Has the importance of the amniote egg been overstated?

The evolution of the amniote egg is commonly regarded as an important milestone in the history of the vertebrates, an innovation that completed the transition from aquatic to fully terrestrial existence by permitting eggs to be laid away from standing water. This view derives ultimately from the recapitulationist theories of Haeckel, and rests on the assumption that extant frogs and salamanders are good models for the reproductive habits of early tetrapods and the ancestors of the amniotes. It also assumes that it is more difficult to lay eggs on land than in water, and that the amniote egg is an adaptation to the physical rigours that eggs encounter in terrestrial environments. Taken together, these assumptions comprise what may be termed the 'Haeckelian framework' for the origin of vertebrate terrestriality. Several independent lines of evidence suggest that the assumptions of the Haeckelian framework are false. There appear to be no theoretical reasons to assume that the evolution of terrestrial egg-laying was difficult, or required a structure as elaborate as the amniote egg. The physical conditions eggs encounter in the terrestrial environments where they are actually laid are quite mild. Land may in fact be an easier place to lay eggs than water. In addition, analysis of the distribution of key reproductive character states among vertebrates provides no evidence that the 'typical amphibian' reproductive mode is primitive for tetrapods. Amniotes are as likely as frogs or salamanders to retain primitive reproductive character states.