The river martins are in some ways intermediate between typical swallows and other passerines: they have stout bills, large feet and relatively strong legs, which is unusual in aerial feeders. They also have a large syrinx (vocal organ) and a different bronchial structure. The extent of their differences from other swallows and the wide geographical separation of these two martins suggest that they are relict populations of a group of species that diverged from the main swallow lineage early in its evolutionary history, and they may be the most primitive of the swallows. Like other early hirundine lineages, they nest in burrows, rather than adopted nest holes or mud nests.