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Distribution extension and new breeding site of Leptodactylus barrioi (Anura: Leptodactylidae)

Scinax cosenzai sp. nov. (Figs 1, 2, 3) Holotype. MNRJ 75462, adult male, collected at Lage das Brom��lias, Serra do Brigadeiro (20 �� 43 ��� 13.5 ���� S and 42 �� 28 ��� 48.7 ���� W; 1385 m above sea level), Municipality of Araponga, state of Minas Gerais, Brazil. Collected by J.V.A. Lacerda, B. Assis and J.S. Dayrell on October 28, 2007. Paratopotypes. MZUFV 3895, adult male collected by Feio, R.N. and Santos-Filho, J.B, on 0 3 September 1999; MZUFV 6653, adult male collected by R.N. Feio, E.F. Oliveira and J.S. Dayrell, on 0 5 December 2005; MZUFV 8275, adult male collected by J.V.A. Lacerda and B. Assis, on 29 January 2008; MNRJ 75463 ��� 64, adult males collected by Feio, R.N. and Santos-Filho, J.B, on 0 3 September 1999; MNRJ 75465, adult male collected by R.N. Feio, E.F. Oliveira, H.C. Costa and L.L. Moraes, on 22 September 2005; MNRJ 75466, adult male collected by C.S. Cassini and R.N. Feio, on 21 February 2006. Referred specimens. Scinax cosenzai sp. nov.: BRAZIL: State of Minas Gerais: Municipality of Araponga (topotypes): MZUFV 4956, 6654, 7682, 7779, 8029, 8092, 8093, 8567, 8568, 11458���11463, 11470��� 11473); Municipality of Erv��lia: Distrito do Care��o (20 �� 51 ��� 9.24 ������ S and 42 �� 31 ��� 5.56 ������ W; 980 m above sea level): MZUFV 11532 ���11537, 11541��� 11543. Diagnosis. Scinax cosenzai sp. nov. belongs to the S. perpusillus group based on the use of bromeliads for reproduction and reduced webbing between toes II and III. The new species is characterized by: (1) moderate-size within the group (males 17.54���20.97 mm SVL, females 22.65���24.02 mm SVL); (2) snout protruding in lateral view and slightly acuminated in dorsal view; (3) snout with a distinct medial process, defined by Silva and Alves- Silva (2008) as a protuberance caused by the cartilaginous medial prenasal process of the nasal tectum under the skin; (4) medium-sized, elliptical choanae; (5) vomerine teeth in two arc shape series between choanae; (6) head longer than wide; (7) protruding eye; (8) canthus rostralis distinct; (9) loreal region concave; (10) tympanum round and pigmented, with diameter less than half the diameter of the eye; (11) thigh plus tibia length shorter than SVL; (12) webbing lacking between toes I and II and vestigial between II and III; (13) few tubercles scattered throughout the dorsal surface, including the head; (14) ventral skin granulated, more intensively near the cloacal region; (15) dorsum different tones of gray with transversal dark bars originating on each inguinal region and converging on the third portion of the dorsum in an inverted V-shaped figure; (16) lateral dark stripe, from the region of the eye to the anterior margin of the inguinal region; (17) head with an interocular dark transversal bar; (18) transversal dark bars along the anterior and posterior members; (19) discreet nuptial glandular pads on the base of thumb; (20) gular region with irregular dark pigmentation; (21) in life, hidden portion of the thigh and tibia region with some irregular bright yellow colored patches (Fig. 3); (22) a distinct vocalization with heterogeneous multipulsed notes (2���14 notes/call, call duration 177.62���2066.75 ms, 1���33 pulses/note and dominant frequency of 3375.91���4571.21 Hz). Advertisement call. We analyzed a total of 45 calls recorded at the type locality belonging to three call series emitted by three individuals that were not collected. We were unable to observe direct interaction of these males with other males and females during the recordings. The duration of the call series was 34.4, 55.1 and 68.9 s, with 11, 16 and 18 calls/call series, call rate of 0.25, 0.29 and 0.32 calls/sec and interval between calls of 2.78 s (1.9���4.32; SD = 0.64; n= 42 intervals). Calls are constituted by 2���14 multipulsed notes with duration of 177.62���2066.75 ms and dominant frequency of 3375.91���4571.21 Hz (Table 1). Note duration of 36.15 ms (1.81���111.56; SD = 23.94; n= 236 notes), 10.53 pulses/note (1���33; SD = 7.08; n= 236 notes) and rate of 315.12 pulses/ second (58.60���571.42; SD = 103.08; n= 236 notes). Notes are heterogeneous in duration, number of pulses and amplitude. Figure 4 shows the sonogram of the call series with 11 calls emitted by a single male and sonograms and spectrograms of each of these calls. Comparison with other species. Adult individuals of Scinax cosenzai sp. nov. may be distinguished from other species of the Scinax perpusillus group by a combination of traits. The new species differs from S. alcatraz by its smaller size (SVL 19.7���24.4 mm in males and 27.0��� 29.8 mm in females of S. alcatraz; Brasileiro et al. 2007 a) and by lacking inguinal glands (easily visible in S. alcatraz; Faivovich et al. 2010). Scinax cosenzai sp. nov. can be distinguished from S. arduous by its smaller size [19.3���22.7 mm (x=21.0; SD = 0.8; n= 20) in males of S. arduous], slender body, less protruding eyes, more evident canthus rostralis, loreal region more concave, THL + TBL shorter than SVL (longer in S. arduous) and a distinct advertisement call. It differs from S. atratus by the larger size of the female (maximum SVL of females of S. atratus 20.0 mm; Peixoto 1988 b) and by its dorsal color pattern typical of the S. perpusillus group (melanic in S. atratus; Peixoto 1988 b). The new species is promptly distinguished from S. belloni by not possessing dorsum of body and limbs densely covered by prominent granules, having trace of markings on dorsum, inguinal region and hidden surfaces of hind limbs, lacking inguinal glands and by the smaller size (19.8 ���23.0 mm SVL in males and 26.3���29.2 in females of S. belloni; Faivovich et al. 2010). It differs from S. faivovichi by its larger SVL (SVL 16.2 ���18.0 mm in 23 males of S. faivovichi and 18.6���21.7 mm in six females; Brasileiro et al. 2007 b), furthermore S. faivovichi has brown undersurface of tarsus and foot (not brown in the other species of the S. perpusillus group; Brasileiro et al. 2007 b). Scinax cosenzai sp. nov. is promptly distinguishable from S. insperatus by the presence of a yellow flash color on inguinal region and hidden surfaces of thigh and tibia (absence of yellow markings in the inguinal region and hidden surfaces of limbs in S. insperatus; Silva & Alves-Silva 2011) and by its smaller SVL (18.5���21.9 mm in males of S. insperatus; Silva & Alves-Silva 2011). The new species differs from S. littoreus by lacking inguinal glands (easily visible in S. littoreus; Faivovich et al. 2010), by the less pronounced eyes, dorsal color pattern, snout profile and width of the head (dorsum brownish colored, rounded snout profile and wider head in S. littoreus). It differs from S. melloi by its larger size (maximum SVL in males of S. melloi 17.00 mm and 18.7 mm in females; Peixoto 1988 b) and by its dorsal color pattern typical of the S. perpusillus group (marble dorsal pattern in S. melloi; Peixoto 1988 b). Scinax cosenzai sp. nov. can be distinguished from S. peixotoi by having tibia larger than thigh (tibia same size as thigh in S. peixotoi; Brasileiro et al. 2007 a), slender body and by the distinct advertisement call. It differs from S. perpusillus by its slender body, less pronounced eyes, snout more acuminated and a distinct advertisement call. The new species is distinguishable from S. tupinamba by the larger size [SVL of 17.8 mm in S. tupinamba (16.1���19.2; SD = 0.9; n= 21); Silva & Alves- Silva 2008], loreal region less concave and by the absence of a yellow stripe on the head of the tadpoles of the new species (the tadpole of S. tupinamba has a distinct, large, and bright yellow stripe between the eyes and the nostrils; Silva & Alves-Silva 2008). Scinax cosenzai sp. nov. differs from S. v-signatus by its smaller SVL [21.4���23.5 mm (x= 22.4; SD = 0.7; n= 20) in males of S. v-signatus], less rugose dorsal skin texture, canthus rostralis less evident, loreal region less concave and with less tubercles. Besides, S. v-signatus has a dark pigmentation on the gular region organized in the form of a Y or V (Lutz 1973; Peixoto 2002), and has a distinctive orange-flash color on the foot, lower leg, thigh, inguinal region, and armpit (H. R. da Silva, pers. com.). Scinax cosenzai sp. nov. has irregular dark pigmentation on the gular region and presents a yellow-flash color only on inguinal region and hidden surfaces of thigh and tibia, lacking on armpits and foot. The advertisement call of S. peixotoi (3���5 notes and duration of 146���232 ms; Brasileiro et al. 2007 a) is shorter than that of the new species (the calls of Scinax cosenzai sp. nov. with 3���5 notes have duration of 512.75���941.81 ms). The advertisement call of S. arduous (4���6 notes and duration of 198���328 ms; Pombal & Bastos 2003) is shorter than that of the new species (the calls of Scinax cosenzai sp. nov. with 4���6 notes have duration of 812.25���1395.75 ms). The territorial call of S. arduous differs from the vocalization of the new species in having only one note (Pombal & Bastos 2003) (the vocalization of Scinax cosenzai sp. nov. has at least 2 notes). The advertisement call of S. perpusillus (3���6 notes and duration of 92���174 ms; Pombal & Bastos 2003) is shorter than that of the new species (the calls of Scinax cosenzai sp. nov. with 3���6 notes have duration of 512.75���1395.75 ms). Pombal and Bastos (2003) described a second type of call of S. perpusillus, also shorter than that of the new species, with 6���12 notes and duration of 314���400 ms (the calls of Scinax cosenzai sp. nov. with 6���12 notes have duration of 892.81���2066.75 ms). These authors also mentioned the existence of a third type of call with only one note and duration of 185 s, 435 s and 508 s (the calls of Scinax cosenzai sp. nov. have at least two notes). Description of holotype. (Figs. 1, 2) Male; body slender; head longer than wide; snout protruding in lateral view and acuminated in dorsal view; nostrils rounded; prominent medial process between the nostrils; canthus rostralis distinct; loreal region concave; eye slightly protuberant; tympanum distinct; discrete subgular vocal sack; tongue round and large; vomerine teeth in two arc shape series between choanae; medium-sized, elliptical choanae. Arms slender; forearms slightly robust; fingers slender and medium-sized, relative lengths IMeasurements of the holotype (mm). SVL 19.34, HEL 7.05, HW 6.94, THL 8.87, TBL 10.01, FL 7.27, HL 5.18, ED 2.53, TD 1.05, IOD 2.25, END 2.42, IND 1.95. nant frequency of Scinax cosenzai sp. nov.. *One of the calls with one pair of merged notes **Two of the calls with one pair of merged notes. 7 (3)* 1338.48 (1247.06���1428.63; 183.68 (30.00��� 372.93; 4027.82 (3877.77���4316.09; SD = 90.79) SD = 121.72; n= 17) SD = 249.72) 10 (2)** 942.75; 1132.19 68.53 (13.75���210.75; 3949.33; 4135.09 SD = 54.74; n= 16) 11 (3)* 1380.33 (1062.75���1580.25; 88.07 (18.18���403.43; 3984.85 (3941.26���4070.48; SD = 278.09) SD = 98.73; n= 29) SD = 74.16) 12 (3)* 1748.08 (1266.12���2066.75; 106.81 (10.37���443.25; 4568.52 (4563.14���4571.21; SD = 242.26) SD = 125.22; n= 32) SD = 4.66) 13 (1) 1790.50 94.24 (23.37���487.81; 4175.47 SD = 134.43; n= 12) 14 (1) 1784.69 87.16 (10.00��� 381.87; 4127.01 SD = 109.69; n= 13) Variation. In preservative, as in life, coloration of dorsum varies from clear-gray to dark-gray (Fig. 2). Interorbital and dorsal bars are not evident in some individuals. Webbing formulae varies among specimens I ��� II 2 ��� 3 + III (2 + ��� 2 -) ��� (3 + ��� 3) IV 3 + ��� 2 V. Females are larger than males. Range, mean and standard deviation of the measurements of 30 males and seven females are given in Table 2. Character Male (N = 30) Female (N = 7) Natural history notes. Calling activity was more common during the rainy season at night. As observed in others species from the S. perpusillus group (Oliveira & Navas 2004; Brasileiro et al. 2007 a), Scinax cosenzai sp. nov. was more frequently found in clustered bromeliads. We observed a few spawnings, with one or two eggs each, stuck to lateral leaves (as also reported for other species of the group by Alves-Silva & Silva 2009) a few centimeters above the water accumulated by the bromeliad. Tadpoles were also observed in the water of bromeliads and frequently two individuals were observed sharing the same leaf axil. When we were noticed by the tadpoles they usually dived to the bottom of the accumulated water, as also reported for Scinax tupinamba by Silva and Alves- Silva (2009). Distribution. The new species seems to be abundant in Alcantarea extensa (Bromeliaceae), a ground bromeliad commonly found in the rocky outcrops from Parque Estadual da Serra do Brigadero���PESB (Fig. 5), a conservation unit with approximately 13,000 ha from the Atlantic Rain Forest biome managed by Instituto Estadual de Florestas���IEF, in the northern part of the Mantiqueira Mountain Range Complex, State of Minas Gerais (Cruz et al. 2007; Caramaschi et al. 2008). We collected specimens from two rocky outcrops: the type locality, located in the PESB, and another outcrop, located in the surroundings of the PESB (Fig. 5). As Alcantarea extensa is commonly found in others rocky outcrops from the Zona da Mata region in the State of Minas Gerais (Versieux & Wendt 2006), we believe that the new species may also occur in some of these other rocky outcrops and may not be in danger of extinction. Etymology. The specific name honors the biologist Braz Ant��nio Pereira Cosenza for his contributions to the studies of flora and fauna of Serra do Brigadeiro. Additional remarks. As observed by Silva and Alves-Silva (2008), species from the Scinax perpusillus group from outside the geographic limits sampled by Peixoto (1987), usually referred to as Scinax perpusillus, Scinax aff. perpusillus or Scinax gr. perpusillus (see Heyer et al. 1990; Pombal & Gordo 2004; Oliveira & Navas 2004; Carvalho-e-Silva et al. 2008; Feio et al. 2008; Bertoluci et al. 2009; Lacerda et al. 2009), may represent species not yet recognized and described. In spite of such problematic taxonomy, the advertisement call of only five species has been somewhat described (Scinax arduous, S. insperatus, S. peixotoi, S. perpusillus and S. v-signatus). However, the vocalization of these species has been described in two different manners: Heyer et al. 1990, Pombal and Bastos (2003) and Brasileiro et al. 2007 a described each call of a call series while Alves-Silva and Silva (2009) and Silva and Alves-Silva (2008) referred to the entire call series as a single call, referring to notes and pulses what previous studies considered respectively as single calls and notes. We agree with Silva and Alves-Silva (2011) in believing that these previous studies did not consider the social interactions during vocalizations and that the described vocalization probably represents only a part of the entire call (herein referred to as call series). We did not compare the vocalization of S. cosenzai sp. nov. to those described for S. insperatus, S. perpusillus and S. v-signatus presented by Alves-Silva and Silva (2009) and Silva and Alves-Silva (2011) because these studies described only duets, or calls emitted when a female approached the male, while the recorded vocalizations from S. cosenzai sp. nov. were emitted by isolated males. We agree with Alves-Silva and Silva (2009) and Silva and Alves-Silva (2011) in considering further investigation necessary in order to establish the universality of call structure for the species from the S. perpusillus group. As observed in Figure 6, the type localities of Scinax alcatraz, S. faivovichi and S. peixotoi are associated to coastal islands from the state of S��o Paulo (until now, considered endemic to these islands). Scinax littoreous, S. perpusillus and S. tupinamba have type localities associated to lowland coastal areas in the state of Rio de Janeiro. Scinax arduous and S. belloni have distribution associated to mountain regions in the state of Esp��rito Santo, while S. insperatus, S. melloi and S. v-signatus are found in the state of Rio de Janeiro and S. atratus in the state of S��o Paulo. Scinax cosenzai sp. nov. is the first species belonging to the S. perpusillus group described in the State of Minas Gerais. Bertoluci et al. 2009 provided the record of specimens referred to as Scinax sp. 3 (gr. perpusillus) in the Esta����o Ambiental de Peti, in Minas Gerais state, but they only cited the field number, not mentioning the deposition of voucher specimens in any public collection. If this identification is c, Published as part of Lacerda, Jo��o Victor A., Peixoto, Oswaldo Luiz & Feio, Renato N., 2012, A new species of the bromeligenous Scinax perpusillus group (Anura; Hylidae) from Serra do Brigadeiro, State of Minas Gerais, Southeastern Brazil, pp. 31-42 in Zootaxa 3271 on pages 32-41, DOI: 10.5281/zenodo.280807, {"references":["Brasileiro, C. A., Haddad, C. F. B., Sawaya, R. J. & Martins, M. (2007 a) A new and threatened species of Scinax (Anura: Hylidae) from Queimada Grande island, southeastern Brazil. Zootaxa, 1391, 47 - 55.","Faivovich, J., Gasparini, J. L. & Haddad, C. F. B. (2010) A New Species of the Scinax perpusillus Group (Anura: Hylidae) from Espirito Santo, Brazil. Copeia, 1, 97 - 102.","Peixoto, O. L. (1988 b) Duas novas especies de Ololygon do Grupo \" perpusilla \" (Amphibia, Anura, Hylidae). Arquivos da Universidade Federal Rural do Rio de Janeiro, 10, 27 - 37.","Brasileiro, C. A., Oyamaguchi, H. M. & Haddad, C. F. B. (2007 b) A new island species of Scinax (Anura; Hylidae) from southeastern Brazil. Journal of Herpetology, 41, 271 - 275.","Silva, H. R & Alves-Silva, R. (2011) A new bromeligenous species of the Scinax perpusillus group from the hills of the State of Rio de Janeiro, Brazil (Anura, Hylidae). Zootaxa, 3043, 54 - 68.","Silva, H. R. & Alves-Silva, R. (2008) New coastal and insular species of the bromeligenous Scinax perpusillus group, from the State of Rio de Janeiro, Brazil (Anura, Hylidae). Zootaxa, 1914, 34 - 44.","Lutz, B. (1973) Brazilian species of Hyla. Austin & London: University of Texas Press: 260 pp.","Peixoto, O. L. (2002) Uma nova especie de Scinax do grupo \" perpusillus \" para Santa Teresa, Estado do Espirito Santo, Brasil (Amphibia, Anura, Hylidae). Boletim do Museu Biologico Mello Leitao, 13, 7 - 15.","Pombal, Jr., J. P. & Bastos R. P. (2003) Vocalizacoes de Scinax perpusillus (A. Lutz & B. Lutz) e S. arduous Peixoto (Anura, Hylidae), com comentarios taxonomicos. Revista Brasileira de Zoologia, 20, 607 - 610.","Oliveira, F. B. & Navas, C. A. (2004) Plant selection and seasonal patterns of vocal activity in two population of Bromeligen treefrog Scinax perpusillus (Anura; Hylidae). Journal of Herpetology, 38, 331 - 339.","Alves-Silva, R. & Silva, H. R. (2009) Life in bromeliads: reproductive behaviour and the monophyly of the Scinax perpusillus species group (Anura: Hylidae). Journal of Natural History, 43, 205 - 217.","Cruz, C. A. G., Feio, R. N. & Cassini, C. S. (2007) Nova especie de Chiasmocleis Mehely, 1904 (Amphibia, Anura, Microhylidae) da Serra da Mantiqueira, Estado de Minas Gerais, Brasil. Arquivos do Museu Nacional, Rio de Janeiro, 65, 33 - 38.","Caramaschi, U., Feio, R. N. & Sao Pedro, V. A. (2008) A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae) from Serra do Brigadeiro, State of Minas Gerais, Southeastern Brazil. Zootaxa, 1861, 44 - 54.","Versieux, L. M. & WENDT, T. (2006) Checklist of the Bromeliaceae of Minas Gerais, Brazil, with notes on taxonomy and endemism. Selbyana, 27, 107 - 146.","Peixoto, O. L. (1987) Caracterizacao do grupo perpusilla e revalidacao da posicao taxonomica de Ololygon perpusilla perpusilla e Ololygon perpusilla v-signata (Amphibia, Anura, Hylidae). Arquivos da Universidade Federal Rural do Rio de Janeiro, 10, 37 - 49.","Heyer, W. R., Rand A. S., Cruz C. A. G., Peixoto O. L. & Nelson C. E. (1990) Frogs of Boraceia. Arquivos de Zoologia, 31, 231 - 410.","Pombal Jr., J. P. & Gordo, M. (2004) Anfibios anuros da Jureia. In: Marques, O. A. V. & Duleba, W. (Eds.) Estacao Ecologica Jureia-Itatins. Editora Holos Ribeirao Preto, pp. 243 - 256.","Carvalho-e-Silva, A. M. T., Silva, G. R. & Carvalho-e-Silva, S. P. (2008) Anuros da reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Biota Neotropica, 8, 199 - 209.","Bertoluci, J., Canelas, M. A. S., Eisemberg, C. C., Palmuti, C. F. S. & Montingelli, G. G. (2009) Herpetofauna da Estacao Ambiental de Peti, um fragmento de Mata Atlantica do estado de Minas Gerais, sudeste do Brasil. Biota Neotropica, 9, 147 - 155.","Lacerda, J. V. A., Assis, B., Santana, D. J. & Feio, R. N. (2009) Anurans in bromeliads, Parque Estadual da Serra do Brigadeiro, state of Minas Gerais, southeastern Brazil. Check List, 5, 800 - 806."]}

Leptodactylus cupreus sp. nov. (Figures 1–3) Holotype: MNRJ 47752. Adult male (Figure 1). Lagoa das Bromélias (20 o 25 ’S, 43 o 29 ’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. São­Pedro, on 0 8 October 2006. 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. São­Pedro, and P.S. Silva, on 24 October 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 surface of the thighs, posterior surface of tarsus smooth or with few tubercles, and sole of foot with prominent 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, delimited 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, dorsal 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. 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 copper 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 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 pattern) 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 distinct 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 sub­elliptical viewed from above (Fig. 2 A), protruding with distinct shelf in profile (Fig. 2 B); 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 distance, 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 posterior 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 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 urostyle. 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 post­commissural gland distinctly golden. Flanks, below the black dorsolateral 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 lateral head, dorsolateral body, arms, and legs are maintained; bucal post­commissural 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. 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. 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. Habitat and habits: The type locality of L. cupreus is essentially the same as recently described for Chiasmocleis mantiqueira (Anura, Microhylidae; see Cruz et al. 2007). The Parque Estadual da Serra do Brigadeiro, in the Atlantic Rain Forest 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 pond at 1,227 m altitude, that completely dryes­ up during the dry season (April to September), but with about 250 m 2 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 tadpoles are unknown. Etymology: The specific epithet, “ cupreus ”, is a Latin adjective referred to the copper general color pattern of the new species.

Leptodactylus cupreus sp. nov. (Figures 1���3) Holotype: MNRJ 47752. Adult male (Figure 1). Lagoa das Brom��lias (20 o 25 ���S, 43 o 29 ���W, 1,227 m above sea level), Parque Estadual da Serra do Brigadeiro, District of Care��o, Municipality of Erv��lia, State of Minas Gerais, Southeastern Brazil. Collected by R.N. Feio and V.A. S��o��Pedro, on 0 8 October 2006. 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. S��o��Pedro, and P.S. Silva, on 24 October 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 surface of the thighs, posterior surface of tarsus smooth or with few tubercles, and sole of foot with prominent 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, delimited 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, dorsal 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. 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 copper 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 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 pattern) 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 distinct 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. 2 A), protruding with distinct shelf in profile (Fig. 2 B); 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 distance, 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 posterior 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 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 urostyle. 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 dorsolateral 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 lateral 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. 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. 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. Habitat and habits: The type locality of L. cupreus is essentially the same as recently described for Chiasmocleis mantiqueira (Anura, Microhylidae; see Cruz et al. 2007). The Parque Estadual da Serra do Brigadeiro, in the Atlantic Rain Forest 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 Brom��lias (local name meaning Bromeliads��� Lake) is a temporary pond at 1,227 m altitude, that completely dryes�� up during the dry season (April to September), but with about 250 m 2 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 tadpoles are unknown. Etymology: The specific epithet, ��� cupreus ���, is a Latin adjective referred to the copper general color pattern of the new species., Published as part of Caramaschi, Ulisses, Feio, Renato N. & S��o-Pedro, Vin��cius A., 2008, A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae) from Serra do Brigadeiro, State of Minas Gerais, Southeastern Brazil, pp. 44-54 in Zootaxa 1861 on pages 45-51, DOI: 10.5281/zenodo.183691, {"references":["Heyer, W. R. (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Natural History Museum of Los Angeles County, Science Bulletin, 29, 1 - 85.","Heyer, W. R., Garcia-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.","Cruz, C. A. G., Feio, R. N. & Cassini, C. S. (2007) Nova especie de Chiasmocleis Mehely, 1904 (Amphibia, Anura, Microhylidae) da Serra da Mantiqueira, Estado de Minas Gerais, Brasil. Arquivos do Museu Nacional, Rio de Janeiro, 65 (1), 33 - 38."]}