High temperatures are associated with substantial reductions in breeding success and offspring quality in an arid-zone bird.

During hot weather, terrestrial animals often seek shaded thermal refugia. However, this can result in missed foraging opportunities, loss of body condition and impaired parental care. We investigated whether such costs could compromise breeding success in a widespread southern African bird: the Southern Yellow-Billed Hornbill Tockus leucomelas. We predicted that hornbills might be especially vulnerable to temperature-dependant reductions in parents' foraging capacity due to extreme asymmetry in sex-specific roles during breeding: females are confined within the nest cavity for most of the nesting period and the burden of provisioning falls solely on the male during this time. We followed 50 hornbill nesting attempts in the Kalahari Desert between 2012 and 2015, collecting data on provisioning rates, adult and nestling body mass, fledging success and size of fledglings. Mean daily maximum air temperatures (Tmax) during nesting attempts ranged from 33.2 to 39.1 °C. The likelihood of successful fledging fell below 50% at mean Tmax > 35.1 °C; a threshold now regularly exceeded at our study site due to recent climate warming. Additionally, offspring fledging following the hottest nesting attempts were > 50% lighter than those fledging following the coolest. Sublethal costs of keeping cool including loss of body condition, production of poor-quality offspring and breeding failure are likely to become issues of serious conservation concern as climate change progresses; even for currently widespread species. Missed-opportunity costs associated with behavioral thermoregulation and direct sublethal costs of temperature exposure should not be overlooked as a potential threat to populations, especially in environments that are already hot.

The costs of keeping cool: behavioural trade-offs between foraging and thermoregulation are associated with significant mass losses in an arid-zone bird.

Avian responses to high environmental temperatures include retreating to cooler microsites and/or increasing rates of evaporative heat dissipation via panting, both of which may affect foraging success. We hypothesized that behavioural trade-offs constrain the maintenance of avian body condition in hot environments, and tested predictions arising from this hypothesis for male Southern Yellow-billed Hornbills (Tockus leucomelas) breeding in the Kalahari Desert. Operative temperatures experienced by the hornbills varied by up to 13 °C among four microsite categories used by foraging males. Lower prey capture rates while panting and reductions associated with the occupancy of off-ground microsites, resulted in sharp declines in foraging efficiency during hot weather. Consequently, male body mass (Mb) gain between sunrise and sunset decreased with increasing daily maximum air temperature (Tmax), from ~ 5% when Tmax < 25 °C to zero when Tmax = 38.4 °C. Overnight Mb loss averaged ~ 4.5% irrespective of Tmax, creating a situation where nett 24-h Mb loss approached 5% on extremely hot days. These findings support the notion that temperature is a major determinant of body condition for arid-zone birds. Moreover, the strong temperature dependence of foraging success and body condition among male hornbills provisioning nests raises the possibility that male behavioural trade-offs translate into equally strong effects of hot weather on female condition and nest success. Our results also reveal how rapid anthropogenic climate change is likely to substantially decrease the probability of arid-zone birds like hornbills being able to successfully provision nests while maintaining their own condition.

Regulation of Heat Exchange across the Hornbill Beak: Functional Similarities with Toucans?

Beaks are increasingly recognised as important contributors to avian thermoregulation. Several studies supporting Allen's rule demonstrate how beak size is under strong selection related to latitude and/or air temperature (Ta). Moreover, active regulation of heat transfer from the beak has recently been demonstrated in a toucan (Ramphastos toco, Ramphastidae), with the large beak acting as an important contributor to heat dissipation. We hypothesised that hornbills (Bucerotidae) likewise use their large beaks for non-evaporative heat dissipation, and used thermal imaging to quantify heat exchange over a range of air temperatures in eighteen desert-living Southern Yellow-billed Hornbills (Tockus leucomelas). We found that hornbills dissipate heat via the beak at air temperatures between 30.7°C and 41.4°C. The difference between beak surface and environmental temperatures abruptly increased when air temperature was within ~10°C below body temperature, indicating active regulation of heat loss. Maximum observed heat loss via the beak was 19.9% of total non-evaporative heat loss across the body surface. Heat loss per unit surface area via the beak more than doubled at Ta > 30.7°C compared to Ta < 30.7°C and at its peak dissipated 25.1 W m(-2). Maximum heat flux rate across the beak of toucans under comparable convective conditions was calculated to be as high as 61.4 W m(-2). The threshold air temperature at which toucans vasodilated their beak was lower than that of the hornbills, and thus had a larger potential for heat loss at lower air temperatures. Respiratory cooling (panting) thresholds were also lower in toucans compared to hornbills. Both beak vasodilation and panting threshold temperatures are potentially explained by differences in acclimation to environmental conditions and in the efficiency of evaporative cooling under differing environmental conditions. We speculate that non-evaporative heat dissipation may be a particularly important mechanism for animals inhabiting humid regions, such as toucans, and less critical for animals residing in more arid conditions, such as Southern Yellow-billed Hornbills. Alternatively, differences in beak morphology and hardness enforced by different diets may affect the capacity of birds to use the beak for non-evaporative heat loss.