Dehydration risk, not ambient incubation, limits nest attendance at high temperatures
High air temperatures have measurable negative impacts on reproduction in wild animal populations, including during incubation in birds. Understanding the mechanisms driving these impacts requires comprehensive knowledge of animal physiology and behaviour under natural conditions. We used a novel combination of a non-invasive doubly-labelled water technique and behaviour observations in the field to examine effects of temperature, rainfall, and group size on physiology and behaviour during incubation in southern pied babblers Turdoides bicolor, a cooperatively-breeding passerine endemic to a semi-arid region in southern Africa. The proportion of time that clutches of eggs were incubated declined as air temperatures increased, traditionally interpreted as a benefit of ambient incubation. However, we show that a) clutches were less likely to hatch when exposed to high air temperatures; b) pied babbler groups incubated their nests almost constantly (97% of daylight hours) except on hot days; c) operative temperatures in unattended nests were substantially higher than air temperatures and frequently exceeded 40.5{degrees}C, above which bird embryos are at risk of death; d) pied babblers incubating for long periods of time failed to maintain water balance on hot days but not cool days; and e) pied babblers from incubating groups did not maintain body mass on hot days. These results suggest that, rather than taking advantage of opportunities for ambient incubation, pied babblers leave the nests during hot periods to avoid dehydration as a consequence of incubating at high operative temperatures. As mean air temperatures increase and extreme heat events become more frequent under climate change, birds will likely incur greater water costs during incubation, leading to compromised nest attendance and increased likelihood of eggs overheating, with implications for nest success and, ultimately, population persistence.