Axel Kwet

High levels of diversity uncovered in a widespread nominal taxon: continental phylogeography of the Neotropical tree frog Dendropsophus minutus

Species distributed across vast continental areas and across major biomes provide unique model systems for studies of biotic diversification, yet also constitute daunting financial, logistic and political challenges for data collection across such regions. The tree frog Dendropsophus minutus (Anura: Hylidae) is a nominal species, continentally distributed in South America, that may represent a complex of multiple species, each with a more limited distribution. To understand the spatial pattern of molecular diversity throughout the range of this species complex, we obtained DNA sequence data from two mitochondrial genes, cytochrome oxidase I (COI) and the 16S rhibosomal gene (16S) for 407 samples of D. minutus and closely related species distributed across eleven countries, effectively comprising the entire range of the group. We performed phylogenetic and spatially explicit phylogeographic analyses to assess the genetic structure of lineages and infer ancestral areas. We found 43 statistically supported, deep mitochondrial lineages, several of which may represent currently unrecognized distinct species. One major clade, containing 25 divergent lineages, includes samples from the type locality of D. minutus. We defined that clade as the D. minutus complex. The remaining lineages together with the D. minutus complex constitute the D. minutus species group. Historical analyses support an Amazonian origin for the D. minutus species group with a subsequent dispersal to eastern Brazil where the D. minutus complex originated. According to our dataset, a total of eight mtDNA lineages have ranges >100,000 km2. One of them occupies an area of almost one million km2 encompassing multiple biomes. Our results, at a spatial scale and resolution unprecedented for a Neotropical vertebrate, confirm that widespread amphibian species occur in lowland South America, yet at the same time a large proportion of cryptic diversity still remains to be discovered.

Stomach-flushing for diet analysis in anurans: an improved protocol evaluated in a case study in Araucaria forests, southern Brazil*

Stomach-flushing is a gentle treatment in analysing the diet of vertebrates. We applied this method in the study of seasonal nutrition of anurans of 15 species at an Araucaria forest habitat on the Serra Geral of Rio Grande do Sul, Brazil. Over 500 frogs and toads of different body size were handled in order to evaluate an improved protocol without narcosis using soft infusion tubes in order to avoid negative effects of the flushing procedure. The specimens were treated soon after capture and then returned to the sampling sites. Our results are discussed with reference to the few studies on stomach flushing in amphibians. The improved technique is recommended to avoid killing of numerous animals for nutritional studies, in particular because of the worldwide threatening of amphibian populations.

The genus Pseudis (Anura: Pseudidae) in Rio Grande do Sul, southern Brazil, with description of a new species

A new species of Pseudis is described from Sa o Francisco de Paula, Rio Grande do Sul, Brazil. It is known from the southern parts of the Serra Geral where it occurs in grassland, inhabiting permanent ponds and still-water zones of slow e owing creeks. It is characterized by a paired vocal sac and a bulbous thumb, considerably widened at thebase. It is distinguished from P. minutusby its different call, body coloration, rounded snout, and a more robust body with shorter hindlimbs. Advertisement call, tadpole, and life history are described. The taxonomy of the family Pseudidae in Rio Grande do Sul is discussed, supporting the synonymy of Lysapsus mantidactylus and P. meridionalis with P. minutus.

Taxonomic Revision of the Scinax alter Species Complex (Anura: Hylidae)

Scinax alter, a taxon belonging to the S. ruber clade, has been previously suggested to represent a species complex. We analyzed variation among populations of Scinax alter using advertisement calls, dorsal color pattern, and external morphology. We identified three diagnosable groups distributed throughout the Atlantic Forest of eastern Brazil, which differ mainly in the advertisement call, dorsal drawing pattern, snout–vent length, and presence of tubercles on tarsus. Scinax alter was restricted to populations from south of Bahia State to Rio de Janeiro State, and two new species were related to the southern populations: Scinax imbegue, from Parque das Nascentes, Municipality of Blumenau (27°03′S, 49°05′W, 412 m a.s.l.), Santa Catarina State, Brazil, and Scinax tymbamirim, from Córrego Grande (27°35′S, 48°31′W, at sea level), Municipality of Florianópolis, Santa Catarina State, Brazil.

First record of Chaunus achavali (Anura, Bufonidae) from Rio Grande do Sul, Brazil, with a key for the identification of the species in the Chaunus marinus group

The bufonid toad Chaunus achavali, a recently described species known only from Uruguay, is recorded for Brazil. This species is morphologically similar to C. ictericus and C. arenarum, and several individuals were labeled in Brazilian scientific collections under these taxa. A lectotype of C. arenarum is designated. Additional field notes on C. achavali and a key for the identification of the species in the Chaunus marinus group are presented.

A new species of Adenomera (Amphibia: Anura: Leptodactylidae) from the Atlantic rain forest in Santa Catarina, southern Brazil

As part of the remarkable biodiversity of anurans in the southern range of the Brazilian Mata Atlântica, Adenomera engelsi sp. nov. is described here as a new taxon occurring on the island of Santa Catarina and adjacent coastal mainland. This litter-inhabiting frog is restricted to the Atlantic rain forest biome, living in primary and secondary woods and semi-open areas from sea level to about 900 m a.s.l., partly in sympatry with its congener A. araucaria. Both species are also referred to the Leptodactylus marmoratus group. Adenomera engelsi sp. nov. is a medium-sized member of the genus (snout–vent length 20.9–22.7 mm in males), characterized by a brown dorsal coloration with a maculated pattern of variable dark spots and blotches and a unique advertisement call, consisting of single, unpulsed notes with duration of 95–160 ms, dominant frequency between 3500 and 4300 Hz, and note repetition rates of 10–24 notes/min. Está descrita, como parte da enorme biodiversidade de anuros na região sul da Mata Atlântica brasileira, Adenomera engelsi sp. nov. que ocorre na Ilha de Santa Catarina e no litoral adjacente. A espécie está restrita ao bioma da Mata Atlântica, onde vive no folhiço em mata primária, secundária e áreas semi-abertas, desde o nível do mar até 900 m, parcialmente em simpatria com A. araucaria. Ambas as espécies também são atribuidas ao grupo Leptodactylus marmoratus. Adenomera engelsi sp. nov., de tamanho medio dentro do gênero (SVL 20.9–22.7 mm em machos), é caraterizada pela coloração dorsal marrom, com manchas escuras de tamanho e forma variável e um canto de anúncio distinto, que consiste de notas únicas e sem pulsos de 95–160 ms com frequência dominante entre 3500–4300 Hz e taxa de repetição de 10–24 notas por minuto.

SPATIAL, SEASONAL AND INTRAPOPULATIONAL VARIATION IN THE DIET OF PSEUDIS CARDOSOI (ANURA: HYLIDAE) FROM THE ARAUCÁRIA PLATEAU OF RIO GRANDE DO SUL, BRAZIL

Abstract The diet of Pseudis cardosoi was studied in the Araucaria forest and adjacent grassland ecosystems of southern Brazilian highlands by stomach content analysis of 119 individuals. Frogs were collected in ponds surrounded by different grassland habitats. Besides diet composition, diet diversity and volume were explored. Stomach contents were compared among samples from different habitats and collection times. Male, female, and juvenile diets were also compared. A total of 29 prey categories were identified. The most frequent items in the diet of P. cardosoi were: Diptera (15.4%), Coleoptera (14.2%), Hymenoptera (14.3%), plant remains (12.5%), Hemiptera (5.5%), Odonata naiad (2.9%), Isoptera (2.4%), and Araneae (2.3%). There were no significant differences in diet composition among areas. Significant differences in diet composition among males, females, and juveniles were detected, whereas diet diversity and diet volume did not differ significantly among these groups. Diet diversity was higher during spring and summer (January, February and October) and lower in winter (June and July). Certain items showed frequency peaks along the year, suggesting that the diet of P. cardosoi could vary in response to prey temporal dynamics.

Validation of Hylodes henselii Peters, 1870, from Southern Brazil and Description of Acoustic Variation in Eleutherodactylus guentheri (Anura: Leptodactylidae)

Abstract Comparison of advertisement calls of Eleutherodactylus guentheri from different localities in southern and southeastern Brazil revealed considerable acoustic variation among populations. Because of significant differences in call parameters, we conclude that the southern form is a distinct species, for which the name Hylodes henselii Peters, 1870, is available. Examination of the holotypes of H. henselii and Hylodes guentheri Steindachner, 1864, along with recently collected material, showed that both species are morphologically similar, possibly sibling species. Eleutherodactylus henselii is the southernmost member of the genus Eleutherodactylus occurring in subtropical Atlantic rain forest and Araucaria forest of the Brazilian states of Rio Grande do Sul and Santa Catarina, and Misiones, Argentina. Bioacoustic data suggest that at least one additional cryptic species is involved in the Eleutherodactylus guentheri group.

The Taxonomic Status of Leptodactylus geminus Barrio, 1973

The advertisement calls of Leptodactylus geminus, Leptodactylus gracilis, Leptodactylus plau- manni, and Leptodactylus fuscus are analyzed and compared with published reports. Based on calling data and examination of type material, Leptodactylus geminus is synonymized with L. plaumanni. To differentiate between the sibling species L. gracilis and L. plaumanni, fieldwork is needed, including recording of ad- vertisement calls. Reliable identification of fixed specimens is not possible based on morphology and col- oration alone. Leptodactylus plaumanni Ahl, 1936 was de- scribed from a single specimen collected in Nova Teutonia (municipality Seara, State of San- ta Catarina, southern Brazil). The holotype was originally deposited in the Deutsches Kolonial- und Ubersee-Museum Bremen and is now in the herpetological collection of the Forschung- sinstitut und Naturmuseum Senckenberg, Frankfurt (Germany). Cochran (1955), Boker- mann (1966), and Mertens (1967) treated L. plau- manni as a synonym of Leptodactylus sibilatrix, which was later considered a synonym of Lep- todactylus fuscus (Heyer, 1978). In a revision of the fuscus group, however, Heyer (1978) as- signed L. plaumanni as a synonym of Leptodac- tylus gracilis based on better matching of mor- phological characters. Heyer (1978) noted that he had been unable to examine the holotype of L. plaumanni and that this specimen could be- long to another species of the same group, namely L. geminus Barrio, 1973. The external morphology of L. geminus was described as al- most identical with that of its sibling species L. gracilis. Barrio (1973) noted only minor differ- ences in mean body length and coloration; he distinguished the new species on the basis of call characteristics. Later, Scrocchi and Lavilla (1986) suggested that the ratio of the outer metacarpal tubercle length in relation to the dis- tance from the union of fourth and fifth toes to the tip of fifth toe (Tec/Dp8) was an appropriate morphological character that could discriminate amongst both species. Cardoso (1985) collected additional material at the type locality of L. plaumanni and found a striped Leptodactylus of the fuscus group, distin-

O status taxonômico de Hyla cochranae Mertens e recaracterização de Aplastodiscus A. Lutz (Anura, Hylidae)

Hyla cochranae Mertens, 1952 is revalidated and the description of the tadpole and vocalization is provided. Based on external morphology, the genus Aplastodiscus A. Lutz, 1950 is recharacterized and a redescription of A. perviridis A. Lutz, 1950 is given. New data on geographical distribution and biology of both species are provided.