We explored the relationship between lipids of the stratum corneum (SC), the barrier to water-vapor diffusion of the skin, and cutaneous water loss (CWL) of species of free-living larks along a temperature-moisture gradient. Our results showed that free fatty acids, cholesterol, and ceramides were the major constituents of SC in larks from different environments including the Netherlands, a mesic environment; Iran, a semiarid region; and several areas in Saudi Arabia, a hot dry desert. We found that CWL was reduced among larks inhabiting deserts, but our data did not support the hypothesis that birds from desert environments have larger quantities of lipids per unit dry mass of the SC than larks from more mesic environments. Instead, larks in arid environments had a higher proportion of ceramides, especially the more polar fractions 4-6, and a smaller proportion of free fatty acids in their SC, an adjustment that apparently reduced their CWL. Subtle changes in the ratios of lipid classes can apparently alter the movement of water vapor through the skin. We hypothesize that desert birds have higher proportions of ceramides in their SC and lower proportions of free fatty acids because this combination allows the lipid lamellae to exist in a more highly ordered crystalline phase and consequently creates a tighter barrier to water-vapor diffusion.

SUMMARY When vertebrates invaded land during the Carboniferous period, they were exposed not only to new ecological opportunities but also to a desiccating environment. To maintain cellular water homeostasis, natural selection modified the integument of pioneering terrestrial animals, enabling them to reduce water loss through the skin. In desert environments, where ambient temperatures (Ta) can reach 50°C, relative humidities are low and drinking water is scarce, integumentary modifications that reduce cutaneous water loss (CWL) could be fundamental to survival. Previous research has shown that hoopoe larks (Alaemon alaudipes) from the Arabian desert reduced CWL when acclimated to 35°C compared with individuals at 15°C, but skylarks (Alauda arvensis) and woodlarks (Lullula arborea), from The Netherlands, and Dunn's larks (Eremalauda dunni), also from the Arabian desert, did not. Here, we test the idea that hoopoe larks acclimated to 35°C would alter the lipid composition of their stratum corneum (SC), resulting in a decrease in CWL, but that skylarks,woodlarks and Dunn's larks would not. Specifically, we hypothesized that hoopoe larks, acclimated to 35°C, would increase the proportions of polar ceramide content and decrease the proportions of free fatty acids in their SC compared with individuals acclimated to 15°C. Results showed that hoopoe larks at 35°C had lower CWL and higher proportions of total ceramides but lower proportions of free fatty acids and sterols in their SC. We demonstrate that adjustments in ratios of lipid classes in the SC are associated with changes in CWL in hoopoe larks.