenced by body size at macroscales.ResultsLarge‐bodied ectotherms (over 100 g) attain a slightly higher Tb and conserve it for longer than small animals (5 g), owing to the lower convective heat dissipation and their greater thermal inertia. While large animals are more at risk of overheating in warm regions, all organisms, irrespective of body size, are constrained by low temperatures experienced at boreal latitudes. Furthermore, because of the nonlinear decrease in EWL with increasing size, larger size allows reduction of water loss of small wet‐skinned ectotherms in more evaporative environments.Main conclusionsSize differences in heat balance offer a mechanistic explanation for the pattern of increasing body size northwards, as well as for the associated decrease in species richness. Water conservation is also identified as a potential mechanism for larger body sizes in dryer areas in wet‐skinned ectotherms such as amphibians. Moreover, our model reveals nonlinear relationships involving size and optimum states of thermal and hydric functions, accounting for complexities that are unnoticeable by traditional, correlative investigations.

Temperature and dehydration effects on metabolism, water uptake and the partitioning between respiratory and cutaneous evaporative water loss in a terrestrial toad.

Terrestrial anurans often experience fluctuations in body temperature and hydration state, which are known to influence evaporative water loss through the skin (EWLSkin) and lungs (EWLResp). These effects arise from associated changes in skin permeability, metabolism and lung ventilation. Herein, we determined the rates of EWLSkin and EWLResp in the terrestrial toad Rhinella diptycha at different temperatures and hydration states. We measured oxygen uptake rates to verify whether alterations in the partitioning between EWLSkin and EWLResp were associated with metabolism-induced changes in pulmonary gas exchange. We also measured the influence of hydration and temperature on water uptake (WU) through the skin. Finally, as estimates of skin resistance to evaporation (Rs) are usually inferred from total evaporative water loss (EWLTotal), under the assumption of negligible EWLResp, we calculated the potential error in accepting this assumption for different temperature and hydration states. EWLSkin and EWLResp increased with temperature, but this response was greater for EWLResp, which was attributed to the temperature-induced elevation in metabolism and lung ventilation. Dehydration caused a decrease in the relative contribution of EWLSkin to EWLTotal, mirrored by the concurrent increase in the contribution of EWLResp, at all temperatures. Thus, Rs increased with dehydration. WU rates were dictated by dehydration with little influence of temperature. The partitioning between EWLSkin and EWLResp was affected by both temperature and hydration state and, under some conditions, considering EWLResp as negligible led to significant errors in the assessment of skin resistance to evaporation.

Avian thermoregulation in the heat: evaporative cooling capacity of arid-zone Caprimulgiformes from two continents.

Birds in the order Caprimulgiformes (nightjars and allies) have a remarkable capacity for thermoregulation over a wide range of environmental temperatures, exhibiting pronounced heterothermy in cool conditions and extreme heat tolerance at high environmental temperatures. We measured thermoregulatory responses to acute heat stress in three species of Caprimulgiformes that nest in areas of extreme heat and aridity, the common poorwill (Phalaenoptilus nuttallii: Caprimulgidae) and lesser nighthawk (Chordeiles acutipennis: Caprimulgidae) in the Sonoran Desert of Arizona, and the Australian owlet-nightjar (Aegotheles cristatus: Aegothelidae) in the mallee woodlands of South Australia. We exposed wild-caught birds to progressively increasing air temperatures (Ta) and measured resting metabolic rate (RMR), evaporative water loss (EWL), body temperature (Tb) and heat tolerance limit (HTL; the maximum Ta reached). Comparatively low RMR values were observed in all species (0.35, 0.36 and 0.40 W for the poorwill, nighthawk and owlet-nightjar, respectively), with Tb approximating Ta at 40°C and mild hyperthermia occurring as Ta reached the HTL. Nighthawks and poorwills reached HTLs of 60 and 62°C, respectively, whereas the owlet-nightjar had a HTL of 52°C. RMR increased gradually above minima at Ta of 42, 42 and 35°C, and reached 1.7, 1.9 and 2.0 times minimum resting values at HTLs in the poorwill, nighthawk and owlet-nightjar, respectively. EWL increased rapidly and linearly as Ta exceeded Tb and resulted in maximum rates of evaporative heat dissipation equivalent to 237-424% of metabolic heat production. Bouts of gular flutter resulted in large transient increases in evaporative heat loss (50-123%) accompanied by only small increments in RMR (<5%). The cavity-nesting/roosting owlet-nightjar had a lower HTL and less efficient evaporative cooling compared with the species that nest and/or roost on open desert surfaces. The high efficiency of gular flutter for evaporative cooling, combined with mild hyperthermia, provides the physiological basis for defending Tb well below Ta in extreme heat and is comparable to the efficient cooling observed in arid-zone columbids in which cutaneous EWL is the predominant cooling pathway.

The cutaneous lipid composition of bat wing and tail membranes: a case of convergent evolution with birds.

The water vapour permeability barrier of mammals and birds resides in the stratum corneum (SC), the outermost layer of the epidermis. The molar ratio and molecular arrangement of lipid classes in the SC determine the integrity of this barrier. Increased chain length and polarity of ceramides, the most abundant lipid class in mammalian SC, contribute to tighter packing and thus to reduced cutaneous evaporative water loss (CEWL). However, tighter lipid packing also causes low SC hydration, making it brittle, whereas high hydration softens the skin at the cost of increasing CEWL. Cerebrosides are not present in the mammalian SC; their pathological accumulation occurs in Gaucher's disease, which leads to a dramatic increase in CEWL. However, cerebrosides occur normally in the SC of birds. We tested the hypothesis that cerebrosides are also present in the SC of bats, because they are probably necessary to confer pliability to the skin, a quality needed for flight. We examined the SC lipid composition of four sympatric bat species and found that, as in birds, their SC has substantial cerebroside contents, not associated with a pathological state, indicating convergent evolution between bats and birds.

Metabolic rate, evaporative water loss and thermoregulatory state in four species of bats in the Negev desert

Life in deserts is challenging for bats because of their relatively high energy and water requirements; nevertheless bats thrive in desert environments. We postulated that bats from desert environments have lower metabolic rates (MR) and total evaporative water loss (TEWL) than their mesic counterparts. To test this idea, we measured MR and TEWL of four species of bats, which inhabit the Negev desert in Israel, one species mainly restricted to hyper-arid deserts (Otonycteris hemprichii), two species from semi-desert areas (Eptesicus bottae and Plecotus christii), and one widespread species (Pipistrellus kuhlii). We also measured separately, in the same individuals, the two components of TEWL, respiratory water loss (RWL) and cutaneous evaporative water loss (CEWL), using a mask. In all the species, MR and TEWL were significantly reduced during torpor, the latter being a consequence of reductions in both RWL and CEWL. Then, we evaluated whether MR and TEWL in bats differ according to their geographic distributions, and whether those rates change with Ta and the use of torpor. We did not find significant differences in MR among species, but we found that TEWL was lowest in the species restricted to desert habitats, intermediate in the semi-desert dwelling species, and highest in the widespread species, perhaps a consequence of adaptation to life in deserts. Our results were supported by a subsequent analysis of data collected from the literature on rates of TEWL for 35 bat species from desert and mesic habitats.

Thermal and moisture habitat preferences do not maximize jumping performance in frogs

Summary Amphibians are suffering population declines globally, and understanding how environmental parameters influence their thermal and moisture preferences and performance at various tasks is crucial to understanding how these animals will be influenced by climate change. Body temperature and hydration affect organismal performance at many fitness‐related tasks. Since amphibians are ectotherms with highly water‐permeable skin, environmental temperature and moisture directly affect their body temperature and hydration. Therefore, amphibians should select habitats with the optimal combination of temperature and moisture to perform tasks necessary for survival. However, interactions between environmental temperature and moisture can influence habitat selection and task performance in different and often unpredictable ways, and this has only infrequently been considered. We tested for interactions between environmental temperature, moisture and organismal hydration on temperature and moisture preferences and jumping performance in green frogs (Lithobates clamitans) in the laboratory, using thermal and moisture gradients, and high‐speed video and force plate data. We then integrated the laboratory experiments with field data. In the thermal and moisture gradients, frogs selected environmental conditions that minimized cutaneous evaporative water loss, hydroregulating more stringently than thermoregulating. These results are consistent with frogs in the field, which had highly variable body temperatures, but were always hydrated above 95% of their standard mass. However, conditions that minimized evaporative water loss frequently did not maximize jumping performance because warmer temperatures conferred greater performance. The ecology of L. clamitans may explain the discrepancy between their preferences and jumping performance optima because the frogs remain in wet environments that serve as refuges from dehydration. In parts of their range where frogs are subjected to warmer and drier conditions, they are likely to select microhabitats that minimize the risk of dehydration, possibly at the expense of their ability to forage and escape from predators.