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Zootaxa 1861: 44–54 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition)

A new species of Fitzinger (Anura, ) from Serra do Brigadeiro, State of Minas Gerais, Southeastern

ULISSES CARAMASCHI1,3, RENATO N. FEIO2 & VINÍCIUS A. SÃO-PEDRO2 1Departamento de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista, 20940-040 Rio de Janeiro, RJ, Brasil. E-mail: [email protected] 2Departamento de Biologia , Universidade Federal de Viçosa, 36571-000 Viçosa, MG, Brasil. E-mail: [email protected] 3Corresponding author. E-mail: [email protected]

Abstract

A new species of Leptodactylus belonging to the L. fuscus species group, and related to the L. mystaceus complex, is described from the Lagoa das Bromélias (20o53’S, 42o31’W; 1,227 m above sea level), Parque Estadual da Serra do Bri- gadeiro, Municipality of Ervália, State of Minas Gerais, Southeastern Brazil. Leptodactylus cupreus sp. nov. is character- ized by the large size for the group (SVL 50.1–55.1 mm in males) and color pattern. The new species has a non-pulsed advertisement call, with call rate about 12 calls/s and a dominant frequency between 2,800 and 3,058 Hz.

Key words: Amphibia, Leptodactylus cupreus sp. nov., advertisement call,

Introduction

A recent molecular phylogeny of (Frost et al. 2006) suggested that the genus Leptodactylus Fitz- inger includes the former genera Adenomera Steindachner, Lithodytes Fitzinger, and Vanzolinius Heyer (pre- viously synonymized with Leptodactylus by De Sá et al., 2005), consisting of a total of 83 species (81 species referred in Frost 2007 plus two new species recently described by Giaretta & Costa 2007 and Berneck et al. 2008). These species have been traditionally clustered in morphological groups (e.g., Heyer 1970, L. mel- anonotus group; Heyer 1973, L. marmoratus group; Heyer 1978, L. fuscus group; Heyer 1979, 2005, L. penta- dactylus group; Heyer 1994, L. podicipinus-L. wagneri group), without a phylogenetic framework. The Leptodactylus fuscus species group is the largest one in terms of the number of species. Members of the L. fuscus group are small to moderate sized ; the toes lack fringe or web, the head is of normal width proportions, and the males lack thumb spines (Heyer 1978). Within this group, an assemblage of morphologi- cally similar species, defined by having two distinct dorsolateral folds (at least indicated by color pattern), a distinct light lip stripe, a distinct longitudinal light stripe on the posterior surface of the thighs, posterior sur- face of tarsus smooth (or with distinct, non-obvious light tubercles), and sole of foot with prominent light tubercles has been identified as the “L. mystaceus complex” (Heyer et al. 1996); this assemblage includes L. mystaceus (Spix), L. elenae Heyer, L. notoaktites Heyer, L. spixi Heyer, and L. didymus Heyer, García-Lopez & Cardoso. Herein, a new species clearly related to the L. mystaceus morphological assemblage is described from the State of Minas Gerais, Brazil.

44 Accepted by M. Vences: 4 Aug. 2008; published: 1 Sep. 2008 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Material and methods

Examined specimens are housed at the Museu Nacional, Rio de Janeiro, Brazil (MNRJ) and Museu de História Natural João Moojen de Oliveira, Viçosa, Brasil (MZUFV). Comparative specimens are referred in the Appendix 1. Measurements were taken with digital calipers. Abbreviations of the measurements (in mm) are: SVL (snout-vent length); HL (head length); HW (head width); IND (internarial distance); END (eye to nostril dis- tance); ED (eye diameter); UEW (upper eyelid width); IOD (interorbital distance); TD (tympanum diameter); HAL (hand length); THL (thigh length); TL (tibia length); FL (foot length). Morphological terminology fol- lows Heyer et al. (1990). Specimens were recorded with a Panasonic RQ-L11 tape recorder on 08 October 2006, at 20 oC air temper- ature, by R. N. Feio. Calls were digitized from the same tape recorder to avoid velocity alterations and quality loss; and the digitized file did not present differences to the actual call when compared to hearing. Calls were digitized at a sampling rate of 22,050 Hz and resolution of 16 bits. Sound analyses were performed with the program Avisoft-SASLab Light for Windows, version 3.74 (www.avisoft.com). A sonogram was produced utilizing the following parameters: Bandwidth = 323 Hz; FFT length = 256; Frame = 100; Window = Flat Top; Overlap = 75; Time resolution = 2.90 ms; Contrast = char3.gray. Oscillograms and power spectrum of the calls were obtained with the program Sound Ruler Acoustic Analysis, version 0.9.6.0 (http://soun- druler.sf.net). Terminology used for call description follows Duellman and Trueb (1994).

Results

Leptodactylus cupreus sp. nov. (Figures 1–3)

Holotype: MNRJ 47752. Adult male (Figure 1). Lagoa das Bromélias (20o25’S, 43o29’W, 1,227 m above sea level), Parque Estadual da Serra do Brigadeiro, District of Careço, Municipality of Ervália, State of Minas Gerais, Southeastern Brazil. Collected by R.N. Feio and V.A. São-Pedro, on 08 October 2006.

FIGURE 1.Leptodactylus cupreus sp. nov., holotype (MNRJ 47752, SVL 51.8 mm), dorsal and ventral views.

NEW SPECIES OF LEPTODACTYLUS FROM BRAZIL Zootaxa 1861 © 2008 Magnolia Press · 45 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 2. Leptodactylus cupreus sp. nov., holotype (MNRJ 47752). A: Dorsal view of head. B: Lateral view of head. C: Hand. D: Foot. Bars equal 5 mm.

Paratypes: Seven adult males: MNRJ 47753–47754, collected with the holotype; MNRJ 50436–50438, MZUFV 8015–8016, collected at the type locality by R.N. Feio, V.A. São-Pedro, and P.S. Silva, on 24 Octo- ber 2007. Diagnosis: A species belonging to the L. fuscus group and related to the L. mystaceus complex by having two distinct dorsolateral folds, a distinct light lip stripe, a distinct longitudinal light stripe on the posterior sur- face of the thighs, posterior surface of tarsus smooth or with few tubercles, and sole of foot with prominent

46 · Zootaxa 1861 © 2008 Magnolia Press CARAMASCHI ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. light tubercles. The new species is characterized by: (1) size large for the group (SVL 50.1–55.1 mm in males); (2) head slightly wider than long, HL 96% of HW, HL 34% of SVL, HW 35% of SVL; (3) general color copper on dorsal surfaces of body and limbs; (4) lateral head black stripe defined, broad, extending from the tip of the snout and passing over the eye and tympanum; (5) lateral head with white stripe defined, delim- ited above and below by black stripes, extending from tip of snout to insertion of the forelimb; (6) a large bucal white gland found posterior to the jaw articulation and to the tympanum; (7) dorsum without spots, dor- sal surface of thighs and tibiae not distinctly barred; (8) dorsolateral folds broad, from the anterior third of the body to groin, light in color and defined by black markings below and above (only in its posterior part); (9) urostyle stripe present, anal region with many large white tubercles; (10) advertisement call not pulsed, call rate about 12 calls/s, dominant frequency between 2,800 and 3,058 Hz.

FIGURE 3. Leptodactylus cupreus sp. nov., color in life.

Comparisons with other species: Leptodactylus cupreus sp. nov. is distinguished from L. didymus, L. elenae, L. mystaceus, L. notoaktites, and L. spixi by the larger size (SVL 50.1–55.1 mm in males of L. cupreus sp. nov.; combined SVL 42.7–50.8 mm in males of the other species, see Heyer 1978 and Heyer et al. 1996); head slightly wider than long (HL 34% of SVL, HW 34.2% of SVL in L. cupreus sp. nov.; HL 36.8–39.8% of SVL, HW 33.6–35.2% of SVL in the other species, see Heyer 1978 and Heyer et al. 1996); general color cop- per on dorsal surfaces of body and limbs (general color brown to greyish brown in the other species); lateral head has a black, clearly defined, broad stripe, passing on the eye and tympanum (lateral head stripe less defined, thinner, grey to dark grey, passing under the eye and over the tympanum in the other species); lateral head with a white, defined stripe, delimited above and below by black stripes (poorly defined and poorly delimited in the other species); presence of a large, white gland behind the jaw articulation (absent or small, poorly developed, in the other species); dorsum without spots, dorsal surface of thighs and tibiae not distinctly barred (dorsum spotted, dorsal surface of thighs and tibiae distinctly barred in the other species); dorsolateral

NEW SPECIES OF LEPTODACTYLUS FROM BRAZIL Zootaxa 1861 © 2008 Magnolia Press · 47 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. folds broad, from the anterior third of the body to groin (dorsolateral folds thin, from the eye to groin in the other species); urostyle stripe present, anal region with many large white tubercles (urostyle stripe absent or indistinct, anal region without or with indistinct tubercles in the other species). Moreover, L. cupreus sp. nov. is distinguished from: (1) the all other species in the complex by an outer metacarpal tubercle divided and smaller than the inner metacarpal tubercle (outer tubercle entire, rounded or somewhat triangular, larger than the inner tubercle in the other species); (2) from L. didymus, L. elenae, L. mystaceus, and L. notoaktites by a smooth dorsal surface of the tibiae (with many small horny spines in these species); and (3) from L. didymus, L. mystaceus, and L. notoaktites by a tarsus and foot with few tubercles (sole of foot and tarsus with many tubercles in these species). Additionally, L. cupreus sp. nov. is distinguished from L. mystacinus (a species not included in the L. mystaceus complex, but with two dorsolateral folds and sometimes with a copper color pat- tern) by the wider head and longer legs, lateral head black broad stripe passing on the eye and tympanum (passing under the eye and not covering the tympanum in L. mystacinus), the thin white stripe on lip (wide in L. mystacinus), the presence of clear dorsolateral stripes (absent in L. mystacinus), dark bars on dorsal surface of thighs, tibiae, and feet poorly defined, fragmented (defined, continuous in L. mystacinus), presence of a dis- tinct light stripe on the posterior surface of thighs (absent in L. mystacinus) and surface of tarsus and foot with tubercles (distinct white tubercles present only on the tarsus in L. mystacinus). Leptodactylus cupreus sp. nov. has the fastest call rate in the assemblage, with about 12 calls/s (combined call rate ranging from 1.0 to 2.3 calls/s in L. didymus, L. elenae, L. mystaceus, L. notoaktites, and L. spixi) and a higher dominant frequency (between 2,800 and 3,058 Hz in L. cupreus sp. nov., ranging from 470 to 2,033 Hz in other species); additionally, L. cupreus sp. nov. may be promptly separated from L. mystaceus by having unpulsed advertisement call (pulsed in L. mystaceus). Additionally, the new species is distinguished from L. mystacinus by the fast call rate (12 calls/s in L. cupreus sp. nov.; 5–6.5 calls/s in L. mystacinus) and a higher dominant frequency (between 2,800 and 3,058 Hz in L. cupreus sp. nov.; 2,200 to 2,500 Hz in L. mystacinus). Description of the holotype: Robust build; head slightly wider than long, HL 96% of HW, HL 34% of SVL, HW 35% of SVL. Snout sub-elliptical viewed from above (Fig. 2A), protruding with distinct shelf in profile (Fig. 2B); canthus rostralis indistinct, rounded; loreal region oblique, slightly concave. Nostrils closer to tip of snout than to eyes; internarial distance slightly larger than eye to nostril distance and smaller than eye diameter. Eye to nostril distance smaller than eye diameter and larger than upper eyelid width, interorbital dis- tance, and tympanum diameter. Upper eyelid width smaller than interorbital distance and tympanum diameter. Tympanum circular, annulus distinct; tympanum largely separated from eye, its diameter smaller than eye diameter, TD 84% of ED. Upper eyelid, head, and dorsal skin smooth; a supratympanic fold from the poste- rior corner of eye, arching downwards posteriorly to tympanum, delimiting the well developed shoulder blade and reaching the arm articulation; a large, longitudinally elongated, jaw articulation gland present; a pair of dorsolateral folds, from the anterior third of the body to groin; flanks barely rugose, a weak dermal fold and sparse tubercles present; ventral skin smooth, belly disk fold distinct; a granular seat patch under thighs; anal region with many rounded tubercles; dorsal surface of tibiae with many small and horny tubercles. Vocal sac poorly developed, subgular; a pair of distinct, developed, lateral vocal folds. Vocal slits present; vomerine teeth in two transverse, almost contacting medially, slightly arched series, located between and just posterior to the choanae. Tongue large, free, slightly notched behind. Hand (Fig. 2 C) with fingers slender, not webbed, tips rounded, not expanded; weak lateral ridges on fingers II and III, absent on the others; fingers lengths IV < II < III < I, first finger much longer than second; subarticular tubercles rounded, with proximal tubercles more developed than distal ones; few supranumerary tubercles present; outer metacarpal tubercle large, longitudi- nally divided, the most outer part about three times the inner part; inner metacarpal tubercle two times larger than outer one, elliptical; no spine or asperities on thumbs, no prepollex visible; no tubercles or crests on fore- arm. Legs robust, tibia length slightly larger than the thigh length; sum of tibia and thigh lengths about 97.5% of SVL. Foot large (Fig. 2 D), foot length larger than tibia and thigh lengths, 75.3% of SVL. Toes slender, not webbed, nor fringed; toes lengths I < II < V < III < IV; toe tips slightly pointed; subarticular tubercles large,

48 · Zootaxa 1861 © 2008 Magnolia Press CARAMASCHI ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. rounded; sole of foot with distinct, approximately aligned tubercles; outer metatarsal tubercle very small, elliptical, flat; inner metatarsal tubercle large, elliptical, slightly elevated; sole of tarsus with few tubercles; inner tarsal fold distinct, approximately equal to the length of the tarsus. Measurements of the holotype: SVL 51.8; HL 17.4; HW 18.1; IND 4.6; END 4.5; ED 5.0; UEW 3.8; IOD 4.4; TD 4.2; HAL 12.2; THL 24.0; TL 26.5; FL 39.0. Color: In life, overall dorsal coloration uniformly copper; two dorsolateral stripes clear copper, from the anterior third of the body to groin, bounded above and below by black stripes. A light copper stripe on the uro- style. A wide black stripe from the tip of snout, passing over the naris, subcanthal region, inferior three fourth of eye, tympanum, and posteriorly bending towards the shoulder, bordering the shoulder blade inferiorly; below and along this black stripe, a golden, well defined stripe is visible; below the golden stripe, the upper and lower lips are dark grey. Bucal post-commissural gland distinctly golden. Flanks, below the black dorso- lateral stripe, copper above and grey copper below, with scattered clear copper spots and flecks. Forelimbs copper above, with a black stripe on anterior and posterior sides of arms and black stains on forearms; fingers clear grey with small yellow flecks above. Legs copper above; the anterior dorsolateral side of thighs with black spots that sometimes fuse with each other; a clear cream longitudinal line runs along the lower limit of the posterior surface of thighs and delimits the granulose seat pad; dorsum of tibiae and outer surface of feet with scattered black stains, without forming defined pattern of bars; toes clear grey with scattered clear copper flecks. Venter and ventral surfaces of forelimbs and legs whitish grey with scattered clear cream, undefined stains; gular region clear pink. Anal region with distinct white tubercles. Eyes copper on superior one third and black below. In preservative, the copper color surfaces become clear brownish grey; the dorsolateral and urostyle stripes become cream, the lateral head stripe become clear cream, almost white, and the black stripes on lat- eral head, dorsolateral body, arms, and legs are maintained; bucal post-commissural gland becomes white; the longitudinal stripe on posterior side of thighs becomes white. Venter greyish white; gular region clear grey; tubercles on anal region white. Variation: Examined specimens are congruent among them respecting the morphological characters and color. Range, mean, and standard deviation of the measurements of eight males are in Table 1.

TABLE 1. Range, mean, and standard-deviation (SD) of the measurements (mm) of Leptodactylus cupreus sp. nov. (n, number of specimens).

Males (n = 8) Characters Range Mean SD SVL 50.1–55.1 52.4 3.96 HL 16.9–18.8 17.9 2.88 HW 17.4–18.5 18.0 2.89 IND 4.5–5.9 5.0 1.62 END 4.3–4.8 4.5 1.49 ED 4.6–5.3 5.1 1.62 UEW 3.0–3.9 3.6 1.29 IOD 4.1–5.4 4.6 1.53 TD 4.0–4.7 4.4 1.47 HAL 12.0–13.1 12.4 2.52 THL 23.7–25.2 24.2 3.19 TL 26.0–27.4 26.1 3.26 FL 38.6–42.6 40.3 3.70

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Advertisement call (Fig. 4): Calls emitted irregularly in a variably long and fast sequence of peeps; call rate about 12 calls/s; call not pulsed, modulated, composed of three harmonics at 3,005 Hz, 5,770 Hz, and 8,527 Hz; fundamental and dominant frequency between 2,800 and 3,058 Hz.

FIGURE 4. Advertisement call of Leptodactylus cupreus sp. nov. A: Oscillogram of 24 calls emitted in 2 seconds. B: Detailed oscillogram of one call. C: Power spectrum of one call. D: Sonogram of a sequence of ten calls.

Geographic distribution: The new species is known only from type locality, associated to the northern part of the Mantiqueira Mountain Range Complex, locally called Serra do Brigadeiro. and habits: The type locality of L. cupreus is essentially the same as recently described for Chi- asmocleis mantiqueira (Anura, Microhylidae; see Cruz et al. 2007). The Parque Estadual da Serra do Briga-

50 · Zootaxa 1861 © 2008 Magnolia Press CARAMASCHI ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. deiro, in the Atlantic Rain biome, is a conservation unit managed by the Instituto Estadual de Florestas of the State of Minas Gerais. The 13,000 ha of the Parque involve the highest portions of a set of mountains integrating the Mantiqueira Mountain Range Complex, with maximum of 1,985 m above sea level. The Lagoa das Bromélias (local name meaning Bromeliads’ Lake) is a temporary at 1,227 m altitude, that com- pletely dryes- up during the dry season (April to September), but with about 250 m2 of water surface in the wet season (October to March). This pond is found in a forest fragment with especially rich epiphitic flora mainly represented by the Bromeliaceae and Orchidaceae plants families. Males of L. cupreus were found calling at night in densities of up to 20 individuals, only during the early month of the wet season (October), when the dryed pond ground is totally covered by herbaceous vegetation. The males call under these plants, inside small burrows excavated in the soil. Females, egg clutches, and tad- poles are unknown. Etymology: The specific epithet, “cupreus”, is a Latin adjective referred to the copper general color pat- tern of the new species.

Discussion

The species of the complex are essentially allopatric taxa found in South America, east of the Andes. However, the boundaries of L. didymus and L. mystaceus are not precise given that these species are separated only on the basis of advertisement call patterns (Heyer et al. 1996); they occur in the western Amazon Basin in far western Brazil, southeastern , and extreme northern , in the Depart- ment Pando (Heyer et al. 1996; Köhler & Lötters 1999; Frost 2007). Although L. didymus and L. mystaceus have been considered as sibling species (Heyer et al. 1996), molecular work demonstrated that they do not show sister-taxa relationships (De Sá et al. 2005). Leptodactylus elenae is a wide-ranging species inhabiting semi-arid in the Chaco, cerrado, caatinga, and dry forest regions from the eastern slope of Bolivia to Mato Grosso, in Brazil, and southern to north-central (Heyer & Heyer 2002). Leptodacty- lus mystaceus is distributed throughout the greater Amazon Basin and cerrados bordering that basin, in Ecua- dor, , Peru, , Guianas, and northern Brazil to the southern limit in central Brazil and Paraguay (Heyer 1978; Frost 2007); Toledo et al. (2005) extended the distribution to southeastern Brazil, in the State of São Paulo, and considered that the record of L. mystaceus (as L. amazonicus) by Heyer (1978) for the western State of Minas Gerais is actually L. mystaceus rather than L. didymus; notwithstanding, the records by Heyer (1978) for northeastern Brazil, in the states of Pernambuco and Alagoas, are actually referred to L. spixi. Leptodactylus notoaktites occurs in southeastern and southern Brazil, in the states of São Paulo, Paraná, and Santa Catarina (Heyer 1978; our data). Finally, is distributed on the east coast of Brazil, associated with the Atlantic Rain Forest, from the State of Ceará to eastern Minas Gerais and Rio de Janeiro (Heyer 1978, as L. mystaceus and part of L. amazonicus; our data). Leptodactylus cupreus is known only from the type locality, in an Atlantic Rain Forest fragment in the eastern State of Minas Gerais, above 1,200 m in altitude. This locality is inside the general distribution of L. spixi, but this species was never collected nor heard in the region. Apparently, L. cupreus is a high-altitude endemic species occurring in the isolated of the Serra do Brigadeiro, a distributional pattern previously reported for Physalaemus maxi- mus (Leiuperidae) and Chiasmocleis mantiqueira (see Feio et al. 1999, and Cruz et al. 2007). Advertisement calls for all species of “L. mystaceus complex” have been described or figured: L. didymus by Heyer et al. (1996) and Köhler and Lötters (1999); L. elenae by Barrio (1965, as L. mystaceus, reiterated by Straughan and Heyer 1976), Gallardo (1987), Gallardo and Valera de Olmedo (1992), Straneck et al. (1993), Marquez et al. (1995), and Heyer et al. (1996); L. mystaceus by Heyer (1978, as L. amazonicus), Heyer et al. (1996), and Toledo et al. (2005); L. notoaktites by Heyer et al. (1996); and L. spixi by Bilate et al. (2006); moreover, the call of L. mystacinus was treated on by Barrio (1965). Leptodactylus cupreus share the

NEW SPECIES OF LEPTODACTYLUS FROM BRAZIL Zootaxa 1861 © 2008 Magnolia Press · 51 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. harmonic and modulated structure of the call with all other species of the complex. The unpulsed call is also shared, except with L. mystaceus, the only species of the assemblage to present pulsed call. On the other hand, L. cupreus has highest call rate values and dominant frequency than any other species of the L. fuscus group (see Bilate et al. 2006, for a resume).

Acknowledgements

We acknowledge Dr. Carlos Alberto G. Cruz for comments on the manuscript, and Paulo Roberto Nascimento for the line drawings. The Instituto Estadual de Florestas de Minas Gerais for permission to collect in the Parque Estadual da Serra do Brigadeiro. The Conselho Nacional de Desenvolvimento Científico e Tec- nológico (CNPq) for the fellowship and grants to UC. Collect permits numbers IBAMA 253/04 and 289/06 - NUFAS/MG, and IEF 055/2005.

References

Barrio, A. (1965) Afinidades del canto nupcial de las especies cavicolas del genero Leptodactylus (Anura–Leptodactyl- idae). Physis, 25(70), 401–410. Berneck, B.M., Costa, C.O.R. & Garcia, P.C.A. (2008) A new speciesof Leptodactylus (Anura: Leptodactylidae) from the Atlantic Forest of São Paulo State, Brazil. Zootaxa, 1795, 46–56. Bilate, M., Wogel, H., Weber, L.N. & Abrunhosa, P.A. (2006) Vocalizações e girino de Leptodactylus spixi Heyer, 1983 (Amphibia, Anura, Leptodactylidae). Arquivos do Museu Nacional, Rio de Janeiro, 64(3), 235–245. Cruz, C.A.G., Feio, R.N. & Cassini, C.S. (2007) Nova espécie de Chiasmocleis Méhelÿ, 1904 (Amphibia, Anura, Micro- hylidae) da Serra da Mantiqueira, Estado de Minas Gerais, Brasil. Arquivos do Museu Nacional, Rio de Janeiro, 65(1), 33–38. De Sá, R.O., Heyer, W.R. & Camargo, A. (2005) A phylogenetic analysis of Vanzolinius Heyer, 1974 (Amphibia, Anura, Leptodactylidae): taxonomic and life history implications. Arquivos do Museu Nacional, Rio de Janeiro, 63(1),707– 726. De Sá, R.O., Camargo, A. & Heyer, W.R. (2005) Are Leptodactylus dydimus and L. mystaceus phylogenetically sibling species (Amphibia, Anura, Leptodactylidae)? Pp. 90–92. In: Ananajeva, A. & Tsinenko, O. (Eds.) Herpetologia Pet- ropolitana. Proceeding of the 12th Ordinary General Meeting of the Societas Europaea Herpetologica, St. Petersburg, Russia. Duellman, W.E. & Trueb, L. (1994) Biology of Amphibians. The Johns Hopkins University Press, Baltimore. xxiv+670 pp. Feio, R.N., Pombal, Jr., J.P. & Caramaschi, U. (1999) A new Physalaemus (Anura, Leptodactylidae) from the Atlantic Rain Forest of Minas Gerais, Brazil. Copeia, 1999(1), 141–145. Frost, D.R. (2007) Species of the World: An Online Reference. Version 5.1 (10 October, 2007). Electronic database accessible at http://research.amnh.org/herpetology/amphibia/index.php. American Museum of Natural His- tory, New York, USA (Accessed 30 June 2008). Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., De Sá, R.O., Channing, A., Wilkinson, M., Donellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370. Gallardo, J.M. (1987) Anfibios argentinos: Guía para su identificación. Biblioteca Mosaico, Buenos Aires. Gallardo, J.M. & Valera de Olmedo, E. (1992) Anfibios de la Republica Argentina: ecología y comportamiento. Fauna de Agua Dulce de la Republica Argentina, 41, 1–116. Giaretta, A.A. & Costa, H.M. (2007) A redescription of Leptodactylus jolyi Sazima and Bokermann (Anura, Leptodac- tylidae) and the recognition of a new closely related species. Zootaxa, 1608, 1–10. Heyer, W.R. (1970) Studies on the frogs of the genus Leptodactylus (Amphibia: Leptodactylidae). VI. Biosystematics of the Melanonotus group. Contributions in Science, Natural History Museum of Los Angeles County, 191, 1–48. Heyer, W.R. (1973) Systematics of the Marmoratus group of the genus Leptodactylus (Amphibia, Leptodactylidae). Contributions in Science, Natural History Museum of Los Angeles County, 251, 1–49. Heyer, W.R. (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Natu- ral History Museum of Los Angeles County, Science Bulletin, 29, 1–85.

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Heyer, W.R. (1979) Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia: Lepto- dactylidae). Smithsonian Contributions to Zoology, (301), 1–43. Heyer, W.R. (1983) Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978 and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae). Proceedings of the Biological Soci- ety of Washington, 96(2), 270–272. Heyer, W.R. (1994) Variation within the Leptodactylus podicipinus - wagneri complex of frogs (Amphibia: Leptodactyl- idae). Smithsonian Contributions to Zoology, (546), 1–124. Heyer, W.R. (2005) Variation and taxonomic clarification of the large species of the Leptodactylus pentadactylus species group (Amphibia: Leptodactylidae) from Middle America, Northern South America, and Amazonia. Arquivos de Zoologia, São Paulo, 37(3), 269–348. Heyer, W.R. & Heyer, M.M. (2002) Leptodactylus elenae Heyer. Catalogue of American Amphibians and Reptiles, 742, 1–5. Heyer, W.R., García-Lopez, J.M. & Cardoso, A.J. (1996) Advertisement call variation in the Leptodactylus mystaceus species complex (Amphibia: Leptodactylidae) with a description of a new sibling species. Amphibia-Reptilia, 17, 7– 31. Heyer, W.R., Rand, A.S., Cruz, C.A.G., Peixoto, O.L. & Nelson, C.E. (1990) Frogs of Boracéia. Arquivos de Zoologia, São Paulo, 31(4), 231–410. Köhler, J. & Lötters, S. (1999) Annotated list of amphibian records from the Departamento Pando, Bolivia, with descrip- tion of some advertisement calls. Bonner Zoologische Beiträge, 48(3–4), 259–273. Marquez, R., De La Riva, I. & Bosch, J. (1995) Advertisement calls of Bolivian Leptodactylidae (Amphibia, Anura). Journal of Zoology, London, 237, 313–336. Straneck, R., Olmedo, E.V. & Carrizo, G.R. (1993) Catalogo de Voces de Anfibios Argentinos. Ediciones L.O.L.A., Bue- nos Aires, 131 pp. Straughan, I.R. & Heyer, W.R. (1976) A functional analysis of the mating calls of the neotropical frog genera of the Lep- todactylus complex (Amphibia, Leptodactylidae). Papéis Avulsos de Zoologia, São Paulo, 29(23), 221–245. Toledo, L.F., Castanho, L.M. & Haddad, C.F.B. (2005) Recognition and distribution of Leptodactylus mystaceus (Anura; Leptodactylidae) in the State of São Paulo, Southeastern Brazil. Biota Neotropica, 5(1), 57–62.

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Appendix 1. Comparative specimens examined

Leptodactylus elenae—BRAZIL: Mato Grosso: Barão de Melgaço, RPPN SESC-Pantanal (MNRJ 43853–43858); Car- andasal (MNRJ 50888–50890). PARAGUAY: Independencia, near Villa Rica (MNRJ 50886–50887). Leptodactylus mystaceus – BRAZIL: Amazonas: Manaus (MNRJ 50880); Manaus, Reserva Ducke (MNRJ 7995); Rio Nhamundá (MNRJ 50905–50907). Pará: Belém (MNRJ 1470); Monte Alegre (MNRJ 50895); Óbidos, Rio Jamac- arú (MNRJ 50896); Serra do Cachimbo (MNRJ 50891); Serra do Macaco, Bacia do Rio Paru de Leste (MNRJ 50908–50910). Tocantins: Tocantins (MNRJ 50892–50893). Goiás: Aliança, Jatobazinho (MNRJ 2699, 12393– 12396); Rio São Miguel (MNRJ 0770). Mato Grosso: Estrada Cáceres-Porto Velho (MNRJ 1839, 10188). Minas Gerais: João Pinheiro (MNRJ 38820). Leptodactylus mystacinus—BRAZIL: Goiás: UHE Serra da Mesa (MNRJ 20167–20186). Distrito Federal: Brasília (MNR 2714, 12423–12424). Mato Grosso do Sul: Volta Grande (MNRJ 43647). Minas Gerais: Conceição do Ibiti- poca (MNRJ 33043); Cristália, Fazenda Curral Velho (MNRJ 40584); João Pinheiro, Fazenda Gameleira (MNRJ 45223, 45225, 38815); Lagoa Santa (MNRJ 1050); Santana do Riacho, Serra do Cipó (MNRJ 38738); Turmalina, Peixe Crú (MNRJ 34493–34496). Bahia: Guaratinga, Fazenda Vista Bela (MNRJ 25675); Itamaraju, Fazenda Novo Pau Brasil (MNRJ 46851). Espírito Santo: Sooretama, Reserva Biológica de Sooretama (MNRJ 35004). Rio de Jan- eiro: Arraial do Cabo, APA de Massambaba (MNRJ 45739); Cabo Frio, Restinga do Peró (MNRJ 47537); Niterói, Itaipú (MNRJ 51993); Saquarema, Palmital (MNRJ 25448–25449); Teresópolis (MNRJ 5411). São Paulo: Botu- catu, Fazenda Lageado (MNRJ 19309–19313); Rio Claro (MNRJ 1049). Rio Grande do Sul: Santa Maria (MNRJ 18762). Leptodactylus notoaktites—BRAZIL: São Paulo: Pedro de Toledo, 4 km (MNRJ 29061–29062); Pirassununga (MNRJ 2107); Ribeirão Branco, Fazenda São Luís (MNRJ 17653, 17692, 18396–18403); Ribeirão Branco (MNRJ 18247– 18248). Paraná: Cerro Azul (MZUFV 4810); Morretes, Estrada Itupava-Porto de Cima (MNRJ 402217). Santa Catarina: Corupá (MNRJ 0396, 0398, 5379–5401); Santa Luzia (MNRJ 2148, paratype; MNRJ 1624, 2690, 8447); São Bento do Sul (MNRJ 48296); Serra Nossa Senhora Catarina (MNRJ 50903). Leptodactylus spixi—BRAZIL: Ceará: Ererê (MNRJ 50894); Guaramiranga, Serra do Baturité (MNRJ 13586). Paraíba: Areias, Mata do Pau Ferro (MNRJ 18079–18081). Pernambuco: Igarassú (MNRJ 2363). Alagoas: São Miguel dos Campos, Fazenda do Prata (MNRJ 9584). Sergipe: Cristinópolis (MNRJ 32866). Bahia: Ilhéus, CEPLAC (MNRJ 33095); Ilhéus, Fazenda Almada (MNRJ 1724, 9300–9302); Ilhéus, Fazenda Pirataquicé (MNRJ 1706); Jussari, RPPN Serra do Teimoso (MNRJ 44966); Mascote, Fazenda São José (MNRJ 40467). Espírito Santo: Cachoeiro do Itapemirim (MNRJ 19367–19368); Linhares, Povoação (MNRJ 29054, 29064–29065, 29066–29067); Santa Teresa (MNRJ 1755, 9387). Rio de Janeiro: Duque de Caxias, Cidade das Meninas (MNRJ 1656, 2861, 8514–8516); Duque de Caxias (MNRJ 1809, 2374, 10126–10129); Mangaratiba (MNRJ 0568, 5708–5709); Niterói, Itaipú (MNRJ 43803); Petrópolis, Correias (MNRJ 0401, 5912); Porciúncula, Fazenda Vargem Alegre (MNRJ 43612); Rio de Janeiro, Parque Natural Municipal da Serra do Mendanha (MNRJ 39080); Rio de Janeiro, Realengo (MNRJ 2427, 11287–11289); Rio de Janeiro (MNRJ 0399, 46667); Santana (MNRJ 2086); Saquarema, Palmital (MNRJ 30309–30310); Teresópolis, Granja Comary (MNRJ 50899–50902); Teresópolis (MNRJ 0397, 5408–5410, 50897– 50898, 50904) . Minas Gerais: Chiador (MNRJ 37258–37259, 37260–37262); Juiz de Fora, Estação Agrícola de Água Limpa (MNRJ 50881–50882, 50883–50885); Viçosa (MZUFV 8047–8048).

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