Cryptobatrachus

Abstract: Egg-brooding frogs (Hemiphractidae) are a group of 105 currently recognized Neotropical species, with a remarkable diversity of developmental modes, from direct development to free-living and exotrophic tadpoles. Females carry their eggs on the back and embryos have unique bell-shaped gills. We inferred the evolutionary relationships of these frogs and used the resulting phylogeny to review their taxonomy and test hypotheses on the evolution of developmental modes and bell-shaped gills. Our inferences relied on a total evidence parsimony analysis of DNA sequences of up to 20 mitochondrial and nuclear genes (analyzed under tree-alignment), and 51 phenotypic characters sampled for 83% of currently valid hemiphractid species. Our analyses rendered a well-resolved phylogeny, with both Hemiphractidae (sister of Athesphatanura) and its six recognized genera being monophyletic. We also inferred novel intergeneric relationships [((Cryptobatrachus, Flectonotus), (Stefania, (Fritziana, (Hemiphractus, Gastrotheca))))], the non-monophyly of all species groups previously proposed within Gastrotheca and Stefania, and the existence of several putative new species within Fritziana and Hemiphractus. Contrary to previous hypotheses, our results support the most recent common ancestor of hemiphractids as a direct-developer. Free-living aquatic tadpoles apparently evolved from direct-developing ancestors three to eight times. Embryos of the sister taxa Cryptobatrachus and Flectonotus share a pair of single gills derived from branchial arch I, while embryos of the clade including the other four genera have two pairs of gills derived from branchial arches I and II respectively. Furthermore, in Gastrotheca the fusion of the two pairs of gills is a putative synapomorphy. We propose a revised taxonomy concordant with our optimal topologies.

Abstract: Three patterns of superficial mandibular musculature are described in hemiphractine hylid frogs. One of these is unique to the morphologically bizarre Hemiphractus. A second pattern is found in Flectonotus and also occurs in some species of Gastrotheca and Stefania. A third pattern involves a differentiated apical element of the m. intermandibularis and is found in Cryptobatrachus, many species of Gastrotheca, and one species of Stefania. Evidence supports the plesiomorphic state of an undifferentiated m. intermandibularis and two derived states of differentiation of that muscle. One of these is the development of supplementary posterolateral elements characteristic of the Phyllomedusinae, whereas the differentiation of an apical element has occurred in at least six independent lineages--the entire Pelodryadinae, three unrelated genera of Hylinae, and two genera of Hemiphractinae. Gastrotheca and Stefania are the only anuran genera known to include species possessing, and others lacking, differentiation of the m. intermandibularis. Vocal sacs and apertures are absent in Cryptobatrachus, Hemiphractus, Stefania, and six species of Gastrotheca.

Abstract: Hemiphractid frogs of the genus Cryptobatrachus are nearly exclusive to Colombia and are distributed over an altitudinal range between 360 and 2400 meters on the three cordilleras and the massif of the Sierra Nevada de Santa Marta. To the three species currently recognized, three additional species are named (C. conditus, C. pedroruizi, and C. ruthveni). The genus is monophyletic and all but the cis-Andean C. nicefori form a clade (defined by the presence of adhesive pads on some subarticular tubercles of the hand and foot). All but C. fuhrmanni and C. nicefori form an additional clade in northern Colombia. Two (or three) species are found on the Sierra Nevada de Santa Marta and two other species are found on the Serrania de Perija. The most widely distributed species (C. fuhrmanni) occurs on all three Andean cordilleras without apparent geographic variation.