Some anamniotic aquatic vertebrates lay eggs in a terrestrial habitat that is hostile to the survival of hatchings or larvae. These terrestrial eggs are ready and able to hatch at a particular developmental time, but do not hatch until presented with suitable conditions for aquatic larval survival. Beyond this time, hatching is possible whenever aquatic conditions occur. The duration of extended terrestrial incubation is dependent on the availability of energy for metabolism from the yolk. Extended incubation is useful for anamniotic eggs laid in terrestrial habitats where conditions suitable for larval survival arrive with unpredictable or variable timing. Examples of anamniotes with delayed hatching and extended terrestrial incubation can be found among teleost fishes, anurans, and caudate amphibians. This paper characterizes the embryonic period, compares this mode with other forms of developmental plasticity in anamniotes, evaluates the constraints and advantages of this life history mode, and examines how some fishes and amphibians are able to obtain the benefits of terrestriality for their eggs when the timing of the return to aquatic conditions is not entirely predictable.

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.

A surprisingly large number of fish and amphibian species reproduce terrestrially despite the absence of the key evolutionary innovation of the amniotic egg. In contrast with shelled eggs of reptiles and birds, eggs of teleost fish and amphibians are typically much smaller and enclosed in relatively simple chorionic membranes. Incubation times may be brief or prolonged, and resultant hatchlings typically require the return to an aquatic habitat. Advantages of terrestrial incubation include the increased availability of warmer temperatures and avoidance of aquatic hypoxia, whereas disadvantages include desiccation, exposure to novel predators, and the risk of hatching into a hostile habitat. Hatching may be environmentally cued. Use of energy in the yolk may require trade-offs between growth of the embryo and extended incubation, as exemplified by a case study of the California Grunion. The physical challenges of terrestrial incubation, constraints for hatching, effects of egg size, and parental care are explored. Eight different types of early life history among anamniotic embryos incubating in a terrestrial environment are identified, with examples of these alternate routes to the invasion of land by vertebrates.