Foot-mediated incubation: Nazca booby (Sula granti) feet as surrogate brood patches

Incubation in most avian species involves transferring heat from parent to egg through a highly vascularized brood patch. Some birds, however, do not develop a brood patch. Unusual among birds, these species hold their eggs under the webs of their feet, but the role of the feet in heat transfer is uncertain. Often the webs are positioned between the feathered abdomen and the egg during incubation, suggesting that either the abdomen, the feet, or both could transfer heat to the egg. We studied heat transfer from foot webs to eggs during incubation in Nazca boobies by spatially separating the feet from the abdomen using an oversized egg. We found that feet transfer heat to eggs independently of any heat that may be transferred from the abdomen. In addition, we found that incubating boobies had significantly greater vascularization in their foot webs, measured as a percentage of web area covered by vessels, than nonincubating boobies. We also found that males, whether incubating or nonincubating, had significantly less web vascularization than females. We concluded that vascularized Nazca booby feet function in the same way during incubation that vascularized brood patches do, acting as surrogate brood patches.