Embryonic stages of Gastrotheca riobambae (Fowler) during maternal incubation and comparison of development with that of other egg-brooding hylid frogs.

A table of development (25 stages) for the period of incubation in the pouch was constructed for Gastrotheca riobambae; it can be used to stage embryos of other egg-brooding hylids. Analysis of embryonic weights during incubation shows that the mother does not contribute nutrients, but gases and other factors are probably exchanged between mother and embryos. According to species, incubation on the back of the mother is carried to the froglet or to the tadpole stages. Development in these hylids is characterized by specialized gills, the bell gills derived from the brachial arches. In some species, the bell gills derive from the first branchial arch and cover less than 50% of the embryo, while in others, the bell gills come from both branchial arches I and II and cover from less than 50% to 100% of the embryo. The most complex bell gills derive from the fusion of the two branchial arches. The majority of egg-brooding hylids live in tropical forests and carry development to the froglet stage. Tadpoles are produced by species of Flectonotus, Fritziana, and Gastrotheca. Tadpole-producing species of Gastrotheca have the most complex reproductive adaptations among egg-brooding hylids Acceleration and retardation in development seem to have played important roles in the evolution of these frogs. The evolutionary trend has been toward direct development, i.e., disappearance of the free-living larval stages through maternal incubation, and later to a recovery of the free-living tadpole stages in species of Gastrotheca with the most complex reproductive adaptations.