Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru.

Molecular phylogenetic analysis revealed the existence of two undescribed species of the hemiphractid genus Gastrotheca in the Andes in northern Peru. Both species are similar morphologically to Gastrotheca dysprosita and G. monticola, but they differ from these species and from one another in subtleties of coloration and minor variances in size and proportions. Gastrotheca aguaruna sp. nov. (6˚10'50"S, 77˚37'01"W, 2480 m) is from humid forested areas in the northern part of the Cordillera Central, whereas G. aratia sp. nov. (6˚14'00"S, 78˚51'24"W, 2560 m ) is known from the northern part of the Cordillera Occidental.

Brazilian marsupial frogs are diphyletic (Anura: Hemiphractidae: Gastrotheca).

Molecular phylogenetic analyses based on expanded taxonomic and geographic sampling support the monophyly of the marsupial frog genera (family Hemiphractidae), resolve six geographically circumscribed lineages within Gastrotheca, and, for the first time, reveal that two divergent lineages of Gastrotheca inhabit the Atlantic Coastal Forests of Brazil. Within Gastrotheca, the earliest diverging clade is confined to northeastern Brazil, whereas the three subsequent diverging lineages are restricted to northern Venezuela (G. walkeri), southeastern Brazil, and northwestern South America. All species in these clades inhabit humid forests at low to mid-elevations, and their life histories are characterized by lacking free-living tadpoles (i.e., direct development). Two derived clades inhabit the Andes, and both contain species with either direct development or tadpoles. One Andean clade of Gastrotheca ranges in the high Andes from Colombia to extreme northern Peru, whereas the other clade inhabits high elevations in the Andes of southern Ecuador, Peru, and Bolivia, and lower elevations in the Andes of northwestern Argentina. The presence of two non-sister lineages on each side of the Amazon Basin suggests that vicariance across this central region played an important role in diversification within Gastrotheca.

Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species.

We review the systematics of the species of Gastrotheca (Anura: Hemiphractidae) in the highlands of the southern Ecuadorian Andes. We analyzed phylogenetic, morphological, ecological, and acoustic data from populations in the region. We provide an updated phylogenetic hypothesis inferred from a database that contains 42 species of Gastrotheca, and sequences from mitochondrial (12S, 16S, and ND1) and nuclear (POMC and RAG-1) genes. We describe four new endemic biphasic species for Ecuador (Gastrotheca cuencana, G. elicioi, G. turnerorum, and G. yacuri). In addition, we redefine G. lojana, recently resurrected from the synonymy of G. monticola, and redescribe G. litonedis. Furthermore, we describe the tadpoles and ontogenetic color changes in six species (G. cuencana, G. elicioi, G. litonedis, G. lojana, G. pseustes, and G. turnerorum). Also, we describe the calls of seven species (G. cuencana, G. elicioi, G. litonedis, G. lojana, G. pseustes, G. testudinea, and G. yacuri). The phylogenetic relationships estimated here are congruent with previous phylogenetic hypotheses for the group, except for the placement of G. galeata, G. plumbea, G. orophylax, G. nicefori, and G. griswoldi. Because most sister species in Gastrotheca are allopatric, the evolutionarly scenarios that likely have intervened in shaping the diversity are the uplift of the Andes and the formation of cross-Andean river valleys, which probably promoted conditions that acted as dispersal barriers that led to speciation. Many species of Gastrotheca (44%) are highly threatened. Most monophasic species have not been seen since the late 1980s, whereas the highland biphasic species have survived the catastrophic events that led to the disappearance of many other sympatric anurans. Research and conservation actions are urgently needed for all surviving species treated herein, five of which are either Critically Endangered or Endangered because of their restricted distributions (10-1600 km2), habitat destruction and fragmentation, climate change, and pathogens. Two of the new species, G. cuencana, and G. elicioi, mostly are known from the cities of Cuenca and Loja, respectively, where urbanization threatens their survival. In order to call attention to conservation issues, we name one new species to refer to the Yacuri National Park, and the other three in honor of the people from Cuenca, the Turner family, and Elicio E. Tapia, whose actions are helping to halt amphibian extinctions.

Advertisement calls and DNA sequences reveal a new species of Scinax (Anura: Hylidae) on the Pacific lowlands of Ecuador.

Scinax is a speciose genus of Neotropical hylid frogs. We describe a new species from western Ecuador (elevations between 0 and 1207 m) using morphology, vocalizations, and DNA sequences. We also present a new phylogeny for Scinax based on mitochondrial DNA genes 12S rRNA, Cytochrome Oxidase sub-unit I, Cytochrome B, 16S rRNA, NADH dehydrogenase subunit 1, and adjacent tRNAs. The new species, Scinax tsachila sp. nov. was previously confused with S. quinquefasciatus, a morphologically similar sympatric species. They differ by having markedly different advisement calls, distinct skin texture in the dorsum, and different bone coloration. The new species is sister to S. elaeochroa, a species that differs in advertisement call and color pattern. We provide an updated species account for Scinax quinquefasciatus and a redescription of its holotype.

Loss and re-evolution of complex life cycles in marsupial frogs: does ancestral trait reconstruction mislead?

Using phylogeny-based methods to identify evolutionary transitions has become an integral part of evolutionary biology. Here, we demonstrate the potential for these methods to give statistically well-supported but misleading inferences about character evolution. We also show how inferences of character evolution can be informed using GIS-based methods to reconstruct ancestral environmental regimes. We reconstruct a phylogeny for marsupial frogs (Hemiphractidae) using nuclear and mitochondrial DNA sequences and estimate patterns of life-history evolution across the resulting tree. We find that Gastrotheca species with complex life cycles (i.e., egg, tadpole, and adult stages) are phylogenetically nested among species and genera with direct development (i.e., egg and adult stages only). Assuming a single rate for gains and losses in likelihood reconstructions, there is strong statistical support for the hypothesis that the tadpole stage was lost early in the phylogeny but reappeared within Gastrotheca. Assuming different rates of gain and loss, the model with significantly higher statistical support, the tadpole stage seems to have been lost multiple times but never regained. Given that both hypotheses cannot be correct, at least one reconstruction model must be giving well-supported but misleading results. Several lines of evidence (including GIS-based reconstructions of the ancestral climatic regime) suggest that the former hypothesis is correct, and that the tadpole stage has evolved from direct development within Gastrotheca, the only known case of such a reversal in frogs.

Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae).

A phylogenetic analysis of sequences from 503 species of hylid frogs and four outgroup taxa resulted in 16,128 aligned sites of 19 genes. The molecular data were subjected to a maximum likelihood analysis that resulted in a new phylogenetic tree of treefrogs. A conservative new classification based on the tree has (1) three families composing an unranked taxon, Arboranae, (2) nine subfamilies (five resurrected, one new), and (3) six resurrected generic names and five new generic names. Using the results of a maximum likelihood timetree, times of divergence were determined. For the most part these times of divergence correlated well with historical geologic events. The arboranan frogs originated in South America in the Late Mesozoic or Early Cenozoic. The family Pelodryadidae diverged from its South American relative, Phyllomedusidae, in the Eocene and invaded Australia via Antarctica. There were two dispersals from South America to North America in the Paleogene. One lineage was the ancestral stock of Acris and its relatives, whereas the other lineage, subfamily Hylinae, differentiated into a myriad of genera in Middle America.

Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal.

Approximately one-half of all species of amphibians occur in the New World tropics, which includes South America, Middle America, and the West Indies. Of those, 27% (801 species) belong to a large assemblage, the eleutherodactyline frogs, which breed out of water and lay eggs that undergo direct development on land. Their wide distribution and mode of reproduction offer potential for resolving questions in evolution, ecology, and conservation. However, progress in all of these fields has been hindered by a poor understanding of their evolutionary relationships. As a result, most of the species have been placed in a single genus, Eleutherodactylus, which is the largest among vertebrates. Our DNA sequence analysis of a major fraction of eleutherodactyline diversity revealed three large radiations of species with unexpected geographic isolation: a South American Clade (393 sp.), a Caribbean Clade (171 sp.), and a Middle American Clade (111 sp.). Molecular clock analyses reject the prevailing hypothesis that these frogs arose from land connections with North and South America and their subsequent fragmentation in the Late Cretaceous (80-70 Mya). Origin by dispersal, probably over water from South America in the early Cenozoic (47-29 million years ago, Mya), is more likely.

Oral structures and their development in egg-brooding hylid frog embryos and larvae: Evolutionary and ecological implications.

Among egg-brooding hylid frogs there is much interspecific variation in the degree of development of the young at hatching. In certain species of Gastrotheca the eggs hatch into free-living tadpoles, whereas in others (and in the genera Amphignathodon, Cryptobatrachus, Stefania and Hemiphractus) the eggs hatch directly into frogs. We examined the oral anatomy of tadpoles and embryos of 22 species of egg-brooding hylids in order to determine the morphological differences between free-living larvae and embryos of species having direct development. All free-living Gastrotheca larvae are morphologically similar and have a large array of oral structures directly associated with a suspension feeding way of life. Among those egg-brooding hylids without free-living larvae there is a complete gradation from those with all of the free-living tadpole oral structures to those with none. Different lineages retain different vestiges of free-living larval morphology, suggesting that direct development has evolved multiple times among these frogs. All of the morphological patterns in the direct-developing embryos can be accounted for by simple truncation or acceleration of the normal tadpole developmental program. We explore the possibility that certain Gastrotheca species with tadpoles may have evolved from species that lack larval stages. The development of oral structures in egg-brooding hylids provides insight into the phylogenetic significance of these charactes in other groups of anurans. Most significantly they reinforce the idea that microhylids evolved from ranoidlike ancestors.