ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah Acta Herpetologica 7(2): 297-308, 2012 Advertisement call of Scinax littoralis and S. angrensis (Amphibia: Anura: Hylidae), with notes on the reproductive activity of S. littoralis Michel V. Garey1, Thais R.N. Costa2, André M.X. de Lima2, Luís F. Toledo3, Maríl- ia T. Hartmann4 1 Departamento de Zoologia e Botânica, Instituto de Biociências, Letras e Ciências Exatas, Universi- dade Estadual Paulista, Brazil. Corresponding author. E-mail: michelgarey@gmail.com 2 Programa de Pós-graduação em Ecologia e Conservação, Universidade Federal do Paraná, Brazil 3 Museu de Zoologia “Prof. Dr. Adão José Cardoso”, Universidade Estadual de Campinas, Brazil 4 Universidade Federal da Fronteira Sul, Campus de Erechim, Brazil Submitted on: 2012, 15th May; revised on: 2012, 13th October; accepted on: 2012, 13th October. Abstract. Scinax littoralis and S. angrensis are poorly known Atlantic forest endemic species from the Scinax catharinae group, which is known from southern and south- eastern Brazil. Herein, we describe the advertisement calls of these two species and compare them to other species within the S. catharinae group. Additionally, we pro- vide information on breeding sites, and calling activity in S. littoralis. Advertisement calls of S. angrensis and S. littoralis are composed of multi-pulsed notes with multiple frequency bands and frequency modulation. Scinax littoralis has a higher minimum and maximum frequency, and lower dominant frequency than S. angrensis, while Scinax littoralis has a longer call. Scinax littoralis appears to be a habitat generalist, occurring from secondary to mature forests, and breeding both in temporary and in permanent ponds. Male S. littoralis were usually observed calling perched on shrub stems, and displayed satellite behavior when the male density was high. Keywords. Bioacoustics, breeding activity, satellite behavior, vocalization, treefrog. INTRODUCTION Vocalization is the most widespread means of communication for frogs, especially for mate attraction (Ryan, 2001). Call types are usually categorized into mating or advertise- ment, territorial, release, distress, and warning calls (Bogert, 1960), but for several species, one signal type may communicate many things to conspecifics (Wells, 2007). The adver- tisement call is mainly a reproductive signal, but may also provide information about spe- 298 Michel V. Garey et al. cies recognition or territory defense (Ryan, 2001; Wells, 2007). Because frogs have spe- cies-specific calls, the study of acoustic signals can be useful for researchers: field research dependent on auditory surveys (e.g., line transects, Zimmerman, 1993), or research focus- ing on behavior (Bee et al., 1999), natural history (Lima et al., 2010) or taxonomic goals (e.g., Cocroft and Ryan, 1995; Pombal et al., 1995; Toledo et al., 2007) can all capitalize on differences in calls among species to gain information relevant to a specific study. Both Scinax littoralis and S. angrensis belong to S. catharinae group, which is with- in the S. catharinae clade (Faivovich, 2002). Both species are endemic to the Brazilian Atlantic forest and are morphologically similar (cryptic species) and apparently allopatric. However, their life history and ecology remain poorly known. The distribution of S. lit- toralis ranges from southern portion of the state of São Paulo (Pombal and Gordo, 1991) to the state of Paraná (Conte et al., 2009) and S. angrensis may be found from the state of Rio de Janeiro (Carvalho-e-Silva et al., 2008) to the northeastern portion of the state of São Paulo (Araújo et al., 2009). The reproductive season of S. littoralis is prolonged, with breeding occurring throughout the year (Narvaes et al., 2009). Males of this species usually call from high perches over permanent ponds (Narvaes et al., 2009). Females dis- play two distinct reproductive modes: eggs may be deposited in ponds or in bromeliad leaf axils, and females have been observed depositing eggs in temporary ponds (Toledo et al., 2012). Scinax angrensis males have been recorded calling around temporary and per- manent ponds and around stream zones (Carvalho-e-Silva et al., 2008). One reproductive mode was observed: females lay eggs in ponds (Hartmann et al., 2010). Herein, we describe the advertisement calls of both S. littoralis and S. angrensis and provide novel data on the reproductive and calling behavior of S. littoralis. These calls are compared to other closely related species, which may provide insight into relationships within the group. MATERIAL AND METHODS Call description The advertisement call of S. littoralis was recorded in Candonga Village (25⁰35’26”S, 48⁰48’47”W, 230 m a.s.l.), in the municipality of Morretes (Fig. 1), state of Paraná, in November 2010, at 2230h – 0100h (air temperature = 23 °C). The advertisement calls of this species were recorded with a TASCAM DR-08 digital recorder with external microphone Seinnheser ME66. The advertisement call of S. angrensis was recorded in the Serra do Mar State Park (23°21’53”S, 44°49’27”W, 50 m a.s.l.), municipality of Ubatuba (Fig. 1), state of São Paulo, in January 1994, at 2000h-2130h (air temperature = 23.5°C). The advertisement calls of this species were recorded with a NAGRA-E tape recorder with an external microphone Seinnheser ME66. Calls were digitalized and analyzed in Raven Pro 1.3 (Cornell Bioacoustics Research Pro- gram) at 16 bits, 44 kHz, a FFT frame length of 512 samples, and Hann window type. The bright- ness was adjusted to 68%, the contrast was adjusted to 80%, and overlap was adjusted to 50%. The terminology used in these descriptions follows Toledo and Haddad (2005a). For S. littoralis, 21 advertisement calls of three males were analyzed (n = 75 notes), and for S. angrensis 17 calls of two individuals were analyzed (n = 43 notes). We analyzed the number of notes per call, number of fre- quency bands, call duration, note duration, number of pulses per note, pulse rate, minimum, maxi- 299Advertisement call of Scinax mum and dominant frequency, inter-note interval and call rate. To analyze the difference in inter- note intervals between S. littoralis advertisement calls, and to determine if there were differences in call parameters among S. littoralis and S. angrensis we used a Student’s t-test. All variables were also tested for normality and variance homogeneity statistical analysis. Data are presented as mean ± standard deviation. The advertisement calls of Scinax littoralis (FNJV 12855-12862) and S. angrensis (FNJV 12863) were deposited in Fonoteca Neotropical “Jacques Vielliard”, Instituto de Biologia, Universi- dade Estadual de Campinas, Campinas, state of São Paulo, Brazil. Voucher specimens of S. littoralis were deposited in the Coleção Herpetológica of the Museu de Zoologia Universidade de São Paulo, São Paulo, Brazil (MZUSP 137947-48). We collected individuals from the same population of S. ang- rensis, however, we did not collect the specimens from which the advertisement calls were recorded. The specimens of the population of S. angrensis from Serra do Mar State Park, nucleus Picinguaba, were deposited in the amphibian collection of Célio F. B. Haddad, Departamento de Zoologia, Insti- tuto de Biociência, São Paulo State University, Rio Claro, Brazil (CFBH 7912-7916). Fig. 1. Map of Southern Brazil showing the municipalities of species type-localities and study sites. White circle: Scinax angrensis type-locality; black circle: S. angrensis study-site; black square: Scinax littoralis type-locality; white and gray square: S. littoralis study-sites. 300 Michel V. Garey et al. Reproductive activity of S. littoralis The reproductive behavior of S. littoralis was studied at Reserva Natural Salto Morato (RNSM; 25º09’S; 48º16’-48º20’W), in the municipality of Guaraqueçaba (Fig. 1), on the north coast of the state of Paraná, southern Brazil. RNSM is composed of a mosaic of landscapes due to its land use history. The protected area cover comprises ~2.500 ha of Atlantic forest, with altitude ranging from 25 to 930 m a.s.l. In this region, the climate according to the Köppen-Geiger classification (Peel et al., 2007) is of the type Cfa: temperate climate with no dry season, and an average annual tempera- tures of around 21 °C. Annual rainfall is elevated; with more than 2,000 mm y-1 and mean relative humidity is 85% (Maack, 2002). We sampled nine lentic water bodies, one per night, using the visual and acoustic survey at breeding sites technique (Scott and Woodward, 1994). Data were collected between September 2006 and April 2007. Surveys generally began at 1900 h and were finished at midnight. We searched for frogs at swamps, temporary and permanent ponds in early secondary forest, late secondary forest and mature forest (see Gatti, 2000), three water bodies in each successional forest stage. The water bodies were located between 20 to 200 m a.s.l. For each male found calling, we measured the perch height and the horizontal distance to the margin of pond. In each survey area, the number of calling males was estimated by counting all individuals in each surveyed area by walking around the perim- eter of each breeding site (Vasconcelos and Rossa-Feres, 2005). Males were observed for approxi- mately one hour per night using animal focal sampling (Altmann, 1974). Differences in perch height and distance to ponds margin among the forest succession stag- es were analysed using ANOVA. Differences among treatments were determined using the Tukey- Kramer HSD test. RESULTS Call description The advertisement call of S. angrensis is composed by 1-7 multi-pulsed notes that pre- sent multiple frequency bands and an irregular frequency modulation (Table 1). The mean duration of the advertisement call is 0.42 ± 0.1 s, the mean note duration is 0.025 ± 0.012 s (first note = 0.025 ± 0.009 s, second note = 0.034 ± 0.011 s, third note = 0.015 ± 0.011 s, fourth note = 0.023 ± 0.024 s, fifth note = 0.034 s, sixth note = 0.005 s, and seventh note = 0.009 s), with 17.88 ± 5.18 pulses per note (first note = 12.6 ± 0.009, second note = 14.4 ± 5.6, third note = 8.7 ± 8.9, fourth note = 10.5 ± 12, fifth note = 15, sixth note = 4, and seventh note = 6 pulses per note). The inter-note interval was 50.13 ± 15.74 ms (23-76, n = 23). Notes have 2-4 frequency bands (Fig. 2). The minimum frequency of the fundamental band is 1.67 ± 0.23 kHz, the maximum frequency of the highest band is 4.51 ± 0.69 kHz, while the dominant frequency is 3.13 ± 0.37 kHz, which is the second band (first note = 3.29 ± 0.33 kHz, second note = 3.39 ± 0.23 kHz, third note = 3.17 ± 0.45 kHz, fourth note = 3.01 ± 0.24 kHz, fifth note = 2.76 kHz, sixth note = 2.41 kHz, and seventh note = 3.49 kHz). The pulse rate is 575 ± 7.5 pulses/s, and pulse rate was higher in the middle of the notes. The call rate was 16 calls/min, and note rate was 0.072 note/s. The last note tends to be shorter than the others when the call is composed of three or more notes. The advertisement call of S. littoralis consists of 3-4 multi-pulsed notes, with 3-10 simultaneously frequency bands (Table 1). The mean duration of the advertisement call is 301Advertisement call of Scinax Ta bl e 1. C al l p ar am et er s of t he S ci na x ca th ar in ae s pe ci es g ro up . N n: n um be r of n ot es /c al l; D ur at io n: d ur at io n of t he a dv er tis em en t ca ll; P ul se s/ no te : n um be r of p ul se s pe r no te ; P ul se s/ s: r at e of p ul se s pe r se co nd ; F m ax : m ax im um fr eq ue nc y; F m in : m in im um fr eq ue nc y; F do m : d om in an t f re qu en cy ; S tr uc : s tr uc tu re o f ca ll, p re se nc e of p ul se s = P an d/ or p re se nc e of h ar m on ic s = H ; N fb = n um be r of fr eq ue nc y ba nd s. D at a ar e sh ow n as : m ea n ± st an da rd d ev ia tio n (r an ge , n ). N n D ur at io n (s ) Pu ls es /n ot e Pu ls es /s Fm ax ( kH z) Fm in ( kH z) Fd om ( kH z) St ru c N fb R ef er en ce Sc in ax a ng re ns is (1 -7 ) 0. 42 ± 0 .1 (0 .1 7- 0. 7, n = 43 ) 17 .8 8 ± 5. 18 (7 -2 8, n = 3 3) 43 4. 65 ± 6 3. 28 (2 89 .4 7- 54 2. 86 , n = 33 ) 4. 51 ± 0 .6 9 (3 .1 2- 5. 75 ; n = 75 ) 1. 67 ± 0 .2 3 (1 .1 1- 2. 13 ; n = 75 ) 3. 13 ± 0 .3 7 (2 .1 5- 3. 7, n = 42 ) P/ H 2- 8 Pr es en t s tu dy S. li tt or al is (3 -4 ) 0. 55 ± 0 .1 2 (0 .2 1- 0. 79 ; n = 75 ) 23 .6 8 ± 2. 73 (8 -3 0, n = 3 7) 46 2. 49 ± 12 9. 43 ( 31 4. 29 - 10 00 , n = 3 7) 4. 65 ± 0 .7 9 (2 .8 7- 5. 57 , n = 42 ) 2. 2 ± 0. 31 (1 .4 7- 2. 79 , n = 42 ) 2. 65 ± 0 .2 9 (1 .8 9- 3. 53 ; n = 75 ) P/ H 3- 10 Pr es en t s tu dy S. a gi lis ( no te “ a” ) 2 0. 39 ± 0 .7 6 (0 .3 8- 0. 40 , n = 12 ) 85 .6 7 ± 5. 07 (7 4- 96 , n = 1 2) - - - 7. 66 ± 0 .1 8 (7 .4 5- 7. 92 , n = 12 ) P 0 N un es e t a l., 2 00 7 S. a gi lis ( no te “ b” ) 2 2. 5 ± 5. 3 (1 .0 6- 3. 2, n = 10 4) 7. 15 ± 1 .2 2 (5 -1 1, n = 1 04 ) 7. 02 ± 0 .4 8 (5 .6 -7 .8 8, n = 10 4) P 0 S. a lb ic an s 1 0. 7 - - 4. 2 2. 8 (3 .3 -4 .1 ) P/ H - Po m ba l e t a l., 1 99 5* S. a rg yr eo rn at us (5 -2 80 ) (0 .8 -2 5) (2 -2 5) - 9. 0 3. 6 (5 .0 -6 .3 ) P 0 Po m ba l e t a l., 1 99 5 S. c an as tr en si s (6 -7 ) - - - 5. 6 1. 4 - P 0 C ar do so a nd H ad da d, 19 82 S. c at ha ri na e - > 2 - - 3. 1 2. 2 - P/ H - Po m ba l e t a l., 1 99 5* S. c en tr al is 2 0. 37 ± 0 .0 6 (n = 4 ) - - 6. 2 2. 8 (3 .2 -4 .6 ) P 0 Po m ba l a nd B as to s, 19 96 S. h ie m al is 2 - - - 7. 0 2. 0 (2 .2 -4 .3 ) P/ H (2 -4 ) H ad da d an d Po m ba l, 19 87 S. m ac ha do i (6 -7 ) 0. 05 - - 5. 2 0. 15 3. 5 H - B ok er m an n an d Sa zi m a, 1 97 3 S. r an ki (4 -5 ) - - - 5. 5 1. 5 - P 0 A nd ra de a nd C ar do so , 19 87 S. r iz ib ili s (7 -2 3) (1 .2 9- 2. 74 ) (1 5- 72 ) - 5. 5 2. 0 (2 .8 -4 .0 ) P 0 Po m ba l e t a l., 1 99 5 S. s ka io s (4 2- 73 ) (4 .4 2- 7. 9) 9. 2 ± 3. 33 (5 -1 6, n =1 0) - - - (2 .2 1- 2. 24 ) P 0 Po m ba l e t a l., 2 01 0 * D at a of o th er a ut ho rs p re se nt ed b y Po m ba l e t a l. 19 95 . 302 Michel V. Garey et al. 0.55 ± 0.12 s, the mean note duration is 0.050 ± 0.013 s (first note = 0.054 ± 0.011 s, sec- ond note = 0.053 ± 0.001 s, third note = 0.047 ± 0.016 s, and fourth note = 0.023 ± 0.001 s), with 23.68 ± 2.73 pulses per note (first note = 21.9 ± 4.17, second note = 22.43 ± 2.11, third note = 18.53 ± 6.81, fourth note = 7 ± 1 pulses per note). The minimum frequen- cy of the fundamental band is 2.20 ± 0.31 kHz, the maximum frequency of the highest frequency band is 4.65 ± 0.79 kHz, whereas the dominant frequency is 2.65 ± 0.29 kHz, which is the third band (first note = 2.58 ± 0.26 kHz, second note = 2.72 ± 0.20 kHz, third note = 2.67 ± 0.20 kHz, and fourth note = 2.47 ± 0.19 kHz). The first two notes were simi- Fig. 2. Waveform (above) and spectrogram (below) of the advertisement calls of S. angrensis (air tempera- ture = 23.5 °C), and Scinax littoralis (air temperature = 23 °C). 303Advertisement call of Scinax lar in irregular frequency modulation. The third note varied: in some cases, it was modu- lated like the first two notes, while in others it was of relatively constant frequency. The pulse amplitude increased across the note. The interval between the first and second notes was 41.69 ± 9.43 ms (24-66; n = 51), and the interval between the third and fourth note was 43.22 ± 8.33 ms (27-63; n = 23). The call rate was 6.37 ± 2.73 call/min (4.8-9.52; n = 3), and note rate was 0.082 note/s. The advertisement call of S. angrensis and S. littoralis present similarities in the struc- ture but some parameters differed. Scinax littoralis has a greater call length (t = 5.97; P = 0.0001, df = 120), note length (t = 9.49; P = 0.0001; df = 120), and number pulses per note (t = 7.20; P = 0.0001; df = 120), than S. angrensis. However, S. angrensis has a higher dominant frequency (t= 4.88; P = 0.0001; df = 120), and lower minimum frequency (t = -10.86; P = 0.0001; df = 120), maximum frequency (t = 2.32; P = 0.022; df = 120), in rela- tion to S. littoralis. These species did not differ in relation to notes per call (t = -0.06; P = 0.949; df = 120), and number of frequency bands (t = 1.38; P = 0.069; df = 120). Fig. 3. Horizontal distance from the water of the perches used for calling by males Scinax littoralis in three vegetation succession stage sites in the Reserva Natural Salto do Morato, municipality of Gua- raqueçaba, state of Paraná, Brazil. Positive values indicate that a calling site was located outside of the breeding pool; negative values indicate that a calling site was located within a pool; zero value is the mar- gin of the pool. Dots indicate the means and bars indicate standard deviation. Early = early secondary for- est; Late = late secondary forest; Old-growth = old growth forest. 304 Michel V. Garey et al. Reproductive activity of S. littoralis Scinax littoralis were found calling in every forest type sampled and bred in both tem- porary and permanent ponds. Males were found calling perched on shrubs ranging from 23-220 cm above the ground (69.73 ± 33.35 cm, n = 49) located 0-308 cm horizontally from pond edges (88.37 ± 95.83 cm, n = 49). The height of the perch used by S. littoralis was the same in all forest stages (F(2, 48) = 2.37, P = 0.106). However, perches used in the mature forest were farther from the water in comparison to the others (Fig. 3; F(2, 48) = 6.50, P = 0.003). Scinax littoralis calls during the night and may exhibits a satellite-male strategy (n = 5). Satellite behavior was observed at nights with density of calling males > 20 males per pond, in the late secondary forest and mature forest. Satellite males had a distance of approximately 19 ± 3 cm (n = 3) from the calling male. Satellite males were standing still near the calling male, not in low posture, however, we did not record any satellite male attempting to intercept gravid females. DISCUSSION Multi-pulse structure, common to both S. littoralis and S. angrensis, is widespread within the genus Scinax (e.g., Haddad and Pombal, 1987; Pombal et al., 1995; present study). Other call features common in S. catharinae group are present in the calls of these two species, such as short note lengths and harmonic structure (Pombal et al., 1995). The presence of harmonics in species of the Scinax catharinae group may be questionable (see Vielliard, 1993). In our results it is possible to identify different frequency bands, however, only two of it would be integer multiples of the fundamental frequency, while the other seems as additional frequency modulation. Therefore, some of these frequency bands may be sidebands (Vielliard, 1993). Despite of its origin, multiple frequency band structure in the advertisement call is common among the congeneric species (e.g., Haddad and Pom- bal, 1987). Anyway, despite morphological and ecological similarities between S. littoralis and S. angrensis, the differences among the call features of those species are evident, espe- cially regarding the dominant frequency level and call rate. Few species in the S. catharinae group have higher dominant frequencies than Scinax angrensis, such as S. agilis (Nunes et al., 2007), and S. argyreornatus (Pombal et al., 1995). These species can be distinguished from S. angrensis by the absence of multiple frequen- cy bands. The advertisement call of S. angrensis has more notes per call than S. albicans (Pombal et al., 1995), S. centralis (Pombal and Bastos, 1996), and S. hiemalis (Haddad and Pombal, 1987), although S. angrensis has lower number of notes per call than S. argyre- ornatus (Pombal et al., 1995), S. canastrensis (Cardoso and Haddad, 1982), S. machadoi (Bokermann and Sazima, 1973), Scinax rizibilis (Pombal et al., 1995), and S. skaios (Pom- bal et al., 2010). Scinax angrensis has a higher minimum frequency than S. machadoi as well, S. machadoi has the shortest call duration documented in the group (Pombal et al., 1995). Scinax angrensis has a shorter call than do S. skaios (Pombal et al., 2010), S. catharinae, S. rizibilis, and S. argyreornatus (Pombal et al., 1995). Scinax angresis differs from S. ranki by harmonic structure of advertisement call. For some species in this group 305Advertisement call of Scinax (e.g., S. catharinae) no full description of advertisement calls exists, thus we are unable to compare their calls to others in the S. catharinae group (Table 1). Some species in the S. catharinae group have higher dominant frequencies than Sci- nax littoralis, such as S. agilis (Nunes et al., 2007), S. albicans (Pombal et al., 1995), and S. argyreornatus (Pombal et al., 1995). Most species have a similar range of dominant fre- quency (Table 1). Scinax littoralis can be distinguished from S. argyreornatus, S. rizibilis (Pombal et al., 1995), and S. skaios (Pombal et al., 2010) by the presence of multiple fre- quency bands, lower number of notes per call, and call length. Scinax littoralis has higher call duration than S. centralis (Pombal and Bastos, 1996), S. catharinae, and S. machadoi (Pombal et al., 1995), although S. littoralis has lower call duration than S. albicans (Pom- bal et al., 1995). Advertisement call of S. ranki has no multiple frequency bands (Andrade and Cardoso, 1987). Scinax hiemalis has higher maximum frequency than S. littoralis (Haddad and Pombal, 1987). Scinax littoralis also has fewer notes per call than S. canast- rensis (Cardoso and Haddad, 1982), and S. machadoi (Bokermann and Sazima, 1973). As already know for the genus (e.g., Toledo and Haddad, 2005b; Abrunhosa et al., 2006), Scinax littoralis either exhibits a prolonged breeding season (sensu Wells, 1977a). The prolonged breeding season is common among tropical amphibians (Wells, 2007), and, despite our study site is at a subtropical zone, it is still likely the pattern already observed for S. littoralis in the type locality (Estação Ecológica Juréia-Itatins, Narvaes et al., 2009). Males of this species may adopt satellite strategy. Satellite males are common with- in the most Neotropical amphibian families (e.g., Haddad, 1991; Barreto and Andrade, 1995; Pombal and Haddad, 2005). Four hypotheses, not mutually exclusive, explain satel- lite behavior in amphibians: 1) satellite males are sexual parasites that try to mate with females attracted by their calling neighbor (Wells, 1977b); 2) satellite males are waiting for a change in the physical condition of the dominant male, due to the high energetic cost of calling (Bevier, 1997; Pombal and Haddad, 2005); 3) satellite males are inferior or subor- dinate males waiting for a calling site to be vacated by the dominant male when he enters amplexus (Wells, 1977b); 4) the satellite strategy can decrease predation risk by acousti- cally oriented predators (e.g., Tuttle et al., 1981). We observed satellite behavior only in high densities, as suggested by Gerhardt and Klump (1988). Since we did not observe any female interception attempt by satellite males (hypothesis 1), we suggest that the other three hypotheses are therefore more likely to happen in this species. Although S. littoralis appears to be constrained to the forest habitats, breeding activity seems to be independent of the successional stage of the forest and of pond stability. Perch heights did not vary between areas, but males vocalized farther from the margin of the water in mature forest. These differences in calling site may be related to the availability of bromeliads around the margin of the spawning site (see Toledo et al., 2012): and bro- meliads were most abundant in mature forest, including around the margins of breeding sites. We found low abundance of bromeliads in late secondary forest, and none in early secondary forest. We present here data on the natural history of two endemic species of Atlantic Forest of Brazil, which is an endangered biome with lacks on the life history knowledge of most amphibian species. The calling comparisons presented here aid in easy distinction among some species of the S. catharinae group in further surveys or to study the ecology and the evolution of the of advertisement call for the reproduction of Scinax clade. 306 Michel V. Garey et al. ACKNOWLEDGEMENTS We are indebted to Kristine Kaiser for helpful suggestions on the manuscript and for review- ing the English. M.V. Garey thanks the Fundação Grupo O Boticário de Proteção à Natureza (agree- ment FBPN-UFPR 623) for financial support and access to the study area. We thank IBAMA for assistance and for our collection permit (No. 107/06-RAN), and Programa de Pós Graduação em Ecologia e Conservação of the Universidade Federal do Paraná for general support. M.T. Hartmann thanks CAPES for financial support through the Programa de Apoio a Projetos Institucionais com a Participação de Recém-Doutores (PRODOC). During the preparation of this paper M.V. 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Toledo, L.F., Haddad, C.F.B. (2005a): Acoustic repertoire and calling behavior of Scinax fuscomarginatus (Anura, Hylidae). J. Herpetol. 39: 455-464. Toledo, L.F., Haddad, C.F.B. (2005b): Reproductive biology of Scinax fuscomarginatus (Anura, Hylidae) in south-eastern Brazil. J. Nat. Hist. 39: 3029-3037. Toledo, L.F., Garcia, P.C.A., Lingnau, R., Haddad, C.F.B. (2007): A new species of Sphaeno- rhynchus (Anura: Hylidae) from Brazil. Zootaxa 1658: 57-68. Tuttle, M.D., Taft, K.L., Ryan, M.J. (1981): Acoustical location of calling frogs by Philander opossum. Biotropica 13: 233-234. Vasconcelos, T.S., Rossa-Feres, D.C. (2005): Diversidade, distribuição espacial e temporal de anfíbios anuros (Amphibia, Anura) na região noroeste do estado de São Paulo, Brasil. Biota Neotrop. 5: 137-150. Vielliard, J. (1993): ‘Side-bands’ artifact and digital sound processing. Bioacoustics 5: 159- 162. Wells, K.D. (1977a): The social behaviour of anuran amphibians. Anim. Behav. 25: 666- 693. Wells, K.D. (1977b): Territoriality and male mating success in the green frog (Rana clami- tans). Ecology 58: 750-762. Wells, K.D. (2007): The ecology and behavior of amphibians. University of Chicago Press, Chicago. Zimmerman, B.L. (1993): Audio strip transects. In: Measuring and Monitoring Biologi- cal Diversity. Standard Methods for Amphibians, pp. 92-97. Heyer, W.R., Donnelly, M.A., McDiarmid, R.W., Hayek, L.A.C., Foster, M.S., Eds, Smithsonian Institution, Washington. Acta Herpetologica Vol. 7, n. 2 - December 2012 Firenze University Press Advertisement call of species of the genus Frostius Cannatella 1986 (Anura: Bufonidae) Flora A. Juncá1, David L. Röhr2, Ricardo Lourenço-de-Moraes3, Flávio J. M. Santos1, Airan S. Protázio1, Ednei A. Mercês1, Mirco Solé4 Amphibians in Southern Apennine: distribution, ecology and conservation notes in the “Appennino Lucano, Val d’Agri e Lagonegrese” National Park (Southern Italy). Antonio Romano1,*, Remo Bartolomei1, Antonio Luca Conte1, Egidio Fulco2 The significance of using satellite imagery data only in Ecological Niche Modelling of Iberian herps Neftalí Sillero1, José C. Brito2, Santiago Martín-Alfageme3, Eduardo García-Meléndez4, A.G. Toxopeus5, Andrew Skidmore5 Reproductive strategy of male and female eastern spiny lizards Sceloporus spinosus (Squamata: Phrynosomatidae) from a region of the Chihuahuan Desert, México Aurelio Ramírez-Bautista1,*, Barry P. Stephenson2, Xóchitl Hernández-Ibarra1, Uriel Hernández-Salinas1, Raciel Cruz-Elizalde1, Abraham Lozano1, and Geoffrey R. Smith3 Morphological variability of the Hermann’s tortoise (Testudo hermanni) in the Central Balkans Katarina Ljubisavljević1, Georg Džukić1, Tanja D. Vukov1, Miloš L. Kalezić1,2 The usefulness of mesocosms for ecotoxicity testing with lacertid lizards Maria José Amaral1,2,*, Rita C. Bicho1, Miguel A. Carretero2, Juan C. Sanchez-Hernandez3, Augusto M. R. Faustino4, Amadeu M. V. M. 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Perry1, Robson Waldemar Ávila3, Reinaldo José da Silva4 Rediscovery and redescription of the holotype of Lygosoma vittigerum (= Lipinia vittigera) Boulenger, 1894 Yannick Bucklitsch1, Peter Geissler1, Timo Hartmann1, Giuliano Doria2 , André Koch1,* Reproductive phenology of the tomato frog, Dyscophus antongili, in an urban pond of Madagascar’s east coast Ori Segev1,*, Franco Andreone2, Roberta Pala2, Giulia Tessa2, Miguel Vences1 Range extension of the critically endangered true poison-dart frog, Phyllobates terribilis (Anura: Dendrobatidae), in western Colombia Roberto Márquez1,*, Germán Corredor2, Carlos Galvis3 Daniel Góez2, & Adolfo Amézquita1 Differences in habitat use of two sympatric species of Ameiva in East Costa Rica Esther Sebastián-González1, Ramón Gómez2 ACTA HERPETOLOGICA Journal of the Societas Herpetologica Italica