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Many birds reduce rest-phase energy demands through heterothermy, physiological responses involving facultative, reversible reductions in metabolic rate and body temperature (Tb). Here, we review the phylogenetic distribution and ecological contexts of avian heterothermy. Heterothermy has been reported in 140 species representing 15 orders and 39 families. Recent work supports the view that deep heterothermy is most pronounced in phylogenetically older taxa whereas heterothermy in passerines and other recently diverged taxa is shallower and confined to minimum Tb > 20°C. The reasons why deep heterothermy is absent in passerines remain unclear; we speculate an evolutionary trade-off may exist between the capacity to achieve low heterothermic Tb and the tolerance of hyperthermic Tb. Inter- and intraspecific variation in heterothermy is correlated with factors including foraging ecology (e.g., territoriality and defense of food resources among hummingbirds), food availability and foraging opportunities (e.g., lunar phase predicts torpor use in caprimulgids), and predation risk. Heterothermy also plays a major role before and during migration. Emerging questions include the magnitude of energy savings associated with heterothermy among free-ranging birds, the role phylogenetic variation in the capacity for heterothermy has played in evolutionary radiations into extreme habitats, and how the capacity for heterothermy affects avian vulnerability to rapid anthropogenic climate change., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Evaporative cooling is a prerequisite for avian occupancy of hot, arid environments, and is the only avenue of heat dissipation when air temperatures ( T a ) exceed body temperature ( T b ). 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 T a We analyzed variation in three response variables: (1) slope of EWL at T a between 40 and 46°C, (2) EWL at T a =46°C and (3) EHL/MHP at T a =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., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Nightjars represent a model taxon for investigating physiological limits of heat tolerance because of their habit of roosting and nesting in sunlit sites during the heat of the day. We investigated the physiological responses of Rufous-cheeked nightjars (Caprimulgus rufigena) and Freckled nightjars (Caprimulgus tristigma) to high air temperatures (T a ) by measuring body temperature (T b ), resting metabolic rate (RMR) and total evaporative water loss (TEWL) at T a ranging from 10 to 56 °C. Both species became hyperthermic at T a  > T b . Lower critical limits of thermoneutrality occurred at T a between 35 and 37 °C, whereas we detected no clear upper critical limits of thermoneutrality. Between T a  ≈ 37.0 and 39.9 °C, rates of TEWL increased rapidly with T a . At T a  ≥ 40 °C, fractional increases in mass-specific TEWL rates were 78-106% of allometric predictions. Increasing evaporative heat dissipation incurred only small metabolic costs, with the RMR of neither species ever increasing by more than 20% above thermoneutral values. Consequently, both species displayed extremely efficient evaporative cooling; maximum evaporative heat dissipation was equivalent to 515% of metabolic heat production (MHP) at T a  ≈ 56 °C in C. rufigena and 452% of MHP at T a  ≈ 52 °C in C. tristigma. Our data reiterate that caprimulgids have evolved an efficient mechanism of evaporative cooling via gular fluttering, which minimizes metabolic heat production at high T a and reduces total heat loads. This likely aids in reducing TEWL rates and helps nightjars cope with some of the most thermally challenging conditions experienced by any bird.

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

Compared to mammals, there are relatively few studies examining heterothermy in birds. In 13 bird families known to contain heterothermic species, the common poorwill (Phalaenoptilus nuttallii) is the only species that ostensibly hibernates. We used temperature-sensitive radio-transmitters to collect roost and skin temperature (Tskin) data, and winter roost preferences for free-ranging poorwills in southern Arizona. Further, to determine the effect of passive rewarming on torpor bout duration and active rewarming (i.e., the use of metabolic heat to increase Tskin), we experimentally shaded seven birds during winter to prevent them from passively rewarming via solar radiation. Poorwills selected winter roosts that were open to the south or southwest, facilitating passive solar warming in the late afternoon. Shaded birds actively rewarmed following at least 3 days of continuous torpor. Average torpor bout duration by shaded birds was 122 h and ranged from 91 to 164 h. Active rewarming by shaded birds occurred on significantly warmer days than those when poorwills remained torpid. One shaded bird remained inactive for 45 days, during which it spontaneously rewarmed actively on eight occasions. Our findings show that during winter poorwills exhibit physiological patterns and active rewarming similar to hibernating mammals. [ABSTRACT FROM AUTHOR]

Endotherms allocate large amounts of energy and water to the regulation of a precise body temperature (Tb), but can potentially reduce thermoregulatory costs by allowing Tb to deviate from normothermic levels. Many data on heterothermy at low air temperatures (Ta) exist for caprimulgids, whereas data on thermoregulation at high Ta are largely absent, despite members of this taxon frequently roosting and nesting in sites exposed to high operative temperatures. We investigated thermoregulation in free-ranging rufous-cheeked nightjars Caprimulgus rufigena and freckled nightjars Caprimulgus tristigma in the southern African arid zone. Individuals of both species showed labile Tb fluctuating around a single modal Tb (Tb-mod). Average Tb-mod was 39.7°C for rufous-cheeked nightjars and 39.0°C for freckled nightjars. In both species, diurnal Tb increased with increasing Ta. At Ta ≥ 38°C, rufous-cheeked nightjar mean Tb increased to 42°C, equivalent to 2.3°C above Tb-mod. Under similar conditions, freckled nightjar Tb was on average only 1.1°C above Tb-mod, with a mean Tb of 40.0°C. Freckled nightjars are one of the most heterothermic caprimulgids investigated to date, but our data suggest that during hot conditions this species maintains Tb within a narrow range above Tb-mod, possibly reflecting an evolutionary tradeoff between decreased thermal sensitivity to lower Tb but increased sensitivity to high Tb. These findings reveal how general thermoregulatory patterns at similar Ta can vary even among closely related species. [ABSTRACT FROM AUTHOR]

In summer, many temperate bat species use daytime torpor, but breeding females do so less to avoid interferences with reproduction. In forest-roosting bats, deep tree cavities buffer roost microclimate from abrupt temperature oscillations and facilitate thermoregulation. Forest bats also switch roosts frequently, so thermally suitable cavities may be limiting. We tested how barbastelle bats ( Barbastella barbastellus), often roosting beneath flaking bark in snags, may thermoregulate successfully despite the unstable microclimate of their preferred cavities. We assessed thermoregulation patterns of bats roosting in trees in a beech forest of central Italy. Although all bats used torpor, females were more often normothermic. Cavities were poorly insulated, but social thermoregulation probably overcomes this problem. A model incorporating the presence of roost mates and group size explained thermoregulation patterns better than others based, respectively, on the location and structural characteristics of tree roosts and cavities, weather, or sex, reproductive or body condition. Homeothermy was recorded for all subjects, including nonreproductive females: This probably ensures availability of a warm roosting environment for nonvolant juveniles. Homeothermy may also represent a lifesaver for bats roosting beneath loose bark, very exposed to predators, because homeothermic bats may react quickly in case of emergency. We also found that barbastelle bats maintain group cohesion when switching roosts: This may accelerate roost occupation at the end of a night, quickly securing a stable microclimate in the newly occupied cavity. Overall, both thermoregulation and roost-switching patterns were satisfactorily explained as adaptations to a structurally and thermally labile roosting environment. [ABSTRACT FROM AUTHOR]

Many bats, including some small-bodied tropical and subtropical Pteropodidae, use torpor to offset energetic constraints. We tested the hypothesis that medium-sized (110-160 g) cave-roosting Egyptian rousette bats (Rousettus aegyptiacus) at the southern extent of their range are able to employ torpor. We measured daytime body temperatures (Tb) of 9 wild individuals using implanted temperature-sensitive radiotransmitters. The bats roosted in a cave on Table Mountain, Cape Town, South Africa (a typically cool and wet area). Daily mean cave temperature (Tc) ranged between 7°C and 12°C (mean 9.3°C). All wild individuals exhibited a circadian cycle in Tb, with an average of 37.7°C upon return from foraging, decreasing to 35.5°C by mid-day. Before emergence for feeding, Tb increased to about 37°C. No individual allowed Tb to drop below 34°C indicating (assuming a threshold temperature of 30°C) that individuals in this population do not use torpor. Bats were active throughout the day within the roost and metabolic heat production may have contributed to the maintenance of rest-phase Tb. Ten individuals implanted with temperature-sensitive passive integrated transponder tags were held in captivity at temperatures of 25-30°C and subjected to food deprivation for 2-3 days. The lowest Tb recorded for any captive individual was 33.4°C despite losing an average of 10% of their initial body mass. Despite exposure to cool Ta in the cave, and often cold, wet, and windy conditions while foraging, or substantial food restriction in captive bats, R. aegyptiacus did not resort to using torpor. [ABSTRACT FROM AUTHOR]