Délio Baêta

The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae)

The leaf or monkey frogs of the hylid subfamily Phyllomedusinae are a unique group of charismatic anurans. We present a molecular phylogenetic analysis that includes 45 of the 60 species of phyllomedusines using up to 12 genes and intervening tRNAs. The aims were to gain a better understanding of the phylogenetic position of Phrynomedusa, test the monophyly and explore the relationships among several putative lineages (Hylomantis, the H. buckleyi Group, Phasmahyla, the four species groups of Phyllomedusa, and the species of Phyllomedusa that remain unassigned to any group), and to examine the implications of our phylogeny for the evolution of several characters in phyllomedusines. The analyses resulted in a well‐supported phylogenetic hypothesis that provides a historical framework for a discussion of the evolution of characters associated with reproductive biology, gliding behaviour, the physiology of waterproofing, and bioactive peptides. Implications include an earlier origin for eggless capsules than for leaf‐folding behaviour during amplexus, two independent origins of gliding, and an earlier origin of reduction in evaporative water loss than uricotelism, which is a result that originally was predicted on the basis of physiology alone. Furthermore, our results support the prediction that bioactive peptides from different peptide families are to be expected in all species of Phyllomedusinae. Hylomantis (as recently redefined) is shown to be paraphyletic and the synonymy of Agalychnis is revised to remedy this problem by including both Hylomantis and Pachymedusa.

Structural diversity of nuptial pads in Phyllomedusinae (Amphibia: Anura: Hylidae).

We studied the morphological variation of the nuptial pads using light microscopy and scanning electron microscopy (SEM) in 26 species of phyllomedusines (Anura: Hylidae), representing the five currently recognized genera. All phyllomedusines have single nuptial pads with dark colored epidermal projections (EPs). Spine‐shaped EPs occur in Cruziohyla calcarifer, Phrynomedusa appendiculata and in one species of Phasmahyla. The other species have roundish EPs. The density of the EPs on the pad is variable. Species in the Phyllomedusa hypochondrialis Group have EPs with a density that varies between 764 ± 58/mm2 and 923 ± 160/mm2. In all other studied species (including the Phyllomedusa burmeisteri and Phyllomedusa perinesos groups, Phyllomedusa camba, Phyllomedusa boliviana, Phyllomedusa sauvagii, Phyllomedusa bicolor, and Phyllomedusa tomopterna) the density of EPs varies between 108 ± 20/mm2 and 552 ± 97/mm2. Pores were observed with SEM in C. calcarifer, Agalychnis lemur, Agalychnis moreletii, but its presence is confirmed through histological sections on several other species. Its visibility using SEM seems to be related with the level of separation between adjacent EPs. The pores in the four studied species of Agalychnis are shown with SEM and histological sections to have a characteristic epidermal rim, that is absent in the otherphyllomedusines. Unlike most previous reports on breeding glands, those of phyllomedusines are alcian blue positive, indicating the presence of acidic mucosubstances on its secretions. J. Morphol. 2012.

Species limits, phylogeographic and hybridization patterns in Neotropical leaf frogs (Phyllomedusinae)

The taxonomy of many species is still based solely on phenotypic traits, which is often a pitfall for the understanding of evolutionary processes and historical biogeographic patterns, especially between closely related species due to either phenotypic conservatism or plasticity. Two widely distributed Neotropical leaf frogs from the Phyllomedusa burmeisteri species group (P. burmeisteri and Phyllomedusa bahiana) constitute a paramount example of closely related species with relatively unstable taxonomic history due to a large phenotypic variation. Herein, we analysed ~260 individuals from 57 localities distributed across the range of the two species to contrast individual phenotypic with an integrative phylogenetic and phylogeographic multilocus approach. We aim to clarify species limits, investigate potential undocumented diversity and examine to what extent taxonomic uncertainties could lead to misleading hypotheses on phylogeographic and interspecific hybridization patterns. Our molecular analysis supports the recognition of the two currently defined species, providing evidences for one novel and highly divergent evolutionary unit within the range of P. burmeisteri, which encompasses its type locality (Rio de Janeiro city). Spatial patterns of genetic and the colour of the hidden areas of the thigh was not congruent, varying considerably both within and between populations of both species. Genetic data showed signs of admixture between both species but do not corroborate the previously inferred wide area of introgression based on the distribution of the intermediate phenotype. Our results suggest that phenotypic variation can result from local adaptations, geographic isolation and/or evolutionary processes and, thus, cannot be used to reliably diagnose P. burmeisteri and P. bahiana. Globally, this study underscores the need of a geographical broad sampling of widespread species and the combination of molecular and phenotypic data to delineate species limits and phylogeographic patterns in species with complex taxonomy.

The submandibular musculature of phyllomedusinae (Anura: Hylidae): A reappraisal

The submandibular musculature of 37 species of the five currently recognized genera of the subfamily Phyllomedusinae (Anura: Hylidae) is described; observations are made on the variation and ontogeny of these muscles. Supplementary apical elements of the m. intermandibularis occur in all phyllomedusines studied, in addition to the supplementary posterolateral elements previously reported. Our observations are discussed in the context of 1) the proposed homology between supplementary apical and posterolateral elements; 2) the homology with the apical elements reported for Pelodryadinae (sister taxon of Phyllomedusinae); and 3) the implications for our understanding of the relationships between Phyllomedusinae and Pelodryadinae. Anatomical differences between the apical and posterolateral elements and their co‐occurrence in phyllomedusines indicate that these supplementary elements are not homologous. Despite differences between phyllomedusines and pelodryadines in the adhesion of supplementary fibers to the principal element of the m. intermandibularis and the occurrence of a broad aponeurosis or a medial raphe, the extensive morphological and developmental resemblances of the apical elements indicate that these structures are homologous, and that the presence of apical elements is a synapomorphy of Phyllomedusinae + Pelodryadinae. J. Morphol., 2011.

The Adhesive Glands during Embryogenesis in Some Species of Phyllomedusinae (Anura: Hylidae)

Abstract. Among anuran embryonic structures, the adhesive (cement) glands appear posterolaterally to the stomodeum and produce a mucous secretion that adheres embryos to surfaces in and out of the egg. In this paper, we study the ontogeny of the adhesive glands in five species of Phyllomedusa representing the two main clades recognized in the genus, plus embryos of Agalychnis aspera and Phasmahyla cochranae. Clutches were collected in the field, and embryos were periodically fixed to obtain complete developmental series and then studied with a stereomicroscope, scanning electron microscopy and routine histological techniques. Structural variations include glands absent (in P. cochranae and Phyllomedusa boliviana), functional club-shaped glands (morphogenetic Type C in Phyllomedusa sauvagii, Phyllomedusa iheringii, and Phyllomedusa tetraploidea), and an unusual Type C-like pattern in Phyllomedusa azurea, characterized by large, oblong glands in a horseshoe-like disposition around the oral disc. This latter gland configuration is similar to that of A. aspera. Interspecific variations also include the arrangement and regression pattern of the secretory region, which are in turn different from those of Type C glands in other clades. To interpret the origin and evolution of gland developmental patterns in the group, we still need information on gland occurrence and development in the basal genera of Phyllomedusinae (Phrynomedusa and Cruziohyla) and in the basal taxa of the two major clades of Phyllomedusa.

Vocal repertory of Scinax littoreus (Anura: Hylidae) with comments on the advertisement call of the Scinax perpusillus species group

The Scinax perpusillus species group consists of thirteen bromeligenous treefrogs, of which only six have had their acoustic parameters appropriately described. In this work, we present the vocal repertory of Scinax littoreus (Peixoto, 1988) based on recordings obtained from three different populations in the state of Rio de Janeiro, Brazil. Based on our observations we suggest that the vocal repertory of S. littoreus is composed of two distinct types of calls. The call named type A, which corresponds to a long series of multipulsed notes, is likely to have a mating function. This type of call is similar to that reported for Scinax arduous Peixoto, 2002, Scinax peixotoi Brasileiro, Haddad, Sawaya & Martins, 2007, and Scinax perpusillus (Lutz & Lutz, 1939). However, the type A call of S. littoreus is nonetheless readily distinguishable from the comparable call observed in other congeneric species. The call named type B, which exhibits a multipulsed structure, presumably has an aggressive function. We observed that different types of calls could be emitted alone or combined according to the social context. Additionally, we discuss problems involving comparisons of call parameters among species belonging to the S. perpusillus species group, provide an updated geographic distribution map, discuss the conservation status of the included species, and emphasize the importance of acoustic data for the recognition of species groups within Scinax.

Amphibian parasites of the Order Dermocystida (Ichthyosporea): current knowledge, taxonomic review and new records from Brazil

Amphibian parasites of the Order Dermocystida (Ichthyosporea) are widespread pathogens known mainly from Europe and North America, which cause primarily a disease of skin and subcutaneous tissue in their hosts. The taxonomy of these organisms has been problematic given their conserved morphology, similar clinical disease and pathology. Currently recognized taxa belong to the three closely related genera, Amphibiocystidium, Amphibiothecum, and Rhinosporidium, whereas species of Dermocystidium and Sphaerothecum destruens include fish parasites. Here, we review the taxonomy of Dermocystida based on a molecular phylogenetic analysis, principally of amphibian parasites, including DNA sequences obtained from amphibian hosts collected in the central-eastern region of South America. A new taxonomic arrangement is proposed, which includes the designation of type material for Dermocystidium pusula, synonymization of Amphibiothecum with Dermocystidium, and the restriction of Amphibiocystidium to its type species A. ranae. We also review the taxonomic status of Dermosporidium hylarum until the present work included in the synonymy of the human and animal pathogen R. seeberi, and considered herein as a valid taxon, however. In addition, a new species of Sphaerothecum parasitic to amphibians is described, being the first record of this genus in the southern hemisphere and in an amphibian host.

On the Monophyly and Relationships of Several Genera of Hylini (Anura: Hylidae: Hylinae), with Comments on Recent Taxonomic Changes in Hylids

Abstract. We present a molecular phylogenetic analysis of the hylid tribe Hylini, with the goals of testing the monophyly of the genera Duellmanohyla, Isthmohyla, and Ptychohyla and providing a discussion on the monophyly of Bromeliohyla, Charadrahyla, Ecnomiohyla, Exerodonta, Megastomatohyla, and Sarcohyla. Our results indicate the paraphyly of Ptychohyla, with Bromeliohyla and Duellmanohyla nested within it, and, as in previous analyses, the paraphyly of Duellmanohyla (due to Ptychohyla legleri and P. salvadorensis being nested within it). To resolve this situation, we restrict the contents of Ptychohyla, redelimit those of Duellmanohyla and Bromeliohyla, and erect two new genera, one to include the former Ptychohyla panchoi and P. spinipollex, and the other for the former Ptychohyla acrochorda, P. sanctaecrucis, P. zoque, and tentatively, P. erythromma. Exerodonta as currently defined is not monophyletic, inasmuch as Exerodonta juanitae is nested within Charadrahyla. Consequently, we transfer this species and, tentatively, E. pinorum to Charadrahyla. Also, we discuss some possible taxonomic problems within Exerodonta. Our results indicate that Isthmohyla is polyphyletic, the bromeliad-dwelling Isthmohyla melacaena being the sister taxon of our only exemplar of Bromeliohyla, B. bromeliacia. For this reason, we transfer I. melacaena to Bromeliohyla, rendering Isthmohyla monophyletic. The former Isthmohyla pictipes Group is shown to be paraphyletic due to having the non-monophyletic I. pseudopuma Group within it. Accordingly, we recognize a redelimited I. pseudopuma Group (contents: I. infucata and I. pseudopuma), an I. zeteki Group (contents: I. picadoi and I. zeteki), and a newly defined I. tica Group (contents: I. angustilineata, I. calypsa, I. debilis, I. graceae, I. lancasteri, I. pictipes, I. tica, I. rivularis, and, tentatively, I. insolita and I. xanthosticta). The three groups of Isthmohyla are supported by molecular evidence with jackknife support values > 90%, and two of them by putative morphological synapomorphies. We discuss the recognition of Dryophytes, Hyliola, Rheohyla, and Sarcohyla and whether it is useful to recognize Anotheca, Diaglena, and Triprion as three distinct, monotypic genera. Finally, we discuss a recent taxonomic proposal involving changes in rank and from ranked to unranked names in hylids that overall we consider to have been poorly justified and only superficially discussed.

Internal Larval Morphology of Two Species of Shining Leaf Frogs (Anura: Phyllomedusidae: Phasmahyla)

Abstract. We describe the buccal cavity and chondrocranial morphology of Phasmahyla cruzi tadpoles in comparison to P. guttata. We also provide a review of some characters for larval phyllomedusids and discuss some evolutionary aspects related to the funnel-mouthed configuration. Our discussion highlights that several characters evolved independently in several taxa with umbelliform oral discs. Nevertheless, we suggest that the dorsal orientation of the suprarostral cartilage and the presence of a prenarial ridge on the buccal roof may represent synapomorphic character states for the genus Phasmahyla.

Review of the Rare Genus Phrynomedusa Miranda-Ribeiro, 1923 (Anura: Phyllomedusidae) With Description of a New Species

Abstract:  We present the first taxonomic review of the genus Phrynomedusa since its description with diagnoses of the genus and species. We present a broad literature review of the genus and provide updates and remarks about the type series, tadpoles, calls, geographic distribution, and natural history of the species of Phrynomedusa. Additionally we describe a new species from município de São Luiz do Paraitinga, state of São Paulo, Brazil. Phrynomedusa dryade was initially identified as Phrynomedusa marginata; however, an integrated analysis of morphological and molecular characters enabled its recognition as a separate new species. For the first time, the advertisement call for one species of Phrynomedusa is described in detail. We describe the tadpole and present some field notes about the activity and biology of this new species.