Arley Camargo

Systematics of the Neotropical Genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-molecular Evidence, and Species Accounts

Abstract. A phylogeny of the species-rich clade of the Neotropical frog genus Leptodactylus sensu stricto is presented on the basis of a total evidence analysis of molecular (mitochondrial and nuclear markers) and non-molecular (adult and larval morphological and behavioral characters) sampled from > 80% of the 75 currently recognized species. Our results support the monophyly of Leptodactylus sensu stricto, with Hydrolaetare placed as its sister group. The reciprocal monophyly of Hydrolaetare and Leptodactylus sensu stricto does not require that we consider Hydrolaetare as either a subgenus or synonym of Leptodactylus sensu lato. We recognize Leptodactylus sensu stricto, Hydrolaetare, Adenomera, and Lithodytes as valid monophyletic genera. Our results generally support the traditionally recognized Leptodactylus species groups, with exceptions involving only a few species that are easily accommodated without proposing new groups or significantly altering contents. The four groups form a pectinate tree, with the Leptodactylus fuscus group diverging first, followed by the L. pentadactylus group, which is sister to the L. latrans and L. melanonotus groups. To evaluate the impact of non-molecular evidence on our results, we compared our total evidence results with results obtained from analyses using only molecular data. Although non-molecular evidence comprised only 3.5% of the total evidence matrix, it had a strong impact on our total evidence results. Only one species group was monophyletic in the molecular-only analysis, and support differed in 86% of the 54 Leptodactylus clades that are shared by the results of the two analyses. Even though no non-molecular evidence was included for Hydrolaetare, exclusion of that data partition resulted in that genus being nested within Leptodactylus, demonstrating that the inclusion of a small amount of non-molecular evidence for a subset of species can alter not only the placement of those species, but also species that were not scored for those data. The evolution of several natural history and reproductive traits is considered in the light of our phylogenic framework. Invasion of rocky outcrops, larval oophagy, and use of underground reproductive chambers are restricted to species of the Leptodactylus fuscus and L. pentadactylus groups. In contrast, larval schooling, larval attendance, and more complex parental care are restricted to the L. latrans and L. melanonotus groups. Construction of foam nests is plesiomorphic in Leptodactylus but their placement varies extensively (e.g., underground chambers, surface of waterbodies, natural or excavated basins). Information on species synonymy, etymology, adult and larval morphology, advertisement call, and geographic distribution is summarized in species accounts for the 30 species of the Leptodactylus fuscus group, 17 species of the L. pentadactylus group, eight species of the L. latrans group, and 17 species of the L. melanonotus group, as well as the three species that are currently unassigned to any species group.

Diet of the South American frog Leptodactylus ocellatus (Anura, Leptodactylidae) in Uruguay

The diet of Leptodactylus ocellatus (L., 1758) was studied in specimens collected at Maldonado Department (Uruguay), where monthly surveys were made between August 1998 and March 2000. A total of 143 frogs were collected, measured, sexed, and dissected, to assess stomach contents. The anurans were grouped into three age classes and four categories (considering sex and capture date). The trophic amplitude for each age class and sex-season category was quantified using Shannon-Weaver Index, and the trophic niche overlap between ages and categories by using the Pianka Index. Comparisons among treatments were made by G-tests. The most important prey items were coleopterans (IRI = 2547), and significant correlations were found between predator and prey sizes. The major differences on diet composition were found between extreme age classes (froglets and adults). Significant differences were also detected between sexes in the cold season but not in the warm season. Any of these changes in diet may be related with the availability of prey.

Environmental and seasonal variation in the diet of Elachistocleis bicolor (Guérin-Méneville 1838) (Anura: Microhylidae) from northern Uruguay.

Abstract We studied the diet of Elachistocleis bicolor captured in pine, eucalypt, and unmodified environments in Uruguay. Comparisons between seasons (active, inactive) and among three environments (pine, eucalypt, unmodified), were made using rarefaction analyses, importance indices, and non-parametric tests. Elachistocleis bicolor has a specialized diet composed mainly of Pheidole and Solenopsis ants and termites. The diet of E. bicolor includes a high number of prey per individual, suggesting active search as a foraging strategy. The consumption of myrmicine ants (i.e., Solenopsis) represents a source for potential toxic skin secretions that in E. bicolor may be used to avoid being attacked in the ant nests used for shelter during aestivation, as occurs in the microhylid Phrynomantis microps. Diets in forested environments were richer in both periods, which may reflect the colonization of opportunistic ant species in these environments. Ants were more important in the eucalyptus plantations, particularly in the inactive period, whereas termites were more important in the pine plantations mainly in the inactive period. These environmental and seasonal differences in diet are consistent with the temperature and humidity tolerances of ants and termites, which are the main prey of E. bicolor.

Trophic Niche Variation and Individual Specialization in Hypsiboas pulchellus (Duméril and Bibron, 1841) (Anura, Hylidae) from Uruguay

ABSTRACT. Intrapopulation variation includes age and sexual variation, but also individual niche specialization. Individual specialization has important effects on dynamics at population or community levels. Environmental variation and intraspecific interactions are factors that can promote or enhance this kind of variation. Despite its relevance, knowledge about individual specialization in amphibians is still scarce, especially in species from temperate regions. In this study we analyzed the intrapopulational variation of the trophic niche in Hypsiboas pulchellus from a population in southern Uruguay. Particularly, we analyzed ontogenetic and sexual variation and individual specialization of the diet. We found ontogenetic variation in diet as a result of a shift from flies to beetles with increasing body size of frogs, but there were no differences in diet between males and females. We also found evidence of individual specialization, which varied seasonally together with population trophic niche width; during the warm season (October–March), individual specialization and population TNW both increased.

High genetic diversity but low population structure in the frog Pseudopaludicola falcipes (Hensel, 1867) (Amphibia, Anura) from the Pampas of South America.

Relative to South Americas ecoregions, the temperate grasslands of the Pampas have been poorly studied from a phylogeographic perspective. Based on an intermediate biogeographic setting between subtropical forest (Atlantic Forest) and arid ecosystems (Chaco and Patagonia), Pampean species are expected to show unstable demographic histories due to the Quaternary climatic oscillations. Herein, we investigate the phylogenetic relatedness and phylogeographic history of Pseudopaludicola falcipes, a small and common frog that is widely distributed across the Pampean grasslands. First, we use molecular data to assess if P. falcipes represents a single or multiple, separately evolving cryptic lineages. Because P. falcipes is a small-size species (<20mm) with extensive coloration and morphological variation, we suspected that it might represent a complex of cryptic species. In addition, we expected strong genetic and geographic structuring within Pseudopaludicola falcipes due to its large geographic distribution, potentially short dispersal distances, and multiple riverine barriers. We found that P. falcipes is a single evolutionary lineage with poor geographic structuring. Furthermore, current populations of P. falcipes have a large effective population size, maintain ancestral polymorphisms, and have a complex network of gene flow. We conclude that the demographic history of P. falcipes, combined with its ecological attributes and the landscape features of the Pampas, favored a unique combination among anurans of small body size, large population size, high genetic variability, but high cohesiveness of populations over a wide geographic distribution.