Brave new propagules: terrestrial embryos in anamniotic eggs.

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.