Zootaxa 2418: 50–60 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

Revalidation and redescription of cesarii (Miranda-Ribeiro, 1920) (Anura: )

LUÍS FELIPE TOLEDO1,2,4, DANIEL LOEBMANN3 & CÉLIO F. B. HADDAD3 1Current address: Museu de Zoologia "Prof. Adão José Cardoso", Universidade Estadual de Campinas (UNICAMP - IB), Rua Albert Einstein s/n, Caixa Postal 6109, CEP 13083-863, Campinas, São Paulo, Brasil. E-mail: [email protected] 2Pós Graduação em Ecologia e Conservação, Universidade Federal do Paraná, Setor de Ciências Biológicas, Centro Politécnico, Jardim das Américas, Curitiba, PR, Caixa Postal 19031, CEP 81531-980 3Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Biociências, UNESP, Rio Claro, São Paulo, Caixa Postal 199, CEP 13506-970, Brasil 4Corresponding author. E-mail: [email protected]

Abstract

Elachistocleis cesarii (Miranda-Ribeiro, 1920), new combination, is revalidated and redescribed based on preserved specimens (syntype series) collected in the state of São Paulo. Furthermore, based on recently collected specimens, we increase the distribution of the species to the states of Minas Gerais and Goiás, Brazil. This species is found in open areas calling during the rainy season (from September to February for populations of the state of São Paulo). The advertisement call is described and some basic natural history data are provided. It distinguishes from all other species of the genus by the combination of the following characters: spotted belly, dominant frequency of the advertisement calls about 3.5 kHz, tympanum concealed, and presence of postcommisural glands. The advertisement call of Elachistocleis piauiensis, a similar species with spotted belly from the Caatinga Biome, was also described to compare it with the advertisement call of E. cesarii.

Key words: Amphibia; Anura; Microhylidae; Gastrophryninae; Elachistocleis cesarii

Introduction

Elachistocleis Parker, 1927 is a genus widespread distributed in South America and currently is composed by six species, which certainly needs taxonomic review. Just for an example, Nelson (1973) reported 12 different calls from 12 different populations under the name (Schneider, 1799) and (Guérin-Méneville, 1838), and suggested that these populations could represent distinct taxa. In a recent paper, Lavilla et al. (2003) clarified many questions about the Elachistocleis ovalis - E. bicolor controversy, but many other aspects remain unclear. We can recognize two groups among the species of the genus Elachistocleis: the species with spotted belly [E. surinamensis (Daudin, 1802), E. piauiensis Caramaschi & Jim, 1983, E. skotogaster Lavilla, Vaira & Ferrari, 2003, and E. erytrogaster Kwet & Di- Bernardo, 1998] and those with immaculate belly (E. bicolor and E. ovalis). In the states of São Paulo, Minas Gerais, and Goiás, from southeastern to central Brazil, there is one morphotype of Elachistocleis with spotted belly that does not match with the descriptions of the other species, but it is related to the former description of Engystoma ovale cesarii (Miranda-Ribeiro, 1920). Therefore, we here redescribe and revalidate this species comparing it with all other species of the genus. In addiction, we describe the advertisement call of Elachistocleis piauiensis, a similar species from the Caatinga Biome.

50 Accepted by M. Vences: 15 Feb. 2010; published: 6 Apr. 2010 Material and methods

Museum abbreviations of specimens are CFBH (Célio F. B. Haddad collection, deposited in the Departamento de Zoologia, UNESP, Rio Claro, São Paulo, Brazil), MNRJ (Museu Nacional, Rio de Janeiro, Brazil), MZUSP (Museu de Zoologia, USP, São Paulo, Brazil), and ZUEC (Museu de Zoologia “Prof. Adão José Cardoso”, UNICAMP, Campinas, São Paulo, Brazil). Abbreviations used in the measurements of adults are SVL (snout-vent length), AGL (axilla-groin length), HL (head length), HW (head width), ED (eye diameter), UEW (upper eyelid width), IOD (interorbital distance: distance between the inner corners of the eyes), END (eye to nostril distance), SED (snout to eye distance), IND (internarial distance), HUL (humerus length: distance from the axilla to the elbow), RUL (radio-ulna length: distance between the elbow and anterior limit of the hand), HNL (hand length: distance from the anterior limit of the hand to the longest finger), THL (thigh length), TBL (tibia length), and FL (foot length: distance from the anterior limit of the metatarsal tubercle to the longest toe). All measurements are presented in millimeters. Most of these measurements follow Duellman (1970) and Cei (1980). For morphometric measurements we used a digital caliper of 0.01 mm of precision and an ocular micrometer in a Zeiss stereomicroscope; measurements were rounded to the nearest 0.1 mm. Male’s vocalizations of Elachistocleis cesarii were recorded with a Marantz cassette tape recorder (PMD222), equipped with an external directional microphone (Audiotecnica AT835b). Male’s vocalizations of Elachistocleis piauiensis were recorded with a Sony cassette tape recorder (TCM 200DV) equipped with an external directional microphone (Yoga HT-81). For all records the microphones were positioned ca. 50 cm from the calling male and it was used chrome cassette tapes at 4.75 cm/s. The recorded calls were digitalized in PC-Desktop equipped with a Creative Sound Blaster Audigy 7.1 with 24 Bits of resolution. We analyzed the calls using Raven 1.2 software (16 bits of resolution, 44 kHz of frequency sampling, FFT and frame length of 256 samples). The terminology for the vocal analysis follows that presented in Toledo and Haddad (2005).

Results

Miranda-Ribeiro (1920; 1926) refers to some individuals deposited in the Museu Paulista (now Museu de Zoologia, University of São Paulo, Brazil) for which he suggested the name Engystoma ovale cesarii. Miranda-Ribeiro (1920) considered it a subspecies of Engystoma ovale together with other two varieties: E. o. concolor and E. o. lineatum. The original description in Portuguese of E. o. cesarii (Miranda-Ribeiro, 1920) is as follows: “Todos os demais se aproximam da segunda variedade, sem comtudo, mostrarem o miniaceo de que falla Natterer, talvez por sua permanencia no alcool, nem o ponctuado que se vê na parte dorsal da figura dada por Steindachner. Poderá ser chamado de E. – o – cesarii Iher. O colorido geral dos exemplares é o denegrido purpureo para o dorso, com ocellos ou manchas brancas na junta anterior da coxa e parte inferior, que têm por côr fundamental o pardo, havendo sempre, embora, as vezes interrompida, a linha posterior branca, das pernas e coxas. A mancha denegrida do queixo do macho apparece aqui, como tambem a observo no exemplar da var. bicolor. E de manchas brancas no dorso, ha-as esparsas na verdade, mais tão pequenas que nem são visíveis a olho nú. Isso nos exemplares magros, pois que este engorda muito.”

The individuals analyzed by Miranda-Ribeiro (1920; 1926) (numbers found in MZUSP: 33, 36–38; 42; 264; 529; 715; 2023–27) lost completely their coloration (Figure 1), turning to almost transparent by now. However, based on the morphology, distribution, and color descriptions of the syntypes it is possible to attribute the name E. cesarii (new combination) to some specimens of Elachistocleis. One of the other two subspecies described by Miranda-Ribeiro (1920, 1926) can be considered a of E. cesarii. Engystoma o. lineatum differs from E. cesarii (in the original description) by presenting a vertebral line from the snout to the vent. However, this character is one of the possible phenotypes of E. cesarii. Therefore, as there are no

REVALIDATION OF ELACHISTOCLEIS CESARII Zootaxa 2418 © 2010 Magnolia Press · 51 more differences between these subspecies, we suggest them to be synonyms, and E. cesarii have priority as appears one page before in the original publication. The remaining subspecies, Engystoma o. concolor, refers to one of the unspotted venter species according the original description; therefore it is related to other group of species that includes E. bicolor and E. ovalis.

FIGURE 1. Lectotype of Elachistocleis cesarii MZUSP 529 in dorsal (A) and ventral views (B).

Elachistocleis cesarii (Miranda-Ribeiro, 1920)

Engystoma ovale cesarii Miranda-Ribeiro, 1920 Engystoma ovale lineatum Miranda-Ribeiro, 1920

Lectotype. MZUSP 529, an adult male, collected at municipality of Piquete, state of São Paulo, southeastern Brazil (approximately 22° 36’ S; 47° 10’ W; 675 m above sea level), on November 1896 by Zech (Figure 1). Paralectotypes. – MZUSP 33 collected on January 1900 by Hempel, at Ipiranga (Ypiranga), municipality of São Paulo; MZUSP 36 (gravid female) collected on October 1896 by Bicego, at municipality of Perus; MZUSP 37 (gravid female) collected on 1897 by Bicego, at municipality of Cubatão; MZUSP 38 collected on January 1898 by Bicego, at Belém, municipality of São Paulo; MZUSP 42 (gravid female) collected on October 1907 by Ruedervalt and Cheubel at Ipiranga (Ypiranga), municipality of São Paulo; MZUSP 264 (gravid female) collected together with the lectotype; MZUSP 715 collected on 1896 by Bicego, at Alto da Serra; MZUSP 2023 (gravid female) collected on 1897 by Bicego at municipality of Cubatão; MZUSP 2024- 27 (gravid females; except for 2025 which resembles an young male) collected on October 1896 by Bicego at municipality of Perus. All municipalities are in the state of São Paulo, southeastern Brazil.

52 · Zootaxa 2418 © 2010 Magnolia Press TOLEDO ET AL. Diagnosis. Elachistocleis cesarii is recognized by the following combination of characters: 1) ventral coloration is white or yellow with gray marks and reticulations; 2) presence of postcommisural glands behind the corner of mouth; 3) tympanum concealed; 4) advertisement call with the mean dominant frequency of about 3.5 kHz.

FIGURE 2 Dorsal and ventral views of: Elachistocleis bicolor from the state of Rio Grande do Sul, Brazil (A and B, respectively); E. erythrogaster from the municipality of São Francisco de Paula, state of Rio Grande do Sul, Brazil (C and D, respectively); E. skotogaster from Aguas Blancas, provincia de Salta, Argentina (E and F, respectively).

REVALIDATION OF ELACHISTOCLEIS CESARII Zootaxa 2418 © 2010 Magnolia Press · 53 FIGURE 3. Dorsal (A) and ventral (B) views of Elachistocleis piauiensis from the municipality of Viçosa do Ceará, state of Ceará, Brazil; dorsal (C) and ventral (D) views of E. cesarii from Itapé, municipality of Rio Claro, state of São Paulo, Brazil (In C an arrow indicates the postcommisural gland); (E) dorsal view of E. surinamensis from Trinidad and Tobago; (F) gelatinous egg mass of Elachistocleis cesarii photographed in nature, where it is possible to visualize the embryos in development inside the egg capsules.

Comparison with other species. In life, Elachistocleis cesarii is readily distinguished from E. erythrogaster by the absence of the reddish coloration of the ventral portion of its body (see Figures 2 C, D and 3 C, D), its smallest body size, and the presence of a well defined femoral stripe (not well defined in E.

54 · Zootaxa 2418 © 2010 Magnolia Press TOLEDO ET AL. erythrogaster). Besides this, the frequency of the advertisement call of E. cesarii is lower than that reported to E. erythrogaster (see Kwet & Di-Bernardo 1998; 1999). E. cesarii is distinguished from E. surinamensis by its smaller size (mean of 32.3 mm in males of E. surinamensis) and by differences in body coloration (see plate XIII in Kenny 1969, and Figures 3 C, D, and E). E. cesarii is distinguished from E. bicolor and E. ovalis by the absence of the immaculate, yellow, ventral coloration (spotted in E. cesarii) (see Lavilla et al. 2003 and Figures 2 A, B and 3 C, D). E. cesarii is distinguished from E. skotogaster by presenting postcommisural glands (absent in E. skotogaster; see Figure 2 E) and by its smaller size (minimum size of 27.5 in males of E. stokogaster). E. cesarii is distinguished from E. piauiensis by some advertisement call parameters: the call frequency is lower and call duration is shorter in E. cesarii than in E. piauiensis (see Table 2). Besides this, E. cesarii has a smaller postcommisural gland (more conspicuous in E. piauiensis) and the blotches of the inguinal region are less conspicuous in E. piauiensis than in E. cesarii. In figures 2 and 3 it is possible to observe dorsal and ventral color patterns of some species of Elachistocleis in life.

TABLE 1. Measurements of the series and additional specimens recently collected of Elachistocleis cesarii. Abbreviations of the measurements are defined in the Material and Methods. Values presented as mean ± standard deviation (range). Lectotype Paralectotype Paralectotype females Other males Other females male Juvenile male (n = 8) (n = 16) (n = 3) SVL 27.8 23.2 32.24 ± 2.27 24.94 ± 1.05 29.15 ± 0.50 (29.0 – 36.0) (22.6 – 26.7) (28.6 – 29.6) AGL 11.6 9.1 15.51 ± 3.08 9.91 ± 0.85 11.97 ± 0.89 (9.4 – 19.3) (8.3 – 11.3) (11.1 – 12.8) HL 6.9 5.3 6.66 ± 0.45 5.90 ± 0.48 6.32 ± 0.34 (6.0 – 7.4) (5.2 –6.8) (6.1 – 6.7) HW 6.1 5.3 7.34 ± 0.58 6.89 ± 0.50 7,33 ± 0.29 (6.5 – 8.0) (6.1 – 8.1) (7.1 – 7.7) ED 1.2 1.0 1.34 ± 0.47 1.21 ± 0.14 1.39 ± 0.23 (1.0 – 2.4) (1.0 – 1.4) (1.2 – 1.7) UEW 1.0 0.9 0.98 ± 0.15 1.18 ± 0.15 1.10 ± 0.15 (1.0 – 1.2) (1.0 – 1.5) (0.9 – 1.2) IOD 2.9 1.8 3.32 ± 0.21 2.78 ± 0.24 3.15 ± 0.09 (2.9 – 3.5) (2.4 – 3.2) (3.1 – 3.2) END 2.1 1.6 2.48 ± 0.37 1.92 ± 0.20 2.25 ± 0.30 (2.0 – 3.0) (1.5 – 2.3) (1.9 – 2.5) SED 3.1 2.9 3.22 ± 0.24 2.92 ± 0.24 3.27 ± 0.27 (3.2 – 3.4) (2.6 – 3.4) (3.0 – 3.5) IND 1.4 0.8 1.28 ± 0.33 1.54 ± 0.10 1.70 ± 0.12 (0.8 – 1.8) (1.4 – 1.7) (1.6 – 1.8) HUL 3.4 2.6 3.22 ± 0.47 3.19 ± 0.35 3.30 ± 0.40 (2.6 – 4.0) (2.3 – 3.7) (3.0 – 3.8) RUL 5.2 3.7 5.37 ± 0.90 4.12 ± 0.38 4.41 ± 0.16 (4.4 – 6.9) (3.2 – 4.8) (4.2 – 4.5) HNL 6.0 5.3 6.96 ± 0.62 5.92 ± 0.49 6.34 ± 0.27 (6.3 – 8.3) (5.1 – 6.9) (6.1 – 6.6) THL 8.9 8.0 11.49 ± 1.10 9.20 ± 0.59 10.89 ± 0.12 (10.4 – 13.5) (7.9 – 10.2) (10.8 – 11.0) TBL 10.6 8.2 11.90 ± 0.85 9.32 ± 0.62 10.39 ± 0.45 (11.0 – 13.5) (7.7 – 10.3) (10.1 – 10.9) FL 11.1 10.0 12.78 ± 0.72 10.65 ± 0.55 12.41 ± 0.57 (11.6 – 14.0) (9.6 – 11.7) (11.8 – 12.8)

REVALIDATION OF ELACHISTOCLEIS CESARII Zootaxa 2418 © 2010 Magnolia Press · 55 FIGURE 4. Spectrograms (above) and waveforms (below) of the advertisement calls of (A) Elachistocleis cesarii recorded in the district of Itapé, municipality of Rio Claro, São Paulo, Brazil, on 29 January 2002, air temperature of 23.1ºC and (B) Elachistocleis piauiensis recorded in the municipality of Caucaia, Ceará, Brazil, on 15 May 2005, air temperature of 27 ºC.

56 · Zootaxa 2418 © 2010 Magnolia Press TOLEDO ET AL. Description of the lectotype. Body robust and ovoid. Texture of the dorsal skin and undersurface of the body smooth. Skin on ventrolateral surface of body and area around the cloacal region smooth. Cloacal flap absent; vent opening positioned posterodorsally. Head small and triangular, broader than long. Cranial crests absent. A conspicuous dorsolateral skin fold from the arms to the groin. Another skin fold crossing the chest between axillae. Eyes small, slightly prominent. Snout pointed in dorsal view and protruding in lateral view. Nostrils directed anterolaterally, slightly protuberant. Choanae relatively small and round. A well developed groove in the terminal part of the lower jaw. Internarial distance small. Canthus rostralis distinct, rounded. Loreal region slightly convex. Interorbital space slightly convex. Lips nearly horizontal when belly is touching the ground. Presence of postcommisural glands behind the corner of mouth. Tympanum concealed. Supratympanic fold absent. Upper jaw projects 1.59 mm beyond the lower jaw. Subgular vocal sac not expanded externally. Vocal slits present. Vomerine odontophores absent. Tongue large, oval. Arms and fingers robust and short. Legs extremely robust; almost equal lengths of thigh, tibia, and foot. Finger lengths I

TABLE 2. Advertisement call parameters of Elachistocleis cesarii (two males) and E. piauiensis (three males). Values presented as mean ± standard deviation (range). Data on environmental temperatures and places of recordings are in the text. E. cesarii E. piauiensis Dominant frequency (kHz) 3.52 ± 0.08 4.53 ± 0.06 (3.60 – 3.45; n = 8) (4.48 – 4.65; n = 18) Minimum frequency (kHz) 3.13 ± 0.09 3.41 ± 0.25 (3.01 – 3.23; n = 8) (3.18 – 3.76; n = 18) Maximum frequency (kHz) 4.03 ± 0.11 5.00 ± 0.09 (3.90 – 4.17; n = 8) (4.86 – 5.10; n = 18) Call duration (s) 1.93 ± 0.46 2.24 ± 0.00 (1.49 – 2.66; n = 8) (2.01 – 2.51; n = 18) Pulse durantion (ms) 4.23 ± 0.16 3,00 ± 0.23 (3.82 – 4.60; n = 50) (2.89 – 3.55; n = 50) Air temperature 23.1 °C 22 – 27 °C

Call descriptions. Advertisement calls of E. cesarii were recorded in Itapé, municipality of Rio Claro, São Paulo state, Brazil, on 29 January 2002, at air temperature of 23.1 ºC (Figure 4). The call consists of a loud, high pitched, sustained whistle, ranging from 1.49 to 2.66 s of duration, with a mean dominant

REVALIDATION OF ELACHISTOCLEIS CESARII Zootaxa 2418 © 2010 Magnolia Press · 57 frequency of 3,515 Hz, with a mean pulse rate of 238.2 per second, and 460.2 pulses per note in average. The frequency ranged from 3.01 to 4.17 kHz. The duration of the pulses did not vary much (mean of 4.23 ± 0.16 ms; n = 50). Further spectral characteristics are presented in Table 2 (see also Figure 4). Advertisement calls of E. piauiensis were recorded in the municipalities of Pacatuba on 13 March 2007 (air temperature 22 ºC), Caucaia on 15 May 2005 (air temperature 27 ºC), and Viçosa do Ceará on 24 April 2007 (air temperature 25 ºC), all of them in the state of Ceará, Northeastern Brazil. The call structure of E. piauiensis is very similar to that of E. cesarii and consist of a loud, high pitched, sustained whistle (ranging from 2.45 to 2.51 s of duration). The duration of the pulses did not vary much (3.00 ± 0.23 ms; n = 50). However, the mean dominant frequency is higher in E. piauiensis (4.53 ± 0.06 Hz) compared to the new species and the frequency ranged from 3.18 to 5.10 kHz (Table 2; Figure 4). Tadpoles. Tadpoles are unknown; however, 10 tadpoles collected in the municipality of Vitória Brasil, state of São Paulo, Brazil, and briefly described under the name Elachistocleis sp. by Rossa-Feres and Nomura (2006), could be E. cesarii. Natural history notes. Elachistocleis cesarii calls during the wet and hot season of the year (summer) as a prolonged breeder; however, after heavy rains during the hot nights it may behave like an explosive breeder (sensu Wells 1977). Males call with the legs in the water and the body out of water, holding the emergent vegetation with the forelimbs. The amplexus is axillar. We collected two amplected pairs that laid eggs in plastic bags. One female deposited 736 eggs and the second deposited 1126 eggs. In nature, the eggs are laid in the water surface as a gelatinous mass attached to floating or emergent vegetation (Figure 3F). Newly metamorphosed specimens have a mean SVL of 11.83 mm ± 0.38 (11.4 – 12.3; N = 6). The ecology of this species has been studied by Thomé and Brasileiro (2007) and a short communication about the defensive posture was published by Toledo (2004), in both articles under the name Elachistocleis cf. ovalis.

FIGURE 5. Geographic distribution map of Elachistocleis cesarii in the states of São Paulo (SP), Minas Gerais (MG), and Goiás (GO), Brazil. Gray triangle represents the type locality (municipality of Piquete, São Paulo) and black circles represent other localities based on specimens deposited in scientific collections.

58 · Zootaxa 2418 © 2010 Magnolia Press TOLEDO ET AL. Geographical distribution. Besides the localities of the type series, the species is known in the municipalities of Rio Claro, Campinas, Itirapina, São Paulo, Angatuba, and Ubatuba, all in the state of São Paulo. In the state of Minas Gerais it can be found in the municipalities of Jaboticatubas, Santana do Riacho, Munhoz, and Viçosa. In the state of Goiás it can be found in Pontalina (Figure 5). Etymology. The meaning of the specific name of E. cesarii was not stated in the original description. Discussion. The taxonomic status of Elachistocleis ovalis remains unclear. This fact limits the recognition of several related still undescribed species in South America. These species are incorrectly being referred to as E. ovalis and even E. bicolor in the past and current literature, as well as in museum collections. The lack of correct identification of all those taxa has direct implication in conservation, as some of them may be endangered or may be endemic to certain localities, countries or biomes. Therefore, a taxonomic review of the genus urges attention and may reveal a great hidden biodiversity. Additional specimens examined. Elachistocleis bicolor: CFBH 3841; 3859; 4010 São Domingos, state of Santa Catarina, Brazil; CFBH 4225 Tabapuã, state of São Paulo, Brazil; CFBH 13608-10 Três Lagoas, state of Mato Grosso do Sul, Brazil; CFBH 18199 Bom Jesus, state of Rio Grande do Sul, Brazil. E. cesarii: CFHB 4132–37; 4147–48; 4209; 4230–31; 4233–37; 4256–7 Rio Claro, state of São Paulo, Brazil; ZUEC 8054 Campinas, state of São Paulo, Brazil; ZUEC 12031 Itirapina, state of São Paulo, Brazil, CFBH 4997 Itirapina, state of São Paulo, Brazil; CFBH 3768 Pontalina, state of Goiás, Brazil; CFBH 6575-6578 Rio Claro, state of São Paulo, Brazil; CFBH 10907 Ubatuba, state of São Paulo, Brazil; CFBH 23136 Angatuba, state of São Paulo, Brazil; ZUEC 599-600 São Paulo, state of São Paulo, Brazil; ZUEC 4246-48, 4250-51, 4263 Munhoz, state of Minas Gerais, Brazil; ZUEC 1417-18, 2848, 2862-63, 3031-32, 3069, 3072-76 Jaboticatubas, state of Minas Gerais, Brazil; ZUEC 6157 (adult male recorded) Viçosa, state of Minas Gerais, Brazil; ZUEC 4995-97 Santana do Riacho, state of Minas Gerais, Brazil. E. erythrogaster: MNRJ 39098 adult male, Paratype. E. piauiensis: CFBH: 15880; 15902-03; 23465-71 Viçosa do Ceará, state of Ceará, Brazil. E. surinamensis: ZUEC 9422–23 Rapidos de Komoiran, Estado Bolivar, Venezuela.

Acknowledgments

Esteban Lavilla and Miguel Vences reviewed the manuscript making valuable suggestions. Axel Kwet provided pictures of Elachistocleis bicolor (dorsal view) and E. erythrogaster (dorsal and ventral views). John C. Murphy provided the picture of E. surinamensis (dorsal view). Diego Baldo, Santiago Nenda, and Esteban Lavilla provided the pictures of E. skotogaster (dorsal and ventral views). Igor J. Roberto provided recordings of the advertisement call of E. piauiensis. José Pombal Jr. and Hussan Zaher allowed the access to MNRJ and MZUSP, respectively. FAPESP and CNPq supported the Herpetology lab, Departamento de Zoologia, UNESP, Rio Claro, São Paulo, Brazil. Authors also thank FAPESP and CNPq for grants and scholarships.

References

Caramaschi, U. & Jim, J. (1983) A new microhylid , genus Elachistocleis (Amphibia, Anura), from northeastern Brazil. Herpetologica, 39(4), 390–394. Cei, J.M. (1980) of Argentina. Monitore Zoologico Italiano (N.S.). Monografia, 2, 1–609. Duellman, W.E. (1970) The hylid of Middle America. Monogr. Museum of Natural History of the University of Kansas, 1, 1–753. Frost, D.R. (2009) Amphibian Species of the World: an Online Reference. Version 5.3 (12 February, 2009). Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/American Museum of Natural History, New York, USA. Kenny, J.S. (1969) The Amphibia of Trinidad. In: Hummelinck, P. W. (Ed.). Studies on the fauna of Curacao and other Caribbean islands, 108, pp. 1–78. Kwet, A. & Di–Bernardo, M. (1998) Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and Environment, 33, 7–18.

REVALIDATION OF ELACHISTOCLEIS CESARII Zootaxa 2418 © 2010 Magnolia Press · 59 Kwet, A. & Di–Bernardo, M. (1999) Pró-Mata – Anfíbios. Amphibien. Amphibians, EDIPUCRS, Porto Alegre, Brasil, 107 pp. Lavilla, E.O., Vaira, M. & Ferrari, L. (2003) A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis ovalis – E. bicolor controversy. Amphibia-Reptilia, 24, 269–284. Miranda-Ribeiro, A. (1920) Os Engystomatideos do Museu Paulista (com um genero e tres especies novos). Revista do Museu Paulista, 12, 280–288, láms. 1–2. Miranda–Ribeiro, A. (1926) Notas para servirem ao estudo dos gymnobatrachios (Anura) brasileiros. Archivos do Museu Nacional do Rio de Janeiro, 27, 1–227 + 22 plates. Nelson, C.E. (1973) Mating calls of the Microhylinae: descriptions and phylogenetic and ecological considerations. Herpetologica, 29(2), 163–176. Rossa-Feres, D.C. & Nomura, F. (2006) Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of São Paulo state, Brazil. Biota Neotropica, 6(1), bn00706012006. Thomé, T.C. & Brasileiro, C.A. (2007) Dimorfismo sexual, uso do ambiente e abundância sazonal de Elachistocleis cf. ovalis (Anura: Microhylidae) em um remanescente de Cerrado do estado de São Paulo, sudeste do Brasil. Biota Neotropica, 7(1), bn00307012007. Toledo, L.F. (2004) Elachistocleis cf. ovalis (Common Oval Frog). Death feigning. Herpetological Review, 35(4), 371– 372. Toledo, L.F. & Haddad, C.F.B. (2005) Acoustic repertoire and calling site of Scinax fuscomarginatus (Anura, Hylidae). Journal of Herpetology, 39, 455–464. Wells, K.D. (1977) The social behaviour of anuran amphibians. Animal Behaviour, 25, 666–693.

60 · Zootaxa 2418 © 2010 Magnolia Press TOLEDO ET AL.