zootaxa, a new species of leptodactylus fitzinger (anura ... · head slightly wider than long (hl...

Accepted by M. Vences: 4 Aug. 2008; published: 1 Sep. 2008 44

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2008 · Magnolia Press

Zootaxa 1861: 44–54 (2008) www.mapress.com/zootaxa/

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

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] de Biologia Animal, Universidade Federal de Viçosa, 36571-000 Viçosa, MG, Brasil. E-mail: [email protected] 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-pulsedadvertisement 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, taxonomy

Introduction

A recent molecular phylogeny of amphibians (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 speciesreferred 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 ofthe L. fuscus group are small to moderate sized frogs; the toes lack fringe or web, the head is of normal widthproportions, 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), adistinct 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 lighttubercles 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 describedfrom the State of Minas Gerais, Brazil.

TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Mario Ribeiro

Sticky Note

Unmarked set by Mario Ribeiro

Rectangle

Mario Ribeiro

Sticky Note

Zootaxa 1861 © 2008 Magnolia Press · 45NEW SPECIES OF LEPTODACTYLUS FROM BRAZIL

Material and methods

Examined specimens are housed at the Museu Nacional, Rio de Janeiro, Brazil (MNRJ) and Museu deHistória Natural João Moojen de Oliveira, Viçosa, Brasil (MZUFV). Comparative specimens are referred inthe 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 20oC air temper-ature, by R. N. Feio. Calls were digitized from the same tape recorder to avoid velocity alterations and qualityloss; and the digitized file did not present differences to the actual call when compared to hearing. Calls weredigitized at a sampling rate of 22,050 Hz and resolution of 16 bits. Sound analyses were performed with theprogram Avisoft-SASLab Light for Windows, version 3.74 (www.avisoft.com). A sonogram was producedutilizing 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 thecalls 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 sealevel), Parque Estadual da Serra do Brigadeiro, District of Careço, Municipality of Ervália, State of MinasGerais, 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.


Rectangle

CARAMASCHI ET AL.46 · Zootaxa 1861 © 2008 Magnolia Press

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 havingtwo 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


Rectangle


light tubercles. The new species is characterized by: (1) size large for the group (SVL 50.1–55.1 mm inmales); (2) head slightly wider than long, HL 96% of HW, HL 34% of SVL, HW 35% of SVL; (3) generalcolor copper on dorsal surfaces of body and limbs; (4) lateral head black stripe defined, broad, extending fromthe 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 largebucal 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 thebody 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, callrate 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. cupreussp. 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% ofSVL, 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); lateralhead has a black, clearly defined, broad stripe, passing on the eye and tympanum (lateral head stripe lessdefined, thinner, grey to dark grey, passing under the eye and over the tympanum in the other species); lateralhead with a white, defined stripe, delimited above and below by black stripes (poorly defined and poorlydelimited 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 distinctlybarred (dorsum spotted, dorsal surface of thighs and tibiae distinctly barred in the other species); dorsolateral


Rectangle


folds broad, from the anterior third of the body to groin (dorsolateral folds thin, from the eye to groin in theother species); urostyle stripe present, anal region with many large white tubercles (urostyle stripe absent orindistinct, 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 andsmaller than the inner metacarpal tubercle (outer tubercle entire, rounded or somewhat triangular, larger thanthe inner tubercle in the other species); (2) from L. didymus, L. elenae, L. mystaceus, and L. notoaktites by asmooth 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 manytubercles in these species). Additionally, L. cupreus sp. nov. is distinguished from L. mystacinus (a species notincluded 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 inL. mystacinus), the presence of clear dorsolateral stripes (absent in L. mystacinus), dark bars on dorsal surfaceof 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 withtubercles (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 (combinedcall rate ranging from 1.0 to 2.3 calls/s in L. didymus, L. elenae, L. mystaceus, L. notoaktites, and L. spixi) anda higher dominant frequency (between 2,800 and 3,058 Hz in L. cupreus sp. nov., ranging from 470 to 2,033Hz in other species); additionally, L. cupreus sp. nov. may be promptly separated from L. mystaceus by havingunpulsed 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 higherdominant 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% ofSVL, HW 35% of SVL. Snout sub-elliptical viewed from above (Fig. 2A), protruding with distinct shelf inprofile (Fig. 2B); canthus rostralis indistinct, rounded; loreal region oblique, slightly concave. Nostrils closerto tip of snout than to eyes; internarial distance slightly larger than eye to nostril distance and smaller than eyediameter. 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 eyediameter, 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 bladeand reaching the arm articulation; a large, longitudinally elongated, jaw articulation gland present; a pair ofdorsolateral folds, from the anterior third of the body to groin; flanks barely rugose, a weak dermal fold andsparse tubercles present; ventral skin smooth, belly disk fold distinct; a granular seat patch under thighs; analregion with many rounded tubercles; dorsal surface of tibiae with many small and horny tubercles. Vocal sacpoorly developed, subgular; a pair of distinct, developed, lateral vocal folds. Vocal slits present; vomerineteeth in two transverse, almost contacting medially, slightly arched series, located between and just posteriorto 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 moredeveloped 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 largerthan 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, notwebbed, nor fringed; toes lengths I < II < V < III < IV; toe tips slightly pointed; subarticular tubercles large,


Rectangle


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 theanterior 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 fourthof 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 upperand 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. Forelimbscopper above, with a black stripe on anterior and posterior sides of arms and black stains on forearms; fingersclear grey with small yellow flecks above. Legs copper above; the anterior dorsolateral side of thighs withblack spots that sometimes fuse with each other; a clear cream longitudinal line runs along the lower limit ofthe posterior surface of thighs and delimits the granulose seat pad; dorsum of tibiae and outer surface of feetwith scattered black stains, without forming defined pattern of bars; toes clear grey with scattered clear copperflecks. Venter and ventral surfaces of forelimbs and legs whitish grey with scattered clear cream, undefinedstains; gular region clear pink. Anal region with distinct white tubercles. Eyes copper on superior one thirdand black below.

In preservative, the copper color surfaces become clear brownish grey; the dorsolateral and urostylestripes 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; thelongitudinal 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 andcolor. 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


Rectangle


Advertisement call (Fig. 4): Calls emitted irregularly in a variably long and fast sequence of peeps; callrate about 12 calls/s; call not pulsed, modulated, composed of three harmonics at 3,005 Hz, 5,770 Hz, and8,527 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 northernpart of the Mantiqueira Mountain Range Complex, locally called Serra do Brigadeiro.

Habitat 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-


Rectangle


deiro, in the Atlantic Rain Forest biome, is a conservation unit managed by the Instituto Estadual de Florestasof the State of Minas Gerais. The 13,000 ha of the Parque involve the highest portions of a set of mountainsintegrating the Mantiqueira Mountain Range Complex, with maximum of 1,985 m above sea level. The Lagoadas Bromélias (local name meaning Bromeliads’ Lake) is a temporary pond 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 wetseason (October to March). This pond is found in a forest fragment with especially rich epiphitic flora mainlyrepresented 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 earlymonth 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 Leptodactylus mystaceus 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 thesespecies are separated only on the basis of advertisement call patterns (Heyer et al. 1996); they occur in thewestern Amazon Basin in far western Brazil, southeastern Peru, and extreme northern Bolivia, in the Depart-ment Pando (Heyer et al. 1996; Köhler & Lötters 1999; Frost 2007). Although L. didymus and L. mystaceushave been considered as sibling species (Heyer et al. 1996), molecular work demonstrated that they do notshow sister-taxa relationships (De Sá et al. 2005). Leptodactylus elenae is a wide-ranging species inhabitingsemi-arid habitats in the Chaco, cerrado, caatinga, and dry forest regions from the eastern slope of Bolivia toMato Grosso, in Brazil, and southern Paraguay to north-central Argentina (Heyer & Heyer 2002). Leptodacty-lus mystaceus is distributed throughout the greater Amazon Basin and cerrados bordering that basin, in Ecua-dor, Colombia, Peru, Venezuela, Guianas, and northern Brazil to the southern limit in central Brazil andParaguay (Heyer 1978; Frost 2007); Toledo et al. (2005) extended the distribution to southeastern Brazil, inthe State of São Paulo, and considered that the record of L. mystaceus (as L. amazonicus) by Heyer (1978) forthe western State of Minas Gerais is actually L. mystaceus rather than L. didymus; notwithstanding, therecords by Heyer (1978) for northeastern Brazil, in the states of Pernambuco and Alagoas, are actuallyreferred to L. spixi. Leptodactylus notoaktites occurs in southeastern and southern Brazil, in the states of SãoPaulo, Paraná, and Santa Catarina (Heyer 1978; our data). Finally, Leptodactylus spixi is distributed on theeast coast of Brazil, associated with the Atlantic Rain Forest, from the State of Ceará to eastern Minas Geraisand Rio de Janeiro (Heyer 1978, as L. mystaceus and part of L. amazonicus; our data). Leptodactylus cupreusis 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 nevercollected nor heard in the region. Apparently, L. cupreus is a high-altitude endemic species occurring in theisolated forests 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. didymusby Heyer et al. (1996) and Köhler and Lötters (1999); L. elenae by Barrio (1965, as L. mystaceus, reiteratedby 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


Rectangle


harmonic and modulated structure of the call with all other species of the complex. The unpulsed call is alsoshared, 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 Nascimentofor the line drawings. The Instituto Estadual de Florestas de Minas Gerais for permission to collect in theParque 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) fromthe 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 siblingspecies (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+670pp.

Feio, R.N., Pombal, Jr., J.P. & Caramaschi, U. (1999) A new Physalaemus (Anura, Leptodactylidae) from the AtlanticRain Forest of Minas Gerais, Brazil. Copeia, 1999(1), 141–145.

Frost, D.R. (2007) Amphibian Species of the World: An Online Reference. Version 5.1 (10 October, 2007). Electronicdatabase 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 NaturalHistory, 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 frog 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.


Rectangle


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 adescription 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 speciesgroup (Amphibia: Leptodactylidae) from Middle America, Northern South America, and Amazonia. Arquivos deZoologia, 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 mystaceusspecies 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.


Rectangle


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); RioNhamundá (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 (MNRJ50908–50910). 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). MinasGerais: 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 (MNRJ45223, 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 NovoPau 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 (MNRJ18762).

Leptodactylus notoaktites—BRAZIL: São Paulo: Pedro de Toledo, 4 km (MNRJ 29061–29062); Pirassununga (MNRJ2107); 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). SantaCatarina: 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 dosCampos, Fazenda do Prata (MNRJ 9584). Sergipe: Cristinópolis (MNRJ 32866). Bahia: Ilhéus, CEPLAC (MNRJ33095); 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 doItapemirim (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); Riode Janeiro, Parque Natural Municipal da Serra do Mendanha (MNRJ 39080); Rio de Janeiro, Realengo (MNRJ2427, 11287–11289); Rio de Janeiro (MNRJ 0399, 46667); Santana (MNRJ 2086); Saquarema, Palmital (MNRJ30309–30310); 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 deÁgua Limpa (MNRJ 50881–50882, 50883–50885); Viçosa (MZUFV 8047–8048).


Rectangle

zootaxa, a new species of leptodactylus fitzinger (anura ... · head slightly wider than long (hl...

Documents