Structure and evolution of the sexually dimorphic integumentary swelling on the hands of dendrobatid poison frogs and their relatives (Amphibia: Anura: Dendrobatoidea).

The hands of adult males of many dendrobatid poison frogs and their relatives possess swelling formed by glandular tissue hypothesized to secrete courtship pheromones delivered to the female during cephalic amplexus. Variation in the occurrence and external morphology of the swelling of finger IV has provided important evidence for dendrobatoid systematics for decades, but its underlying structure has not been investigated. We undertook a detailed comparative analysis of the integument of the hand, including both external morphology and histology, of 36 species representing the diversity of dendrobatoid frogs and several close relatives. The swelling is caused by four densely packed, hypertrophic, morpho-histochemical types of specialized mucous glands (SMGs). We observed type I SMGs on fingers II-V and the wrist, including areas that are not swollen, types II and IV exclusively on finger IV, and type III on finger IV and the wrist. Type I SMGs occur either in isolation or together with types II, III, or IV; types II, III, and IV never occur together or without type I. We delimited 15 characters to account for the variation in external morphology and the occurrence of SMGs. Our data suggest that type I SMGs are a new synapomorphy for Dendrobatoidea and that type II SMGs originated in either the most recent common ancestor (MRCA) of Dendrobatidae or independently in the MRCAs of Aromobatidae and Colostethinae, respectively, while types III and IV are restricted to Anomaloglossus. The discovery of these SMGs adds a new dimension to studies of poison frog reproductive biology, which have investigated acoustic, visual, and tactile cues in courtship, mating, and parental care across the diversity of Dendrobatoidea for decades but have almost entirely overlooked the possible role of chemical cues.

Parotoid, radial, and tibial macroglands of the frog Odontophrynus cultripes: differences and similarities with toads

Anuran integument is characterized by the presence of glands, some of which are responsible for toxin production. In some species these glands accumulate in parts of the body strategically located against predators, forming structures known as macroglands. This is the case for parotoid macroglands, on the dorsum of the head, tibial macroglands, on the rear limbs, and radial macroglands, on the forelimbs of toads and some other anurans. The toad Rhinella jimi, for example, simultaneously displays all three types of macroglands, which is unusual even among bufonids. Interestingly, considering the phylogenetic distance, the frog Odontophrynus cultripes (Odontophrynidae) also presents these three macroglandular types. In this study we analyze the morphology of 0. cultripes macroglands and the chemical composition of their poison using an interdisciplinary approach. In this species, the parotoid, tibial, and radial macroglands consist of aggregates of elongated and juxtaposed poison glands, arranged in a honeycomb style, very similar to that of toads. Comparative analysis of these three macrogland types shows significant differences in both the morphology of secretory granules and biochemical composition. The present work on 0. cultripes contributes to the evidence that amphibians, or at least anurans, share a basic design for all cutaneous glandular accumulations. The determinant factor for macroglandular formation may be the selective pressure for defense against predators.