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The study of early development in Neotropical Leiuperinae frogs (Anura, Leptodactylidae) has been addressed by several works in recent times. However, a comparative developmental approach under a phylogenetic context was not available. Herein we analyzed the morphological and ontogenetic diversity of embryos belonging to 22 species of the three largest genera in Leiuperinae. We find that in most cases, variations fit with the phylogeny at the inter- and intrageneric levels. Embryo kyphosis and whitish color are synapomorphies for the clade grouping Physalaemus and Engystomops . The presence of a third lower tooth row on the oral disc is plesiomorphic for Leiuperinae, only changing in derived clades. The configurations and developmental trajectories of the lower lip are exceptionally diverse. The developmental sequences optimized on the phylogenetic tree recover an early differentiated first lower tooth row a synapomorphy of Pseudopaludicola and Physalaemus , and an early differentiated second row as synapomorphy of Pleurodema . On the other hand, few features are highly conserved in the subfamily, such as the adhesive glands universally present in a type-C configuration. Our results also suggest that the morphology and ontogeny of embryos is in some cases associated to the environment where they develop. A large body size, poorly developed transient respiratory structures, large yolk provision and delayed development of the digestive tract occur convergently in embryos inhabiting cold, oxygenated environments. Embryos that develop in warmer water bodies in xeric environments show more complex and persistent transient respiratory structures and an early onset of hind limbs development. Our survey highlights that morphology and early development of anurans can be a valuable source of information for phylogenetic studies, and provide fundamental bases to explore and discuss how evolutionary changes can be shaped by environmental conditions.
The study of early development in Neotropical Leiuperinae frogs (Anura, Leptodactylidae) has been addressed by several works in recent times. However, a comparative developmental approach under a phylogenetic context was not available. Herein we analyzed the morphological and ontogenetic diversity of embryos belonging to 22 species of the three largest genera in Leiuperinae. We find that in most cases, variations fit with the phylogeny at the inter- and intrageneric levels. Embryo kyphosis and whitish color are synapomorphies for the clade grouping Physalaemus and Engystomops . The presence of a third lower tooth row on the oral disc is plesiomorphic for Leiuperinae, only changing in derived clades. The configurations and developmental trajectories of the lower lip are exceptionally diverse. The developmental sequences optimized on the phylogenetic tree recover an early differentiated first lower tooth row a synapomorphy of Pseudopaludicola and Physalaemus , and an early differentiated second row as synapomorphy of Pleurodema . On the other hand, few features are highly conserved in the subfamily, such as the adhesive glands universally present in a type-C configuration. Our results also suggest that the morphology and ontogeny of embryos is in some cases associated to the environment where they develop. A large body size, poorly developed transient respiratory structures, large yolk provision and delayed development of the digestive tract occur convergently in embryos inhabiting cold, oxygenated environments. Embryos that develop in warmer water bodies in xeric environments show more complex and persistent transient respiratory structures and an early onset of hind limbs development. Our survey highlights that morphology and early development of anurans can be a valuable source of information for phylogenetic studies, and provide fundamental bases to explore and discuss how evolutionary changes can be shaped by environmental conditions.
The Patagonian rainforest has environmental conditions that may have been appropriate for the evolution of different reproductive strategies in anurans. Despite its low species richness, the endemic family Batrachylidae has a wide diversity of reproductive modes. All species have exotrophic tadpoles, but oviposition sites may vary from lakes/ponds to the forest floor and tree holes. Here we study the embryonic development of five species to explore the morphological diversity and heterochronic shifts related to aquatic vs. terrestrial development. The main differences concern the length of the intracapsular period, morphology at tailbud stage, and gut and hind-limb development. Terrestrial embryos of Batrachyla have some peculiar features also reported in species with endotrophic development, such as yolk-rich eggs, poorly developed gills, precocious development of hind limbs and delayed gut coiling. Furthermore, observations on embryos releasing from the jelly layers with the vitelline membrane intact plus the likely absence of hatching gland cells at hatching stages make it likely that differences can occur in the hatching mechanism. Our results highlight that batrachylids and other neoaustraranans offer an excellent opportunity to investigate the evolution and diversity of reproductive and developmental strategies divergent from aquatic/exotrophic development.
Anurans have served as models in evolutionary studies on reproduction and development. Several generalizations have emerged but it is unclear whether they represent the diversity of the entire group. We studied the development of two species of the Neotropical family Cycloramphidae. Frogs in this family have peculiar spawning sites and developmental biologies. Whereas some cycloramphid tadpoles are exotrophic, quasi-terrestrial and thrive in films of water that flow over rocky outcrops, others are endotrophic, terrestrial and complete their development outside the eggs laid on the forest floor, under logs or in crevices. Cycloramphid tadpoles have aspects of their morphology distinct from those of pond and stream dwellers; they hatch in more advanced stages of development and have typical tadpole structures, either in a modified form or absent. Herein we present a study of the tadpoles of C. lithomimeticus and T. miliaris, in order to document and discuss alterations in the ontogeny of several larval structures. These changes seem to be associated with adaptations to life in the peculiar habitats where the tadpoles thrive. We infer that the observations we present are explained by development being associated with aspects of terrestrialization and endotrophism.
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