4 SPECIES OF BRAZILIAN POISON FROGS RELATED TO EPIPEDOBATES-PICTUS (DENDROBATIDAE) - TAXONOMY AND NATURAL-HISTORY OBSERVATIONS

We present morphological and natural history data on Epipedobates pictus and three additional taxa currently regarded as synonyms of the former: Epipedobates braccatus, E. flavopictus, and E. hahneli. Topotypes of the four species were examined. Morphological data on tadpoles and adults, in addition to distribution patterns and vocalizations, indicate that the four taxa represent distinct species. Two groups are evident within them: (1) the slender-bodied, Amazonian E. hahneli with small flash marks on the thighs and a low-finned tadpole, and (2) the robust bodied, almost extra-Amazonian E. braccatus, E. flavopictus, and E. pictus with large flash marks on the thighs and tadpoles with fins of moderate height.

Reproductive modes of Atlantic forest hylid frogs: A general overview and the description of a new mode

This paper reports a new reproductive mode in anurans, observed for the green treefrog, Hyla leucopygia, a species from the Atlantic Forest of Brazil and a member of the H. albosignata group. The reproductive mode of H. leucopygia is defined as follows: aquatic eggs and embryos in subterranean constructed nests; subsequent to flooding, feeding tadpoles in ponds or streams. We suggest that the new reproductive mode is widespread among the species in the H. albosignata group. The specialized courtship behaviors of H. leucopygia, characterized by vocalizations and a stereotyped sequence of mutual touches between male and female, are described. A review of the reproductive modes of hylid frogs is provided, with 11 modes recognized for the Atlantic Forest hylids and 12 modes For hylids around the world.

RESPONSE TO TEMPERATURE IN THE OXYGEN-UPTAKE OF AWAKE AND DORMANT FROGS (AMPHIBIA, LEPTODACTYLIDAE)

In southeastern Brazil the leptodactylid frogs Leptodactylus fuscus and Physalaemus fuscomaculatus enter dormancy during the dry season. Oxygen uptake was measured in awake and dormant groups of both species in a temperature range at which these frogs are usually exposed throughout the year. Oxygen uptake was lower for dormant groups at high temperature, and a lack of response to temperature was reported between 20 - 25 C in the dormant group of both species. This temperature-intensitive range can be considered an adaptive feature to save fat reserves during dormancy.

Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Four species of Brazilian poison frogs related to Epipedobates pictus (Dendrobatidae): taxonomy and natural history observations

Presents morphological and natural history data on Epipedobates pictus and three additional taxa currently regarded as synonyms of the former: Epipedobates braccatus, E. flavopictus and E. hahneli. Morphological data on tadpoles and adults, in addition to distribution patterns and vocalizations, indicate that the four taxa represent distinct species. -from Authors

Frogs of the genus Paratelmatobius (Anura: Leptodactylidae) with descriptions of two new species

Two new species of Paratelmatobius are described from the Atlantic Forest of Sao Paulo State, southeastern Brazil. The vocalization, tadpole, and natural history of one new species are described. Natural history observations of P. poecilogaster are reported, including the description of a specialized reproductive mode. Diagnoses, measurements, figures, color in life, and distributional data for the species of Paratelmatobius are provided.

Testes size in leptodactylid frogs and occurrence of multimale spawning in the genus Leptodactylus in Brazil

Although rare, some anurans show multimale spawning. Herein we present data on testes mass relative to body mass for 11 leptodactylids and report multimale spawning in Leptodactylus chaquensis and Leptodactylus podicipinus. Testes mass averaged 4.13% of body mass for L. chaquensis and 0.75% for L. podicipinus. Testes size of both leptodactylid species with multimale spawning show that they are large and close to values recorded for rhacophorid frogs with multimale breeding (0.71-7.79%). Large testes size in frogs with multimale spawning supports the sperm competition hypothesis.

Canto de anúncio de Hyla jimi e Hyla elianeae (Anura, Hylidae) na região de Botucatu, São Paulo, Brasil

In this work we present a description of the advertisement call of H. jimi, the locality type, and of H. elianeae, until now unknown in the literature. Field activities were carried out from August 1997 through June 1999; recordings were made in two open-area environments in the Botucatu region, São Paulo State. Vocalizations of 100 individuals (49 H. jimi and 51 H. elianeae) were recorded; nine characteristics of the advertisement call were examined. The advertisement calls of both species consist of consecutive series of simple notes with relatively fast repetition rates. Males of H. jimi and H. elianeae presented two patterns of note emission: one emitted by individuals beginning vocalization activity or isolated from the aggregate, and another emitted by males in chorus activity interacting with closely neighboring males. A significant difference was verified in the temporal structure of the two vocalization patterns.

Reproductive modes in frogs and their unexpected diversity in the Atlantic forest of Brazil

Amphibians, and especially the anurans (frogs and toads), exhibit a greater diversity of reproductive modes than other tetrapod vertebrates. Twenty-nine reproductive modes have been recognized for the anurans; we propose 10 more, elevating by more than 34% the number of reproductive modes known for anurans worldwide. These newly recognized reproductive modes for the frogs elevate by almost 48% the number of anuran reproductive modes known for the Neotropics. The highly complex topography of the Atlantic forest, breaking up the biome into many small microhabitats, and the high humidity, which reduces desiccation risks, have enabled the evolution of reproductive specializations such as eggs or tadpoles that develop out of water. Nearly 90% of the Atlantic forest has been cleared, and because several anurans are endemic to this region or have specialized reproductive modes dependent on the forest, this partly explains the generalized population declines and large numbers of species that have disappeared in the last few decades. © 2005 American Institute of Biological Sciences.

Size-fecundity relationships and reproductive investment in female frogs in the Pantanal, south-western Brazil

The ovarian complement of anurans exhibiting different reproductive modes is highly diverse, and intraspecific variation in reproductive output of anurans is generally related to variation in female properties and/or environmental conditions. The size-fecundity relationships, reproductive investment, and correlation between ovary mass and fat body mass were investigated for females of an anuran assemblage in the Pantanal, Brazil. Female body size was positively correlated with clutch size among seven of the eight species analyzed. However, these results seem to be influenced by seasonal variation in fecundity. Interspecific size-fecundity relationships revealed that female SVL was positively correlated with clutch size and egg size regardless of reproductive mode. Among 11 species analyzed, the reproductive investment (RI: ovary mass relative to body mass) varied from 5.5 to 18%, and there were no differences among reproductive modes and activity patterns (explosive/prolonged). RI correlated negatively with female size. Among three Leptodactylus species examined, negative correlation between ovary mass and fat body mass was verified for two species. Variations in the reproductive strategies are closely related to the reproductive activity patterns and reproductive modes exhibited by individual species, but are also influenced by environmental conditions. However, regardless of reproductive mode or activity pattern, each species seems to exhibit a reproductive strategy that allows them to respond differently to the same environmental restrictions.

Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision

Hylidae is a large family of American, Australopapuan, and temperate Eurasian treefrogs of approximately 870 known species, divided among four subfamilies. Although some groups of Hylidae have been addressed phylogenetically, a comprehensive phylogenetic analysis has never been presented. The first goal of this paper is to review the current state of hylid systematics. We focus on the very large subfamily Hylinae (590 species), evaluate the monophyly of named taxa, and examine the evidential basis of the existing taxonomy. The second objective is to perform a phylogenetic analysis using mostly DNA sequence data in order to (1) test the monophyly of the Hylidae; (2) determine its constituent taxa, with special attention to the genera and species groups which form the subfamily Hylinae, and c) propose a new, monophyletic taxonomy consistent with the hypothesized relationships. We present a phylogenetic analysis of hylid frogs based on 276 terminals, including 228 hylids and 48 outgroup taxa. Included are exemplars of all but 1 of the 41 genera of Hylidae (of all four nominal subfamilies) and 39 of the 41 currently recognized species groups of the species-rich genus Hyla. The included taxa allowed us to test the monophyly of 24 of the 35 nonmonotypic genera and 25 species groups of Hyla. The phylogenetic analysis includes approximately 5100 base pairs from four mitochondrial (12S, tRNA valine, 16S, and cytochrome b) and five nuclear genes (rhodopsin, tyrosinase, RAG-1, seventh in absentia, and 28S), and a small data set from foot musculature. Concurring with previous studies, the present analysis indicates that Hemiphractinae are not related to the other three hylid subfamilies. It is therefore removed from the family and tentatively considered a subfamily of the paraphyletic Leptodactylidae. Hylidae is now restricted to Hylinae, Pelodryadinae, and Phyllomedusinae. Our results support a sister-group relationship between Pelodryadinae and Phyllomedusinae, which together form the sister taxon of Hylinae. Agalychnis, Phyllomedusa, Litoria, Hyla, Osteocephalus, Phrynohyas, Ptychohyla, Scinax, Smilisca, and Trachycephalus are not monophyletic. Within Hyla, the H. albomarginata, H. albopunctata, H. arborea, H. boons, H. cinerea, H. eximia, H. geographica, H. granosa, H. microcephala, H. miotympanum, H. tuberculosa, and H. versicolor groups are also demonstrably nonmonophyletic. Hylinae is composed of four major clades. The first of these includes the Andean stream-breeding Hyla, Aplastodiscus, all Gladiator Frogs, and a Tepuian clade. The second clade is composed of the 30-chromosome Hyla, Lysapsus, Pseudis, Scarthyla, Scinax (including the H. uruguaya group), Sphaenorhynchus, and Xenohyla. The third major clade is composed of Nyctimantis, Phrynohyas, Phyllodytes, and all South American/West Indian casque-headed frogs: Aparasphenodon, Argenteohyla, Corythomantis, Osteocephalus, Osteopilus, Tepuihyla, and Trachycephalus. The fourth major clade is composed of most of the Middle American/Holarctic species groups of Hyla and the genera Acris, Anotheca, Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca, and Triprion. A new monophyletic taxonomy mirroring these results is presented where Hylinae is divided into four tribes. Of the species currently in Hyla, 297 of the 353 species are placed in 15 genera; of these, 4 are currently recognized, 4 are resurrected names, and 7 are new. Hyla is restricted to H. femoralis and the H. arborea, H. cinerea, H. eximia, and H. versicolor groups, whose contents are redefined. Phrynohyas is placed in the synonymy of Trachycephalus, and Pternohyla is placed in the synonymy of Smilisca. The genus Dendropsophus is resurrected to include all former species of Hyla known or suspected to have 30 chromosomes. Exerodonta is resurrected to include the former Hyla sumichrasti group and some members of the former H. miotympanum group. Hyloscirtus is resurrected for the former Hyla armata, H. bogotensis, and H. larinopygion groups. Hypsiboas is resurrected to include several species groups - many of them redefined here - of Gladiator Frogs. The former Hyla albofrenata and H. albosignata complexes of the H. albomarginata group are included in Aplastodiscus. New generic names are erected for (1) Agalychnis calcarifer and A. craspedopus; (2) Osteocephalus langsdorffii; the (3) Hyla aromatica, (4) H. bromeliacia, (5) H. godmani, (6) H. mixomaculata, (7) H. taeniopus, (8) and H. tuberculosa groups; (9) the clade composed of the H. pictipes and H. pseudopuma groups; and (10) a clade composed of the H. circumdata, H. claresignata, H. martinsi, and H. pseudopseudis groups. Copyright © American Museum of Natural History 2005.

Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae)

The known diversity of dart-poison frog species has grown from 70 in the 1960s to 247 at present, with no sign that the discovery of new species will wane in the foreseeable future. Although this growth in knowledge of the diversity of this group has been accompanied by detailed investigations of many aspects of the biology of dendrobatids, their phylogenetic relationships remain poorly understood. This study was designed to test hypotheses of dendrobatid diversification by combining new and prior genotypic and phenotypic evidence in a total evidence analysis. DNA sequences were sampled for five mitochondrial and six nuclear loci (approximately 6,100 base pairs [bp]; x=3,740 bp per terminal; total dataset composed of approximately 1.55 million bp), and 174 phenotypic characters were scored from adult and larval morphology, alkaloid profiles, and behavior. These data were combined with relevant published DNA sequences. Ingroup sampling targeted several previously unsampled species, including Aromobates nocturnus, which was hypothesized previously to be the sister of all other dendrobatids. Undescribed and problematic species were sampled from multiple localities when possible. The final dataset consisted of 414 terminals: 367 ingroup terminals of 156 species and 47 outgroup terminals of 46 species. Direct optimization parsimony analysis of the equally weighted evidence resulted in 25,872 optimal trees. Forty nodes collapse in the strict consensus, with all conflict restricted to conspecific terminals. Dendrobatids were recovered as monophyletic, and their sister group consisted of Crossodactylus, Hylodes, and Megaelosia, recognized herein as Hylodidae. Among outgroup taxa, Centrolenidae was found to be the sister group of all athesphatanurans except Hylidae, Leptodactyidae was polyphyletic, Thoropa was nested within Cycloramphidae, and Ceratophryinae was paraphyletic with respect to Telmatobiinae. Among dendrobatids, the monophyly and content of Mannophryne and Phyllobates were corroborated. Aromobates nocturnus and Colostethus saltuensis were found to be nested within Nephelobates, and Minyobates was paraphyletic and nested within Dendrobates. Colostethus was shown to be rampantly nonmonophyletic, with most species falling into two unrelated cis- and trans-Andean clades. A morphologically and behaviorally diverse clade of median lingual process-possessing species was discovered. In light of these findings and the growth in knowledge of the diversity of this large clade over the past 40 years, we propose a new, monophyletic taxonomy for dendrobatids, recognizing the inclusive clade as a superfamily (Dendrobatoidea) composed of two families (one of which is new), six subfamilies (three new), and 16 genera (four new). Although poisonous frogs did not form a monophyletic group, the three poisonous lineages are all confined to the revised family Dendrobatidae, in keeping with the traditional application of this name. We also propose changes to achieve a monophyletic higher-level taxonomy for the athesphatanuran outgroup taxa. Analysis of character evolution revealed multiple origins of phytotelm-breeding, parental provisioning of nutritive oocytes for larval consumption (larval oophagy), and endotrophy. Available evidence indicates that transport of tadpoles on the dorsum of parent nurse frogs-a dendrobatid synapomorphy-is carried out primitively by male nurse frogs, with three independent origins of female transport and five independent origins of biparental transport. Reproductive amplexus is optimally explained as having been lost in the most recent common ancestor of Dendrobatoidea, with cephalic amplexus arising independently three times. © American Museum of Natural History 2006.