Direct impacts of climatic warming on heat stress in endothermic species: seabirds as bioindicators of changing thermoregulatory constraints

There is now abundant evidence that contemporary climatic change has indirectly affected a wide-range of species by changing trophic interactions, competition, epidemiology and habitat. However, direct physiological impacts of changing climates are rarely reported for endothermic species, despite being commonly reported for ectotherms. We review the evidence for changing physiological constraints on endothermic vertebrates at high temperatures, integrating theoretical and empirical perspectives on the morphology, physiology and behavior of marine birds. Potential for increasing heat stress exposure depends on changes in multiple environmental variables, not just air temperature, as well as organism-specific morphology, physiology and behavior. Endotherms breeding at high latitudes are vulnerable to the forecast, extensive temperature changes because of the adaptations they possess to minimize heat loss. Low-latitude species will also be challenged as they currently live close to their thermal limits and will likely suffer future water shortages. Small, highly-active species, particularly aerial foragers, are acutely vulnerable as they are least able to dissipate heat at high temperatures. Overall, direct physiological impacts of climatic change appear underrepresented in the published literature, but available data suggest they have much potential to shape behavior, morphology and distribution of endothermic species. Coincidence between future heat stress events and other energetic constraints on endotherms remains largely unexplored but will be key in determining the physiological impacts of climatic change. Multi-scale, biophysical modeling, informed by experiments that quantify thermoregulatory responses of endotherms to heat stress, is an essential precursor to urgently-needed analyses at the population or species level.