Beach-spawning fishes, terrestrial eggs, and air breathing.

Many fishes have independently evolved beach spawning with oviposition at the water's edge. These include intertidal, subtidal, and estuarine, as well as a few freshwater, species. Their spectacular reproductive behavior at the boundary of water and land has focused attention on adults, but they emerge either briefly or not at all. The need for air breathing is more apparent in the eggs, and the reasons for emergence are more applicable to eggs than to the adults of most beach-spawning fishes. There is little evidence of air breathing in the adults, unless they are regularly emerged at other times as well. Conversely, eggs metabolize in air and show substantial emergence tolerance. We consider beach spawning a form of parental care in fishes. The adults place eggs so they will be emerged into air during part or all of incubation, providing increased temperatures, oxygen availability, and protection. Beach spawning provides habitat segregation at different points in the life history, with air emergence early in the life cycle and a return to water at hatching. The parents take great risks to spawn at the water's edge to give their offspring the most advantageous beginning in life.