Diversity and evolution of the parotoid macrogland in true toads (Anura: Bufonidae)

Skin glands in amphibians are either distributed throughout the skin or aggregated in multiglandular structures such as the parotoids typical of most species of Bufonidae. Although many early divergent and derived bufonids lack a discrete parotoid in the postorbital–supratympanic (PoSt) region, they have a great macroscopic diversity in the skin morphology of this region. To understand the origin and evolution of this diversity, in particular of the parotoids, we describe the histomorphology of the skin of the PoSt and dorsal regions in 17 species of bufonids, with or without external evident parotoid, and compare it with previously published descriptions. The survey results in 27 characters that were optimized on a phylogenetic hypothesis of Bufonidae. Our results reveal that the PoSt region has a noteworthy morphological diversity of types of glands, spatial organization and differences in the secretion products. Some morphological characters represent putative synapomorphies of internal clades of Bufonidae and are related to the progressive differentiation towards defined structures (macroglands, parotoids). These morphological results, along with published information on the toxicity of the skin secretions and defensive behaviours in some representative species, allow us to infer possible relationships between these features.