Variations in anuran embryogenesis: yolk-rich embryos of Hyperolius puncticulatus (Hyperoliidae).

Anuran development is usually described using model species, most notably Xenopus laevis and Rana pipiens. We describe the development of the East African Reed Frog, Hyperolius puncticulatus, a species displaying development that is highly divergent from the "classic" anuran developmental pattern. Although having small eggs, the eggs of H. puncticulatus are characterized by a large amount of yolk, and embryonic development is reminiscent of species with large eggs. The eggs are teleolecithal and the cleavage is holoblastic, with a "pseudo-meroblastic" pattern. Gastrulation proceeds primarily at the dorsal lip and is characterized by reduced embryonic cavities. Gastrulation ends with a thickened "embryonic mantle" that sits upon a large yolk mass and forms most of the tissues of the embryo. The embryonic axis curves across the yolk mass, instead of the typical lengthening of anuran embryos. The tadpole hatches with a large ventral yolk mass which is gradually absorbed. We hypothesize about the developmental mechanisms that underlie this unusual development, based on comparisons with other anuran and fish species. We suggest that this type of development is not unique to this species, but can be found in many species of different anuran taxonomic groups. Comparing the development of H. puncticulatus and similar species to what is known about the development of model species, such as X. laevis, shows us the variation in early anuran embryogenesis. Knowing the existing diversity is a prerequisite to understanding the evolution of early anuran development and the changes in patterning mechanisms in different lineages.