Adaptive variation in the upper limits of avian body temperature.

Physiological performance declines precipitously at high body temperature (Tb), but little attention has been paid to adaptive variation in upper Tb limits among endotherms. We hypothesized that avian maximum tolerable Tb (Tbmax) has evolved in response to climate, with higher Tbmax in species exposed to high environmental heat loads or humidity-related constraints on evaporative heat dissipation. To test this hypothesis, we compared Tbmax and related variables among 53 bird species at multiple sites in South Africa with differing maximum air temperature (Tair) and humidity using a phylogenetically informed comparative framework. Birds in humid, lowland habitats had comparatively high Tbmax (mean ± SD = 45.60 ± 0.58 °C) and low normothermic Tb (Tbnorm), with a significantly greater capacity for hyperthermia (Tbmax - Tbnorm gradient = 5.84 ± 0.77 °C) compared with birds occupying cool montane (4.97 ± 0.99 °C) or hot arid (4.11 ± 0.84 °C) climates. Unexpectedly, Tbmax was significantly lower among desert birds (44.65 ± 0.60 °C), a surprising result in light of the functional importance of hyperthermia for water conservation. Our data reveal a macrophysiological pattern and support recent arguments that endotherms have evolved thermal generalization versus specialization analogous to the continuum among ectothermic animals. Specifically, a combination of modest hyperthermia tolerance and efficient evaporative cooling in desert birds is indicative of thermal specialization, whereas greater hyperthermia tolerance and less efficient evaporative cooling among species in humid lowland habitats suggest thermal generalization.

Sub-lethal impacts of lead poisoning on blood biochemistry, immune function and delta-aminolevulinic acid dehydratase (δ-ALAD) activity in Cape (Gyps coprotheres) and white-backed (G. africanus) Vulture chicks.

Although the prevalence of lead poisoning in southern Africa's Gyps vultures is now well-established, its finer physiological effects on these endangered species remain poorly characterised. We evaluated the sub-lethal impact of acute lead exposure on Cape and White-backed Vulture chicks from two breeding colonies in South Africa, by analysing its possible effects on key blood biochemistry parameters, immune function, packed cell volume and δ-aminolevulinic acid dehydratase (δ-ALAD) activity. All 37 White-backed Vulture nestlings sampled displayed elevated lead levels (>10 µg/dL), and seven had blood [Pb] >100 µg/dL. Eight of 28 Cape Vulture nestlings sampled had blood [Pb] exceeding background exposure, with one showing blood [Pb] >100 µg/dL. Delta-aminolevulinic acid dehydratase (δ-ALAD) activity was significantly and negatively related to blood [Pb] in nestlings from both species, with 50% inhibition of the enzyme predicted to occur at blood [Pb] = 52.8 µg/dL (White-backed Vulture) and 18.8 µg/dL (Cape Vulture). Although no significant relationship was found between % packed cell volume (PCV) and blood [Pb], the relatively lower mean PCV of 32.9% in White-backed Vulture chicks, combined with normal serum protein values, is likely indicative of depression or haemolytic anaemia. The leukogram was consistent in both species, although the presence of immature heterophils suggested an inflammatory response in White-backed Vulture chicks with blood [Pb] >100 µg/dL. Values for cholesterol, triglycerides, total serum protein, albumin, globulin, albumin/globulin ratio, alanine aminotransferase (ALT) and gamma-glutamyl transferase (GGT) were consistent with values previously reported. Calcium and phosphorus concentrations suggested no adverse effects on bone metabolism. A significant decrease in urea: uric acid (U:UA) ratio at blood [Pb] >100 µg/dL in White-backed Vulture chicks, brought about by a decrease in urea production, raises the possibility of hepatic abnormality. These results suggest that δ-ALAD activity may serve as a sensitive biomarker of lead toxicity in both species, while highlighting the need to better understand the significant variability in sensitivity that is observed, even between closely related members of the same genus.

Reporting guidelines for terrestrial respirometry: Building openness, transparency of metabolic rate and evaporative water loss data.

Respirometry is an important tool for understanding whole-animal energy and water balance in relation to the environment. Consequently, the growing number of studies using respirometry over the last decade warrants reliable reporting and data sharing for effective dissemination and research synthesis. We provide a checklist guideline on five key sections to facilitate the transparency, reproducibility, and replicability of respirometry studies: 1) materials, set up, plumbing, 2) subject conditions/maintenance, 3) measurement conditions, 4) data processing, and 5) data reporting and statistics, each with explanations and example studies. Transparency in reporting and data availability has benefits on multiple fronts. Authors can use this checklist to design and report on their study, and reviewers and editors can use the checklist to assess the reporting quality of the manuscripts they review. Improved standards for reporting will enhance the value of primary studies and will greatly facilitate the ability to carry out higher quality research syntheses to address ecological and evolutionary theories.

An evaluation of a biophysical model for predicting avian thermoregulation in the heat.

Survival and reproduction of endotherms depend on their ability to balance energy and water exchange with their environment, avoiding lethal deficits and maximising gains for growth and reproduction. At high environmental temperatures, diurnal endotherms maintain body temperature (Tb) below lethal limits via physiological and behavioural adjustments. Accurate models of these processes are crucial for predicting effects of climate variability on avifauna. We evaluated the performance of a biophysical model (NicheMapR) for predicting evaporative water loss (EWL), resting metabolic rate (RMR) and Tb at environmental temperatures approaching or exceeding normothermic Tb for three arid-zone birds: southern yellow-billed hornbill (Tockus leucomelas), southern pied babbler (Turdoides bicolor) and southern fiscal (Lanius collaris). We simulated metabolic chamber conditions and compared model outputs with thermal physiology data collected at air temperatures (Tair) between 10 and 50°C. Additionally, we determined the minimum data needed to accurately model diurnal birds' thermoregulatory responses to Tair using sensitivity analyses. Predicted EWL, metabolic rate and Tb corresponded tightly with observed values across Tair, with only minor discrepancies for EWL in two species at Tair≈35°C. Importantly, the model captured responses at Tair=30-40°C, a range spanning threshold values for sublethal fitness costs associated with sustained hot weather in arid-zone birds. Our findings confirm how taxon-specific parameters together with biologically relevant morphological data can accurately model avian thermoregulatory responses to heat. Biophysical models can be used as a non-invasive way to predict species' sensitivity to climate, accounting for organismal (e.g. physiology) and environmental factors (e.g. microclimates).

Respirometry protocols for avian thermoregulation at high air temperatures: stepped and steady-state profiles yield similar results.

Relationships between air temperature (Tair) and avian body temperature (Tb), resting metabolic rate (RMR) and evaporative water loss (EWL) during acute heat exposure can be quantified through respirometry using several approaches. One involves birds exposed to a stepped series of progressively increasing Tair setpoints for short periods (<20-30 min), whereas a second seeks to achieve steady-state conditions by exposing birds to a single Tair for longer periods (>1-2 h). To compare these two approaches, we measured Tb, RMR and EWL over Tair=28°C to 44°C in the dark-capped bulbul (Pycnonotus tricolor). The two protocols yielded indistinguishable values of Tb, RMR and EWL and related variables at most Tair values, revealing that both are appropriate for quantifying avian thermal physiology during heat exposure over the range of Tair in the present study. The stepped protocol, however, has several ethical and practical advantages.

Chronic, sublethal effects of high temperatures will cause severe declines in southern African arid-zone birds during the 21st century.

Birds inhabiting hot, arid regions are among the terrestrial organisms most vulnerable to climate change. The potential for increasingly frequent and intense heat waves to cause lethal dehydration and hyperthermia is well documented, but the consequences of sublethal fitness costs associated with chronic exposure to sustained hot weather remain unclear. Using data for species occurring in southern Africa's Kalahari Desert, we mapped exposure to acute lethal risks and chronic sublethal fitness costs under past, present, and future climates. For inactive birds in shaded microsites, the risks of lethal dehydration and hyperthermia will remain low during the 21st century. In contrast, exposure to conditions associated with chronic, sublethal costs related to progressive body mass loss, reduced nestling growth rates, or increased breeding failure will expand dramatically. For example, by the 2080s the region will experience 10-20 consecutive days per year on which Southern Pied Babblers (Turdoides bicolor) will lose ∼4% of body mass per day, conditions under which this species' persistence will be extremely unlikely. Similarly, exposure to air temperature maxima associated with delayed fledging, reduced fledgling size, and breeding failure will increase several-fold in Southern Yellow-billed Hornbills (Tockus leucomelas) and Southern Fiscals (Lanius collaris). Our analysis reveals that sublethal costs of chronic heat exposure are likely to drive large declines in avian diversity in the southern African arid zone by the end of the century.

Thermoregulation in desert birds: scaling and phylogenetic variation in heat tolerance and evaporative cooling.

Evaporative heat dissipation is a key aspect of avian thermoregulation in hot environments. We quantified variation in avian thermoregulatory performance at high air temperatures (Ta) using published data on body temperature (Tb), evaporative water loss (EWL) and resting metabolic rate (RMR) measured under standardized conditions of very low humidity in 56 arid-zone species. Maximum Tb during acute heat exposure varied from 42.5±1.3°C in caprimulgids to 44.5±0.5°C in passerines. Among passerines, both maximum Tb and the difference between maximum and normothermic Tb decreased significantly with body mass (Mb). Scaling exponents for minimum thermoneutral EWL and maximum EWL were 0.825 and 0.801, respectively, even though evaporative scope (ratio of maximum to minimum EWL) varied widely among species. Upper critical limits of thermoneutrality (Tuc) varied by >20°C and maximum RMR during acute heat exposure scaled to Mb0.75 in both the overall data set and among passerines. The slope of RMR at Ta>Tuc increased significantly with Mb but was substantially higher among passerines, which rely on panting, compared with columbids, in which cutaneous evaporation predominates. Our analysis supports recent arguments that interspecific within-taxon variation in heat tolerance is functionally linked to evaporative scope and maximum ratios of evaporative heat loss (EHL) to metabolic heat production (MHP). We provide predictive equations for most variables related to avian heat tolerance. Metabolic costs of heat dissipation pathways, rather than capacity to increase EWL above baseline levels, appear to represent the major constraint on the upper limits of avian heat tolerance.

Hot days are associated with short-term adrenocortical responses in a southern African arid-zone passerine bird.

Relatively little effort has been directed towards elucidating the role of physiological stress pathways in mediating avian responses to global heating. For free-ranging southern pied babblers, Turdoides bicolor, daily maximum air temperatures (Tmax) between ∼35 and ∼40°C result in reduced foraging efficiency, loss of body mass and compromised breeding success. We tested the hypothesis that very hot days are experienced as stressors by quantifying relationships between Tmax and faecal glucocorticoid metabolite (fGCM) levels in naturally excreted droppings. On days when Tmax<38°C, fGCM levels were independent of Tmax (mean±s.d. 140.25±56.92 ng g-1 dry mass). At Tmax>38°C, however, fGCM levels increased linearly with Tmax and averaged 190.79±70.13 ng g-1 dry mass. The effects of Tmax on fGCM levels did not carry over to the following morning, suggesting that very hot days are experienced as acute stressors.

How hornbills handle heat: sex-specific thermoregulation in the southern yellow-billed hornbill.

At a global scale, thermal physiology is correlated with climatic variables such as temperature and aridity. There is also evidence that thermoregulatory traits vary with fine-scale microclimate, but this has received less attention in endotherms. Here, we test the hypothesis that avian thermoregulation varies with microclimate and behavioural constraints in a non-passerine bird. Male and female southern yellow-billed hornbills (Tockus leucomelas) experience markedly different microclimates while breeding, with the female sealing herself into a tree cavity and moulting all her flight feathers during the breeding attempt, becoming entirely reliant on the male for provisioning. We examined interactions between resting metabolic rate (RMR), evaporative water loss (EWL) and core body temperature (Tb) at air temperatures (Ta) between 30°C and 52°C in male and female hornbills, and quantified evaporative cooling efficiencies and heat tolerance limits. At thermoneutral Ta, neither RMR, EWL nor Tb differed between sexes. At Ta >40°C, however, RMR and EWL of females were significantly lower than those of males, by ∼13% and ∼17%, respectively, despite similar relationships between Tb and Ta, maximum ratio of evaporative heat loss to metabolic heat production and heat tolerance limits (∼50°C). These sex-specific differences in hornbill thermoregulation support the hypothesis that avian thermal physiology can vary within species in response to fine-scale microclimatic factors. In addition, Q10 for RMR varied substantially, with Q10 ≤2 in some individuals, supporting recent arguments that active metabolic suppression may be an underappreciated aspect of endotherm thermoregulation in the heat.

The Physiology of Heat Tolerance in Small Endotherms.

Understanding the heat tolerances of small mammals and birds has taken on new urgency with the advent of climate change. Here, we review heat tolerance limits, pathways of evaporative heat dissipation that permit the defense of body temperature during heat exposure, and mechanisms operating at tissue, cellular, and molecular levels.

Phenotypic flexibility of metabolic rate and evaporative water loss does not vary across a climatic gradient in an Afrotropical passerine bird.

Small birds inhabiting northern temperate and boreal latitudes typically increase metabolic rates during cold winters or acclimation to low air temperatures (Taccl). Recent studies suggest considerable variation in patterns of seasonal metabolic acclimatization in birds from subtropical and tropical regions with milder winters, but there remains a dearth of acclimation studies investigating metabolic flexibility among lower-latitude birds. We used short-term thermal acclimation experiments to investigate phenotypic flexibility in basal metabolic rate (BMR), thermoneutral evaporative water loss (EWL) and summit metabolism (Msum) in three populations of white-browed sparrow-weavers (Plocepasser mahali) along a climatic and aridity gradient. We allocated individuals to one of three Taccl treatments (5, 20 and 35°C; n=11 per population per Taccl) for 28 days, and measured post-acclimation BMR, EWL and Msum using flow-through respirometry. Our data reveal the expected pattern of lower BMR and EWL (∼12% and 25% lower, respectively) in birds at Taccl=35°C compared with cooler Taccl treatments, as observed in previous acclimation studies on subtropical birds. We found no variation in the reaction norms of BMR and EWL among populations in response to acclimation, suggesting previously documented differences in seasonal BMR acclimatization are the result of phenotypic flexibility. In contrast to higher-latitude species, Msum did not significantly vary in response to thermal acclimation. These findings support the idea that factors other than enhancing cold tolerance may be driving patterns of metabolic variation in subtropical birds.

Extreme and variable torpor among high-elevation Andean hummingbird species.

Torpor is thought to be particularly important for small endotherms occupying cold environments and with limited fat reserves to fuel metabolism, yet among birds deep torpor is both rare and variable in extent. We investigated torpor in hummingbirds at approximately 3800 m.a.s.l. in the tropical Andes by monitoring body temperature (Tb) in 26 individuals of six species held captive overnight and experiencing natural air temperature (Ta) patterns. All species used pronounced torpor, with one Metallura phoebe reaching a minimum Tb of 3.26°C, the lowest yet reported for any bird or non-hibernating mammal. The extent and duration of torpor varied among species, with overnight body mass (Mb) loss negatively correlated with both minimum Tb and bout duration. We found a significant phylogenetic signal for minimum Tb and overnight Mb loss, consistent with evolutionarily conserved thermoregulatory traits. Our findings suggest deep torpor is routine for high Andean hummingbirds, but evolved species differences affect its depth.

Mapping evaporative water loss in desert passerines reveals an expanding threat of lethal dehydration.

Extreme high environmental temperatures produce a variety of consequences for wildlife, including mass die-offs. Heat waves are increasing in frequency, intensity, and extent, and are projected to increase further under climate change. However, the spatial and temporal dynamics of die-off risk are poorly understood. Here, we examine the effects of heat waves on evaporative water loss (EWL) and survival in five desert passerine birds across the southwestern United States using a combination of physiological data, mechanistically informed models, and hourly geospatial temperature data. We ask how rates of EWL vary with temperature across species; how frequently, over what areas, and how rapidly lethal dehydration occurs; how EWL and die-off risk vary with body mass; and how die-off risk is affected by climate warming. We find that smaller-bodied passerines are subject to higher rates of mass-specific EWL than larger-bodied counterparts and thus encounter potentially lethal conditions much more frequently, over shorter daily intervals, and over larger geographic areas. Warming by 4 °C greatly expands the extent, frequency, and intensity of dehydration risk, and introduces new threats for larger passerine birds, particularly those with limited geographic ranges. Our models reveal that increasing air temperatures and heat wave occurrence will potentially have important impacts on the water balance, daily activity, and geographic distribution of arid-zone birds. Impacts may be exacerbated by chronic effects and interactions with other environmental changes. This work underscores the importance of acute risks of high temperatures, particularly for small-bodied species, and suggests conservation of thermal refugia and water sources.

Non-invasive monitoring of physiological stress in an afrotropical arid-zone passerine bird, the southern pied babbler.

Using faecal matter to monitor stress levels in animals non-invasively is a powerful technique for elucidating the effects of biotic and abiotic stressors on free-living animals. To validate the use of droppings for measuring stress in southern pied babblers (Turdoides bicolor) we performed an ACTH challenge on captive individuals and determined the effect of temporary separation from their social group on their faecal glucocorticoid metabolite (fGCM) concentration. Additionally, we compared fGCM concentrations of captive babblers to those of wild conspecifics and examined the effects of dominance rank on fGCM concentration. We found droppings to be a suitable matrix for measuring physiological stress in babblers and that individual separation from the group caused an increase in fGCM levels. In addition, babblers temporarily held in captivity had substantially higher fGCM concentrations than wild individuals, indicating that babblers kept in captivity experience high levels of stress. In wild, free-living individuals, dominant males showed the highest levels of stress, suggesting that being the dominant male of a highly territorial social group is stressful. Non-invasive sampling allows field-based researchers to reduce disturbance related to monitoring adrenocortical function, thereby avoiding artificially increasing circulating corticosterone concentration as it is not necessary to physically restrain study animals.

Reaction norms for heat tolerance and evaporative cooling capacity do not vary across a climatic gradient in a passerine bird.

There is increasing evidence for considerable phenotypic flexibility in endotherm thermal physiology, a phenomenon with far-reaching implications for the evolution of traits related to heat tolerance. Numerous studies have documented intraspecific variation in avian thermoregulatory traits, but few have revealed the shapes of thermoregulatory reaction norms or how these might vary among populations. We investigated phenotypic flexibility in the ability of a model Afrotropical passerine bird (the white-browed sparrow-weaver, Plocepasser mahali) to handle high air temperatures (Ta). We allocated birds from three sites varying by ~ 11 °C in mean daily summer maximum Ta to three acclimation temperature (Taccl) treatments (daytime Taccl ≈ 30 °C, 36 °C or 42 °C respectively; n ≈ 10 per site per Taccl). After an acclimation period of 30 days, heat tolerance and evaporative cooling capacity was quantified by exposing birds to progressively higher Ta until they approached severe hyperthermia (body temperature [Tb] = 44.5 °C; Ta range: 38-54 °C). We measured metabolic rate and evaporative water loss using open flow-through respirometry, and Tb using temperature-sensitive passive-integrated transponder tags. Hyperthermia threshold Ta (Ta,HT) was significantly higher and Tb significantly lower in birds acclimated to the hottest Taccl compared to those from milder acclimation treatments. Population (i.e., site of capture) was not a significant predictor of any thermoregulatory variables or hyperthermia threshold Ta (Ta,HT) after acclimation, revealing that the shape of reaction norms for heat tolerance and evaporative cooling capacity does not vary among these three populations.

Avian thermoregulation in the heat: metabolism, evaporative cooling and gular flutter in two small owls.

The thermoregulatory responses of owls to heat stress have been the subject of few studies. Although nocturnality buffers desert-dwelling owls from significant heat stress during activity, roost sites in tree and cactus cavities or in deep shade provide only limited refuge from high environmental temperatures during the day. We measured thermoregulatory responses to acute heat stress in two species of small owls, the elf owl (Micrathene whitneyi) and the western screech-owl (Megascops kennicottii), which occupy the Sonoran Desert of southwestern North America, an area of extreme heat and aridity. 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 limits (HTL; the maximum Ta reached). Comparatively low RMR values were observed in both species, Tb approximated Ta at 40°C and mild hyperthermia occurred as Ta was increased toward the HTL. Elf owls and screech-owls reached HTLs of 48 and 52°C, respectively, and RMR increased to 1.5 and 1.9 times thermoneutral values. Rates of EWL at the HTL allowed for the dissipation of 167-198% of metabolic heat production (MHP). Gular flutter was used as the primary means of evaporative heat dissipation and produced large increases in evaporative heat loss (44-100%), accompanied by only small increases (<5%) in RMR. These small, cavity-nesting owls have thermoregulatory capacities that are intermediate between those of the open-ground nesting nightjars and the passerines that occupy the same ecosystem.

Avian thermoregulation in the heat: phylogenetic variation among avian orders in evaporative cooling capacity and heat tolerance.

Little is known about the phylogenetic variation of avian evaporative cooling efficiency and heat tolerance in hot environments. We quantified thermoregulatory responses to high air temperature (Ta) in ∼100-g representatives of three orders, namely, the African cuckoo (Cuculus gularis, Cuculiformes), lilac-breasted roller (Coracias caudatus, Coraciiformes) and Burchell's starling (Lamprotornis australis, Passeriformes). All three species initiated respiratory mechanisms to increase evaporative heat dissipation when body temperature (Tb) approached 41.5°C in response to increasing Ta, with gular flutter observed in cuckoos and panting in rollers and starlings. Resting metabolic rate and evaporative water loss increased by quantitatively similar magnitudes in all three species, although maximum rates of evaporative water loss were proportionately lower in starlings. Evaporative cooling efficiency [defined as the ratio of evaporative heat loss (EHL) to metabolic heat production (MHP)] generally remained below 2.0 in cuckoos and starlings, but reached a maximum of ∼3.5 in rollers. The high value for rollers reveals a very efficient evaporative cooling mechanism, and is similar to EHL/MHP maxima for similarly sized columbids which very effectively dissipate heat via cutaneous evaporation. This unexpected phylogenetic variation among the orders tested in the physiological mechanisms of heat dissipation is an important step toward determining the evolution of heat tolerance traits in desert birds.

Avian thermoregulation in the heat: evaporative cooling capacity and thermal tolerance in two Australian parrots.

Avian orders differ in their thermoregulatory capabilities and tolerance of high environmental temperatures. Evaporative heat loss, and the primary avenue whereby it occurs, differs amongst taxa. Although Australian parrots (Psittaciformes) have been impacted by mass mortality events associated with extreme weather events (heat waves), their thermoregulatory physiology has not been well characterized. We quantified the upper limits to thermoregulation under extremely hot conditions in two Australian parrots: the mulga parrot (Psephotellus varius; ∼55 g) and the galah (Eolophus roseicapilla; ∼265 g). At air temperatures (Ta) exceeding body temperature (Tb), both species showed increases in Tb to maximum values around 43-44°C, accompanied by rapid increases in resting metabolic rate above clearly defined upper critical limits of thermoneutrality and increases in evaporative water loss to levels equivalent to 700-1000% of baseline rates at thermoneutral Ta Maximum cooling capacity, quantified as the fraction of metabolic heat production dissipated evaporatively, ranged from 1.71 to 1.79, consistent with the known range for parrots, similar to the corresponding range in passerines, and well below the corresponding ranges for columbids and caprimulgids. Heat tolerance limit (the maximum Ta tolerated) ranged from 44 to 55°C, similar to the range reported for passerines, but lower than that reported for columbids and caprimulgids. Our data suggest that heat tolerance in parrots is similar to that in passerines. We argue that understanding how thermoregulatory capacity and heat tolerance vary across avian orders is vital for predicting how climate change and the associated increase in frequency of extreme weather events may impact avian populations in the future.

Avian thermoregulation in the heat: is evaporative cooling more economical in nocturnal birds?

Evaporative cooling is a prerequisite for avian occupancy of hot, arid environments, and is the only avenue of heat dissipation when air temperatures (Ta) exceed body temperature (Tb). Whereas diurnal birds can potentially rehydrate throughout the day, nocturnal species typically forgo drinking between sunrise and sunset. We hypothesized that nocturnal birds have evolved reduced rates of evaporative water loss (EWL) and more economical evaporative cooling mechanisms compared with diurnal species, permitting nocturnal species to tolerate extended periods of intense heat without becoming lethally dehydrated. We used phylogenetically informed regressions to compare EWL and evaporative cooling efficiency [ratio of evaporative heat loss (EHL) and metabolic heat production (MHP); EHL/MHP] among nocturnal and diurnal birds at high Ta We analyzed variation in three response variables: (1) slope of EWL at Ta between 40 and 46°C, (2) EWL at Ta=46°C and (3) EHL/MHP at Ta=46°C. Nocturnality emerged as a weak, negative predictor, with nocturnal species having slightly shallower slopes and reduced EWL compared with diurnal species of similar mass. In contrast, nocturnal activity was positively correlated with EHL/MHP, indicating a greater capacity for evaporative cooling in nocturnal birds. However, our analysis also revealed conspicuous differences among nocturnal taxa. Caprimulgids and Australian owlet-nightjars had shallower slopes and reduced EWL compared with similarly sized diurnal species, whereas owls had EWL rates comparable to those of diurnal species. Consequently, our results did not unequivocally demonstrate more economical cooling among nocturnal birds. Owls predominately select refugia with cooler microclimates, but the more frequent and intense heat waves forecast for the 21st century may increase microclimate temperatures and the necessity for active heat dissipation, potentially increasing owls' vulnerability to dehydration and hyperthermia.