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1 Zootaxa 4061 (3): Copyright 2016 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) A new species of Scinax Wagler (Anura: Hylidae) from Mato Grosso, Brazil KATYUSCIA ARAUJO-VIEIRA 1,4, PAULA HANNA VALDUJO 2 & JULIÁN FAIVOVICH 1,3 1 División Herpetología, Museo Argentino de Ciencias Naturales Bernardino Rivadavia -CONICET, Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina 2 Laboratório de Ecologia da Paisagem, WWF-Brasil, SHIS QL 6/8, Conjunto E, Lago Sul, Brasília, DF, , Brazil 3 Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos Aires, Argentina 4 Corresponding author. katy.vieira@gmail.com Abstract A new species of the Scinax ruber Clade is described from Municipality of Barra do Garças, State of Mato Grosso, Brazil. It is diagnosed by its size (SVL mm in males); dorsum with a background that varies from light and dark gray to dark brown, with round and irregular dark blotches; hidden surfaces of thigh and shank light or dark brown, with lighter, large and irregular blotches (yellow in life); and nuptial pad that covers the medial margin of Metacarpal I, and obscures the posterior portion and the outer margin of the inner metacarpal tubercle. It is most similar with Scinax rupestris, from which it differs by having larger adult males; less developed nuptial pad; and hidden surfaces of thigh and light or dark brown, with lighter, large and irregular blotches (yellow in life). The new species occurs near the headwaters of permanent or temporary streams, where males call from vegetation or rocks at the margins of small streams. Key words: Scinax haddadorum sp. nov., morphology, taxonomy, Brazilian Cerrado Introduction The neotropical hylid genus Scinax Wagler includes 114 described species ranging from Mexico to East-Central Argentina, including Trinidad and Tobago, and St. Lucia (Frost 2015). The genus comprises two large clades, the S. catharinae and the S. ruber clades (Faivovich 2002; Faivovich et al. 2005). The latter is the most species rich including 68 species, of which 57 are not included in the two monophyletic groups currently recognized: the S. rostratus and S. uruguayus groups (Faivovich et al. 2005; Frost 2015). The species of the S. ruber Clade occupy diverse habitats, including both open and forested areas, and most frequently reproduce on temporary or permanent lentic water bodies (e.g. Duellman 1970; Cardoso & Sazima 1980; Pombal et al. 1995a). Fifteen species of this clade have been recorded from the Brazilian Cerrado domain: Scinax acuminatus (Cope), S. cabralensis Drummond, Baêta & Pires, S. constrictus Lima, Bastos & Giaretta, S. curicica Pugliese, Pombal & Sazima, S. fuscomarginatus (Lutz), S. fuscovarius (Lutz), S. maracaya (Cardoso & Sazima), S. montivagus Juncá, Napoli, Nunes, Mercês & Abreu, S. nasicus (Cope), S. pinima (Bokermann & Sazima), S. similis (Cochran), S. rogerioi Pugliese, Baêta & Pombal, S. rupestris Araujo-Vieira, Brandão & Faria, S. squalirostris (Lutz), and S. tigrinus Nunes, Carvalho & Pereira. In this paper, we describe a new species of the Scinax ruber Clade collected during fieldwork in the Municipality of Barra do Garças, State of Mato Grosso, Brazil. It is the sixteenth Scinax species that occurs in the Brazilian Cerrado domain. Material and methods Adult specimens were fixed in 10% formalin and stored in 70% ethanol. Webbing formula follows Savage & Heyer Accepted by V. Orrico: 20 Nov. 2015; published: 7 Jan

2 (1967) as modified by Myers & Duellman (1982). Dorsal and profile outline standards of snout shape follow Duellman (1970). Measurements (in millimeters) were taken with a digital caliper under a stereomicroscope and rounded to the nearest 0.1 mm. Ten measurements are those employed by Duellman (1970): SVL (snout-vent length), HL (head length), HW (head width), IND (internarial distance), IOD (interorbital distance), ED (eye diameter), END (eye-nostril distance), TD (tympanum diameter), TL (tibia length), and FL (foot length). We also took two other measurements, 3FD (third finger disc diameter) and 4TD (fourth toe disc diameter) employed by Napoli & Caramaschi (1998). Sex was determined by examination of secondary sexual characters (nuptial pads, vocal slits, and expansion of the vocal sac). Illustrations were made with a drawing tube attached to a Nikon steromicroscope (SMZ-800). Institutional abbreviations follow Sabaj Pérez (2014). Results Scinax haddadorum sp. nov. (Figs. 1 3) Scinax aff. cruentommus Valdujo et al. (2012: 76). Species list associated to an analysis of amphibian species distribution in the Brazilian Cerrado. Holotype. MZUSP , adult male, from Fazenda Água Limpa (15 41'34'' S, 52 20'59'' W, 675 m a.s.l.), Municipality of Barra do Garças, State of Mato Grosso, Brazil, collected on January 31, 2009 by Paula Hanna Valdujo, Renato Recoder, and Mauro Teixeira Júnior. Paratypes. Twelve adults collected in four localities in the Municipality of Barra do Garças, State of Mato Grosso, Brazil. All males unless otherwise stated: MZUSP , collected together with the holotype. MZUSP , , Parque Estadual Serra Azul (15 51'28'' S, 52 15'16'' W) collected on November 15, 2008 by Paula Hanna Valdujo, Renato Recoder, and Agustín Camacho. CFBH 39054, Serra do Roncador 6 km E BR-158 (15 20'37'' S, 52 14'05'' W) collected on December 17, 2008 by Paula Hanna Valdujo and Mara Souza de Albuquerque e Silva. MZUSP , CFBH 39055, (female), 17 km ESE BR- 158 (15 41'29'' S, 52 24'48'' W) collected on February 1, 2009 by Paula Hanna Valdujo, Renato Recoder, and Mauro Teixeira Júnior. Diagnosis. Scinax haddadorum sp. nov. can be diagnosed by the following set of characters: SVL in males mm; snout slightly acuminate in dorsal view, rounded in profile; vocal sac single, median, subgular that ventrally does not reach the pectoral region; nuptial pad that covers the medial margin of Metacarpal I, and obscures the posterior portion and the outer margin of the inner metacarpal tubercle; tibia length/snout-vent length = ; absence of conspicuous pointed tubercle on the heel, and pointed tubercles on the lower jaw; iris golden yellowish, with many thin, darker reticulations; dorsum with a background that varies from light and dark gray to dark brown, with round and irregular dark blotches; and hidden surfaces of thigh and shank light or dark brown, with lighter, large and irregular blotches (yellow in life). Comparison with other species. The structure for the comparison is based first on obvious size differences in adults (that is, no overlapping nor a minimal gap between size ranges), followed by more detailed comparisons with species that cannot be differentiated on the basis of size, or in case that there are conspicuous external morphological characters. The SVL in males ( ) promptly distinguish the new species from Scinax acuminatus ( ; Lutz 1973), S. altae ( ; Duellman 1970), S. cabralensis ( ; Drummond et al. 2007), S. castroviejoi (male holotype 45.0; De la Riva 1993), S. cruentommus ( ; Duellman & Wiens 1993), S. danae ( ; Duellman 1986), S. eurydice ( ; Bokermann 1968), S. exiguus ( ; Duellman 1986), S. fuscovarius ( ; Cei 1980), S. fuscomarginatus ( ; Brusquetti et al. 2014), S. hayii ( ; Lutz 1973), S. karenanneae ( ; Pyburn 1993), S. madeirae ( ; Brusquetti et al. 2014), S. maracaya ( ; Cardoso & Sazima 1980), S. staufferi ( ; Duellman 1970), S. villasboasi ( ; Brusquetti et al. 2014), and S. wandae ( ; Pyburn & Fouquette 1971). Scinax haddadorum sp. nov. differs from all species in the S. rostratus group (sensu Faivovich 2002) by the absence of conspicuous pointed tubercle on the heel and pointed tubercles on the lower jaw, and by its rounded snout in profile (elongate, pointed snout in profile; Duellman 1972). 262 Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

3 FIGURE 1. Scinax haddadorum sp. nov., holotype (MZUSP ; SVL 30.4 mm). A: Dorsal view. B: Ventral view. The golden yellowish iris, with many thin, darker reticulations distinguishes the new species from Scinax cruentommus (silvery bronze iris with a median horizontal red streak; Duellman 1972), S. funereus (greenish bronze with brown flecks; Duellman & Wiens 1993), S. oreites (dull bronze with median horizontal brown streak; Duellman & Wiens 1993), and the species in the S. uruguayus group (bicolored iris; Bokermann & Sazima 1973; Faivovich et al. 2005). The dorsal color pattern with a background that varies from light and dark gray to dark brown, with round and irregular dark blotches differentiates the new species from Scinax altae, S. cardosoi, S. fuscomarginatus, S. madeirae, S. pachycrus, S. quinquefasciatus, S. ruber, S. squalirostris, S. staufferi, and S. villasboasi (dorsum with a variable number of dorsal and/or lateral stripes; Duellman 1970; Duellman 1972; Duellman & Wiens 1993; Lutz 1973; Heyer et al. 1990; Carvalho-e-Silva & Peixoto 1991, Brusquetti et al. 2014), S. alter, S. auratus, S. cretatus, S. crospedospilus, S. cuspidatus, S. imbegue, S. juncae, and S. tymbamirim (light or dark dorsal continuous or broken stripes, sometimes delimiting a central darker area; Bokermann 1969; Lutz 1973; Nunes & Pombal 2010, 2011; Nunes et al. 2012), S. blairi (few brown markings and blotches, or small scattered dark dots; Fouquette & Pyburn 1972), S. boesemani (dorsum with or without small white and brown dots; Lescure & Marty 2000), S. cabralensis (dorsum with small dark spots homogeneously distributed; Drummond et al. 2007), S. caldarum, S. curicica, and S. duartei (two irregular longitudinal stripes arising from an interocular marking; Pugliese et al. 2004), S. chiquitanus (small and scattered grayish dots and marks; De la Riva 1990), S. danae (small scattered dark brown dots), S. dolloi and S. perereca (dorsum with indistinct light pattern or with dark spots, interocular marking and a pair of inverted dorsolateral parentheses; Pombal et al. 1995b), S. elaeochrous (dorsum with darker markings and dots, sometimes forming irregular longitudinal stripes; Duellman 1970), S. granulatus (dorsum with clear spots and dark longitudinal X-shaped lines and an interocular marking; Kwet & Di-Bernardo 1999), S. ictericus (dorsum with or without dark brown interocular marking and irregular mostly transverse marks; Duellman & Wiens 1993), S. iquitorum (small and scattered dark brown dots and blotches; Moravec et al. 2009), S. lindsayi (a few small scattered reddish brown dots and blotches), S. maracaya and S. tigrinus (large dark blotches; Cardoso & Sazima 1980; Nunes et al. 2010), and S. rogerioi (brown dorsal blotches extending as a pair of longitudinal irregular and interrupted blotches/stripes from head to inguinal region and inverted brown triangular interocular blotch; Pugliese et al. 2009). NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 263

4 FIGURE 2. Scinax haddadorum sp. nov., holotype (MZUSP ). A: Head in profile. B: Right hand in ventral view. C: Right foot in ventral view. Scale bar = 2 mm. 264 Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

5 FIGURE 3. Nuptial pad of the left hand of Scinax haddadorum sp. nov., holotype (MZUSP ). A: Dorsal view. B: Ventral view. Scale bar = 1 mm. FIGURE 4. Dorsal pattern variation in the type series of Scinax haddadorum sp. nov.. A: MZUSP (SVL 35.4 mm). B: MZUSP (SVL 32.4 mm). C: MZUSP (SVL 31.2 mm). D: MZUSP (SVL 34.1 mm). E: CFBH (SVL 31.8 mm). F: MZUSP (SVL 31.2 mm). The vocal sac single, median, subgular, and that ventrally does not reach the pectoral region differentiates the new species from Scinax baumgardneri, S. exiguus, S. fuscomarginatus, S. madeirae, S. manriquei, S. villasboasi, and S. wandae (vocal sac large that reaches the anterior pectoral region; Barrio-Amorós et al. 2004; Brusquetti et al. 2014); S. camposseabrai, S. karenanneae and S. sateremawe (bilobed vocal sac; Pyburn 1993; Caramaschi & Cardoso 2006; Sturaro & Peloso 2014). NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 265

6 Scinax haddadorum sp. nov. differs from S. nasicus and S. similis for having an slightly acuminate snout in dorsal view (rounded in dorsal view). Also, it differs from S. nasicus by having generally a longer shank in relation to its SVL (TL/SVL = in the new species, and TL/SVL = , n = 20 in S. nasicus). The holotype of Scinax x-signatus (Spix 1824) is considered lost (Hoogmoed & Gruber 1983; Glaw & Franzen 2006), a neotype has still not been designated, and apparently there are several species to which this name has been applied in the State of Bahia, Brazil (its type locality is Provinciae Bahiae, meaning current State of Bahia east of the river San Francisco; Spix 1824; Vanzolini 1981; see Pombal et al. 1995b). The dorsal color pattern which consists of a background that varies from light and dark gray to dark brown, with round and irregular dark blotches in S. haddadorum sp. nov. differentiates this species from those to which the name S. x-signatus could potentially be applied in northeastern Brazil (X-shaped dorsal pattern composed of one or two pairs of inverted parentheseslike marks; Lutz 1973). The new species is distinguished from Scinax montivagus for having a slightly acuminate snout in dorsal view (rounded in dorsal view; Juncá et al. 2015); dorsal surface of forelimbs and thighs with irregular, transverse dark brown markings (absence of brown markings; Juncá et al. 2015); and hidden surfaces of thigh light or dark brown, with lighter, large and irregular blotches, which are yellow in live specimens (immaculate beige, without yellow coloration in live specimens; Juncá et al. 2015). Scinax haddadorum sp. nov. is similar with S. rupestris (see Figs. 1 3 in Araujo-Vieira et al. 2015) from which it differs mainly for having SVL mm in males (SVL ; Araujo-Vieira et al. 2015); a nuptial pad that covers the medial margin of Metacarpal I, and obscures the posterior portion and the outer margin of the inner metacarpal tubercle (wide nuptial pad that covers almost the complete dorsal surface of Metacarpal I and obscures nearly half of the inner metacarpal tubercle; Araujo-Vieira et al. 2015); and hidden surfaces of thigh and shank light or dark brown, with lighter, large and irregular blotches (light or dark brown, with light, small diffuse blotches; Araujo-Vieira et al. 2015). Description of holotype. Body moderately robust; head as large as wide, 40.1% of SVL. Snout rounded in profile, slightly acuminated in dorsal view. Nostrils dorsolateral, elliptical, slightly protruded; distance between nostrils 79.3% of IOD. Canthus rostralis slightly evident and convex. Loreal region slightly concave. Eyes protuberant, ED 34.5% larger than IOD, almost equal to END. Tympanum nearly rounded, separated from eye by a distance almost half TD. TD 59% of ED. Tympanic annulus rounded, with upper portion hidden by supratympanic fold. Supratympanic fold barely evident, from the upper portion of the tympanum to the insertion of the arm. Vocal sac single, median, subgular, externally evident by the loose skin on the sides of jaw, ventrally not reaching the pectoral region. Vocal slits present, located diagonally to the longitudinal body axis, originating laterally to the tongue and running towards the corner of the mouth. Tongue ovoid, free laterally and posteriorly notched. Vomerine teeth in two slightly convex series between and only just posterior to the choanae, each bearing seven (right) and six (left) teeth. Choanae oval. Axillary membrane absent. Upper arm more slender than forearm. A series of small, round, low, and enameled ulnar tubercles (right forearm: one tubercle; left forearm: five tubercles in row). Fingers, short and slender, fringed. Relative finger length I<II< IV<III. Discs elliptical, wider than long; disc on Finger I smaller than the others. Subarticular tubercles single, conical on fingers I and II; rounded and quadrangular on fingers III and IV, respectively. Subarticular tubercle on Finger IV larger than the others. Supernumerary tubercles small, single, and rounded. Inner metacarpal tubercle single, medium-sized, elliptical; outer metacarpal tubercle flat, bilobed (right hand) or divided (left hand), the two parts are elongated in the latter condition. Webbing absent between fingers I and II; basal between fingers II, III, and IV. The lateral fringe on Finger IV is continued in the lateral margin of the palm by an enameled dermal ridge, from the penultimate subarticular tubercle back to the level of distal margin of outer metacarpal tubercle. Thick, wide, light colored nuptial pad covering only the medial margin of Metacarpal I, and obscuring the outer margin of the inner metacarpal tubercle (Fig. 3A B). Hind limbs robust; TL 51.6% of SVL. Toes slender, fringed. Relative toe length I< II< V III< IV. Discs elliptical, wider than long. Subarticular tubercles single, conical, and rounded. Supernumerary tubercles small, single, and rounded. Inner metatarsal tubercle single, oval; outer metatarsal tubercle simple, elliptical, one third the size of inner metatarsal tubercle, located with distal margin at the level of distal margin of inner metatarsal tubercle or slightly offset posteriorly. Webbing formula I II 1 1/3 2 III 1 1/2 2 + IV V. Fringe on lateral margin on Toe V continued along the margin of the sole by a poorly developed ridge that reaches the base of the metatarsal. Along the medial margin of the tarsus, an inconspicuous medial tarsal ridge, barely evident as a thickening of the skin. A 266 Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

7 series of small, round, inconspicuous tarsal tubercles. Three discrete, low tubercles, next to the tibiotarsal articulation on the right tarsus; tubercles absent on the left tarsus. Skin on dorsum, upper eyelid, and dorsal region of limbs smooth, with small, scattered, rounded tubercles on head and upper eyelid. Small, round, and white pigmented tubercles present on the posterior portion of the tympanum, near the insertion of the arm. Vocal sac smooth; pectoral region, abdomen and subcloacal regions densely covered with rounded, low granules. Ventral surfaces of arms and tibiae smooth. Pectoral fold with preand postaxillar elements. There is a white, slightly thickened area in the pectoral region that is apparently glandular, but this requires histological corroboration. Coloration in life of the holotype. There are no notes of coloration in life of the holotype. Coloration in preservative of the holotype. Dorsum light brown with scattered round, irregular, and elongate dark blotches; tympanic membrane finely spotted; canthus rostralis dark brown, loreal region with same coloration as dorsum. Two diagonal dark brown blotches on upper eyelid. Upper lip light cream with diffuse limits. Dorsal surface of fore- and hindlimbs with irregular, transverse dark brown markings. Dorsal surface of hands and discs with dark brown dots and irregular transversal blotches on fingers III and IV, like those on forearms. Flanks and inguinal region lighter with irregular dark blotches. Hidden area of thigh and shank dark brown with some large and irregular light blotches. Ventrally, soles brown, palms light brown; other ventral areas creamy white, immaculate, but margins of the gular region, around arm insertion, and bordering areas with the dorsal coloration finely spotted with brownish gray. The iris is black, with many thin, silver reticulations. Measurements of the holotype (mm). SVL 30.4; HL 12.2; HW 11.8; IND 2.3; IOD 2.9; ED 3.8; END 3.6; TD 2.3; TL 15.7; FL 13.7; 3FD 1.7; 4TD 1.6. Variation in the type series. Some measurements are presented in Table 1. The number of vomerine teeth in males varies between four and seven on both the right and left processes. In one male paratype (MZUSP ) the right process (five teeth) and the left process (six teeth) are more separated than those of the other specimens (about half of the process size). Dorsally, small, scattered, round tubercles are evident mainly on head and body, sometimes on upper eyelid, and less commonly on limbs. Ulnar tubercles are absent in two individuals; when present they can be arranged in line (seven individuals; between two and six tubercles), or scattered (three individuals; between two and five tubercles). In six individuals these tubercles are white. The white dermal ridge on lateral margin of the palm is complete (five individuals) or interrupted (eight individuals). Two specimens (MZUSP ; ) have the nuptial pad thicker than those of other individuals. The granules on the subcloacal region are white in nine individuals. TABLE 1. Some measurements (in mm) of the type series of Scinax haddadorum sp. nov. Measurements Males (n = 12) Females (n = 1) Mean ± SD Range SVL 31.8 ± HL 12.3 ± HW 11.6 ± IND 2.2 ± IOD 3.2 ± ED 3.8 ± END 3.5 ± TD 2.2 ± TL 16.9 ± FL 14.0 ± FD 1.5 ± TD 1.5 ± Tarsal tubercles on the external surface of the tarsus are absent in four individuals. When present these tubercles can be enameled (two individuals). The number of tubercles present on the internal margin of the tarsus, NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 267

8 next to the tibio-tarsal articulation, varies from one to five; the distal one is elongated in MZUSP ; and the tubercles are enameled in five individuals. Small calcar tubercles, round and low are present in six individuals. Webbing formulae among paratypes varies as follows: I ( ) (2 + 2) II (1 1/2 1 - ) (2 + 2) III (1 1/2 1 - ) (2 ½ 2) IV ( ) ( ) V. The only available female (SVL 35.7 mm) is slightly larger than the largest male paratype (MZUSP ; SVL 35.4 mm). Similar to Scinax rupestris (see Araujo-Vieira et al. 2015), some specimens have a thickening of the ventral portion of the webbing between toes IV and V that is also evident on the dorsal aspect as a number of wrinkles or an ampulla-like swelling. Unidentified eggs are present in the webbing in two individuals (MZUSP ). This is apparently common in several species of Scinax (Faivovich pers. obs.). A photograph in life is available only from paratype MZUSP (Fig. 6A). The specimen is dorsally yellowish tan with brown markings. Two diagonal blotches on upper eyelid, an interocular fragmented blotch continuous to the diagonal blotches. Loreal region with the same coloration as dorsum, and a light brown canthal line. Small round blotches, and larger, irregular blotches in the dorsum with a pair of elongate, irregular blotches on scapular and sacral regions. Hidden surfaces of thigh light brown with irregular yellow blotches. Iris golden yellowish, with many thin, darker reticulations. In preservative, the dorsal color pattern of head, body and limbs varies from light and dark gray to dark brown, with round and irregular dark blotches (Fig. 4). A dark brown canthal line is present in seven individuals, and it is fragmented in two individuals (MZUSP ; ). The diagonal dark blotches on the upper eyelids can be lighter or darker than in the holotype. They are absent in MZUSP and The interocular region can show an inverted triangular or V-shaped dark blotch, sometimes fragmented. Upper lip light cream with diffuse limits finely or conspicuously pigmented. A pair of elongate blotches is present in the scapular and sacral regions of some individuals. Transversal blotches on limbs are continuous or fragmented. They are absent on the shanks in MZUSP Flanks and inguinal region light colored, with irregular dark blotches. Hidden surfaces of thigh and shank light or dark brown, with lighter, large and irregular blotches. Ventrally, soles and palms, margins of the gular region, arm insertion, and bordering areas with the dorsal coloration vary from creamy white to dark brown. The pectoral region can be finely pigmented with dark brown in some individuals. One individual (MZUSP ) has a series of lighter regions on the body, as if the dorsal pattern has faded, with several lightly pigmented areas; we are uncertain if this represents a natural condition or a fixation artifact. Tadpoles and advertisement call. The tadpole is unknown, and the advertisement call has not been recorded. Geographic distribution and Natural history. Scinax haddadorum sp. nov. has been found in four different localities in the Municipality of Barra do Garças, State of Mato Grosso, Brazil (Fig. 5). All streams where they occur flow to either Pindaíba or Araguaia rivers, both in Araguaia-Tocantins River Basin, a major tributary of the Amazon River. The headwaters of Pindaíba and Araguaia rivers are located in the Cerrado domain, forming a complex landscape. In this scenery, small streams flow from their headwaters at the top of low mountain ranges (less than 1000 m a.s.l.) to floodplains, where marginal lagoons and flooded forests along rivers cross the prevailing savannah vegetation. Males of the new species were found calling from vegetation or rocks at the margins of small streams (Fig. 6A B). Vegetation varied from open (scarce bushes) to dense (gallery forest), and, in one occasion this species was found calling from young buriti palm tree (Mauritia flexuosa) in a Vereda, a riparian Cerrado vegetation type characterized by a line of buriti palm trees along a flooded headwater, a swamp or a seasonally flooded area (Figs. 5C D). No male was found calling perched above 1.5 m high. All individuals were found above 500 m a.s.l., near the headwaters of permanent or temporary streams, although several areas below 400 m were also sampled during the same fieldwork season. Aquatic environments below 400 m comprised ponds, wide rivers and flooded forests/grasslands. The other anuran species found in the same localities where Scinax haddadorum sp. nov. has been collected are Ameerega berohoka (Dendrobatidae), Leptodactylus furnarius, Leptodactylus syphax, Pseudopaludicola ameghini (Leptodactylidae), and Pristimantis ventrigranulosus (Craugastoridae). Etymology. The specific epithet honors our colleague and friend Dr. Célio Fernando Baptista Haddad, in recognition to his remarkable contributions to the study of the Brazilian anuran fauna, and his family, including his wife, Patricia L. Morelatto, their daughter, Alice Haddad, and son, Andre Haddad, who have been kind hosts to many herpetologists visiting Rio Claro, São Paulo. 268 Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

9 FIGURE 5. Known geographic distribution of Scinax haddadorum sp. nov. The red triangle indicates the type locality (Fazenda Água Limpa, 15 41'34'' S, 52 20'59'' W). 1: 17 km ESE BR-158 (15 41'29'' S, 52 24'48'' W). 2: Parque Estadual Serra Azul (Protected area, 15 51'28'' S, 52 15'16'' W). 3: Serra do Roncador 6 km E BR-158 (15 20'37'' S, 52 14'05'' W). FIGURE 6. A: Dorsal color pattern of Scinax haddadorum sp. nov. in life, male paratype (MZUSP ). B, C and D: habitats of S. haddadorum sp. nov. Remarks. There are 36 available names in the synonymy of valid species of the Scinax ruber Clade (see Frost 2015). From these, most can be safely discarded as potential names for the new species, for having type material that has been revised relatively recently, and/or having published photos or illustrations. Hyla nigra Cope (1887) deserves special attention, for having its type locality ( Chupada [Chapada dos Guimarães], thirty miles north-east of Cuyabá [currently known as Cuiabá], and near the headwaters of the Xingu, an important tributary of the NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 269

10 Amazon, State of Mato Grosso, Brazil), about 360 Km WNW from the type locality of the new species described here. Hyla nigra has been included in the synonymy of Hyla nasica Cope (1862) [currently, Scinax nasicus] by Duellman (1974), who noticed similarities between the holotype of the former and the two syntypes of the latter, including protruding, subacuminate snouts; subtruncate discs on the fingers, bifid palmar tubercles; outer fingers basally webbed; and webbing vestigial between first and second toes. Photographs of the holotype, ANSP 11269, kindly provided by Mr. Ned Gilmore, indicate that the specimen is relatively well preserved, although the toes seem to have suffered from dehydration sometime in the past (Fig. 7). The specimen differs from both Scinax haddadorum sp. nov. and S. nasicus in that the finger discs are rounded rather than wider than long (those on toes are deformed due to past dehydration), as is common in most species of Scinax (Duellman & Wiens 1992). Furthermore, its hindlimbs are clearly longer than in S. nasicus. Its pattern, already described by Cope (1887) as superior surfaces uniform black but better described today as uniform dark brown, also differentiates the holotype of Hyla nigra from both Scinax haddadorum sp. nov. and S. nasicus. FIGURE 7. Hyla nigra, holotype (ANSP 11269). A: Dorsal view. B: Ventral view. C: Right hand in dorsal view. D: Right foot in ventral view. Scale bars = 10 mm (upper) and 2 mm (lower). Photos courtesy of Ned Gilmore. 270 Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

11 While our observations discard Hyla nigra as an available name for Scinax haddadorum sp. nov., they suggest that Hyla nigra should not be considered a junior synonym of Scinax nasicus. The foot webbing that does not reach the first subarticular tubercle of Toe I suggests that indeed Hyla nigra could be associated with Scinax. In the meantime we propose to consider Hyla nigra Cope (1887) as an incerta sedis. We have observed in all males of the type series a white, slightly thickened area in the pectoral region that is apparently glandular. Similar thickened areas occur in some other species of the Scinax ruber Clade (Faivovich & Araujo-Vieira pers. obs.). It is possible that these are similar structures to the pectoral glands that have been described in S. fuscovarius, S. nasicus, and at least some populations from the states of Bahia and Ceará assigned to S. x-signatus as pectoral plaques (Müller & Hellmich 1936; Lutz 1973; Cei 1980). These are quite evident in S. fuscovarius when soil particles became adhered to their secretion product. A thorough study of the histological structure of these glands, and its variation both taxonomic and seasonal is required to understand its biological role, diagnostic value and evolutionary history. Acknowledgments For access to collections and institutional specimen loans, we thank Célio Fernando Baptista Haddad (CFBH), Georges Lenglet (IRSBN), Linda Trueb and William E. Duellman (KU), José P. Pombal, Jr. (MNRJ), Hussam Zaher and Taran Grant (MZUSP), W. Ronald Heyer (USNM), and Jonathan A. Campbell (UTA). We thank Mr. Ned Gilmore (Academy of Natural Sciences of Drexel University) for kindly providing photographs of the holotype of Hyla nigra. For collecting permits, we thank ICMBio ( / ). For their help and company in the field, we thank Renato Recoder, Mauro Teixeira Júnior, Agustín Camacho, and Mara Souza de Albuquerque e Silva. We thank Santiago J. Nenda for the photographs of the type series; and Agustín J. Elías-Costa and Roberta Graboski for the line drawings and the photographs of the holotype, respectively. For logistical support, we thank pequi - Pesquisa e Conservação do Cerrado. We are grateful to Grants #2006/ , # 2007/ , #2012/ , and #2013/ , São Paulo Research Foundation (FAPESP), Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET, PIP ), PICT 1895/ 2011 and 404/2013, and OEA for financial support and fellowships. References Araujo-Vieira, K., Brandão, R.A. & Faria, D.C.C. (2015) A new species of rock-dwelling Scinax Wagler (Anura: Hylidae) from Chapada dos Veadeiros, Central Brazil. Zootaxa, 3915 (1), Barrio-Amorós, C.L. (2004) Amphibians of Venezuela. Systematic list, distribution and references, an update. Revista Ecologica Latino Americana, 9, Bokermann, W.C.A. (1968) Three new Hyla from the Plateau of Maracás, central Bahia, Brazil. Journal of Herpetology, 1, Bokermann, W.C.A. (1969) Notas sobre Hyla aurata Wied, 1824 (Anura, Hylidae). Revista Brasileira de Biologia, 29, Brusquetti, F., Jansen, M., Barrio-Amorós, C.L., Segalla, M. & Haddad, C.F.B. (2014) Taxonomic review of Scinax fuscomarginatus (Lutz, 1925) and related species (Anura; Hylidae). Zoological Journal of the Linnean Society, 171, Caramaschi, U. & Cardoso, M.C.S. (2006) Taxonomic status of Hyla camposseabrai Bokermann, 1968 (Anura: Hylidae). Journal of Herpetology, 40, Cardoso, A.J. & Sazima, I. (1980) Nova espécie de Hyla do sudeste brasileiro (Amphibia, Anura, Hylidae). Revista Brasileira de Biologia, 40, Carvalho-e-Silva, S.P. & Peixoto, O.L. (1991) Duas novas espécies de Ololygon para os Estados do Rio de Janeiro e Espírito Santo (Amphibia, Anura, Hylidae). Revista Brasileira de Biologia, 51, Cei, J.M. (1980) Amphibians of Argentina. Monitore Zoologico Italiano, New Series, Monografia 2, Cope, E.D. (1862) Catalogues of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo and Uraguay Rivers, by Capt. Thos. J. Page, U.S.N.; and of those procured by Lieut. N. Michler, U.S. Top. Eng., Commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 271

12 Cope, E.D. (1887) Synopsis of the Batrachia and Reptilia obtained by H. H. Smith, in the Province of Mato Grosso, Brazil. Proceedings of the American Philosophical Society, 24, De la Riva, I. (1993) A new species of Scinax (Anura, Hylidae) from Argentina and Bolivia. Journal of Herpetology, 27, De la Riva, I. (1990) Una especie nueva de Ololygon procedente de Bolivia. Revista Española de Herpetología, 4, Drummond, L.O., Baêta, D. & Silvério-Pires, M.R. (2007) A new species of Scinax (Anura, Hylidae) of the S. ruber clade from Minas Gerais, Brazil. Zootaxa, 1612, Duellman, W.E. (1970) The Hylid frogs of Middle America. Monographs of the Museum of Natural History, University of Kansas, 1, Duellman, W.E. (1972) A new species of Hyla from Amazonian Ecuador. Copeia, 2, Duellman, W.E. (1974) A reassessment of the taxonomic status of some neotropical hylid frogs. Occasional Papers of the Museum of Natural History, University of Kansas, 27, Duellman, W.E. (1986) Two new species of Ololygon (Anura: Hylidae) from the Venezuelan Guyana. Copeia, 4, Duellman, W.E. & Wiens, J.J. (1992) The status of the hylid frog genus Ololygon and the recognition of Scinax Wagler, Occasional Papers of the Museum of Natural History, University of Kansas, 151, Duellman, W.E. & Wiens, J.J. (1993) Hylid frogs of the genus Scinax Wagler, 1830 in Amazonian Ecuador and Peru. Occasional Papers of the Museum of Natural History, University of Kansas, 153, Faivovich, J. (2002) A cladistic analysis of Scinax (Anura, Hylidae). Cladistics, 18, Faivovich, J., Haddad, C.F.B., Garcia, P.C.A., Frost, D.R., Campbell, J.A. & Wheeler, W.C. (2005) Systematic review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of Natural History, 294, Fouquette, M.J. Jr. & Pyburn, W.F. (1972) A new Colombian treefrog of the Hyla rubra complex. Herpetologica, 28, Frost, D.R. (2015) Amphibian Species of the World, An Online Reference. Version 5.2. American Museum of Natural History, New York, USA Available from: (accessed 20 May 2015) Heyer, W.R., Rand, A.S., Cruz, C.A.G., Peixoto, O.L. & Nelson, C.E. (1990) Frogs of Boracéia. Arquivos de Zoologia, 31, Juncá, F.A., Napoli, M.F., Nunes, I. Mercês, E.A. & Abreu, R.O. (2015) A new species of the Scinax ruber clade (Anura, Hylidae) from the Espinhaço Range, Northeastern Brazil. Herpetologica, 71, Kwet, A. & Di-Bernardo, M. (1999) Pró-Mata. Anfíbios Amphibien - Amphibians. EDIPUCRS, Porto Alegre, 107 pp. Lescure, J. & Marty, C. (2000) Atlas des Amphibiens de Guyane. Collections Patrimoines Naturels, 45, Lutz, B. (1973) Brazilian Species of Hyla. Austin and London, University of Texas Press, Austin and London, XVIII pp. Moravec, J., Tuanama, I.A., Pérez-Peña, P.E. & Lehr, E. (2009) A new species of Scinax (Anura: Hylidae) from the area of Iquitos, Amazonian Peru. South American Journal of Herpetology, 4, Müller, L. & Hellmich, W. (1936) Wissenschaftliche Ergebnisse der Deutschen Gran Chaco-Expedition (Leiter: Professor Dr. Hans Krieg, München). Amphibien und Reptilien. I. Teil: Amphibia, Chelonia, Loricata. Strecker und Schröder, Stuttgart, 120 pp. Myers, C.W. & Duellman, W.E. (1982) A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from Western Panama. American Museum Novitates, 2752, Napoli, M.F. & Caramaschi, U. (1998) Duas novas espécies de Hyla Laurenti, 1768 do Brasil central afins de H. tritaeniata Bokermann, 1965 (Amphibia, Anura, Hylidae). Boletim do Museu Nacional, Nova Série, Zoologia, 391, Nunes, I. & Pombal, J.P. Jr. (2010) A new Scinax Wagler (Amphibia, Anura, Hylidae) from the Atlantic Rain Forest remains of southern State of Bahia, north-eastern Brazil. Amphibia-Reptilia, 31, Nunes, I., Carvalho, R.R. Jr. & Pereira, E.G. (2010) A new species of Scinax Wagler (Anura: Hylidae) from Cerrado of Brazil. Zootaxa, 2514, Nunes, I. & Pombal, J.P. Jr. (2011) A new snouted treefrog of the speciose genus Scinax Wagler (Anura, Hylidae) from northeastern Brazil. Herpetologica, 67, Nunes, I., Kwet, A. & Pombal Jr., J.P. (2012) Taxonomic revision of the Scinax alter species complex (Anura: Hylidae). Copeia, 3, Pombal, J.P. Jr., Bastos, R.P. & Haddad, C.F.B. (1995a) Vocalizações de algumas espécies do gênero Scinax (Anura, Hylidae) do sudeste do Brasil e comentários taxonômicos. Naturalia, 20, Pombal, J.P. Jr., Haddad, C.F.B. & Kasahara, S. (1995b) A new species of Scinax (Anura: Hylidae) from Southeastern Brazil, with comments on the genus. Journal of Herpetology, 29, Pugliese, A., Pombal, J.P. Jr. & Sazima, I. (2004) A new species of Scinax (Anura: Hylidae) from rocky montane fields of the Serra do Cipó, southeastern Brazil. Zootaxa, 688, Zootaxa 4061 (3) 2016 Magnolia Press ARAUJO-VIEIRA ET AL.

13 Pugliese, A., Baêta, D. & Pombal, J.P. Jr. (2009) A new species of Scinax (Anura: Hylidae) from rocky montane fields in southeastern and central Brazil. Zootaxa, 2269, Pyburn, W.F. (1973) A new hylid from the Llanos of Colombia. Journal of Herpetology, 7, Pyburn, W.F. (1993) A new species of dimorphic tree frog, genus Hyla (Amphibia: Anura: Hylidae), from the Vaupe s River of Colombia. Proceedings of the Biological Society of Washington, 106, Rivero, J.A. (1961) Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology, Cambridge, Massachusetts, 126, Sabaj Pérez, M.H. (2014) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 5.0. American Society of Ichthyologists and Herpetologists, Washington, DC. Available from: (accessed 22 September 2014) Savage, J.M. & Heyer, R.W. (1967) Variation and distribution of the tree-frog genus Phyllomedusa in Costa Rica. Beiträge zur Neotropical Fauna, 5, Spix, J.B. (1824) Animalia nova sive species novae Testudinum et Ranarum, quas in itinere per Brasiliam annis MDCCCXVII MDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavariae Regis. Typis Francisci Seraphi Hübschmanni, Monachii, XXXIX + 53pp. Sturaro, M.J. & Peloso, P.L.V. (2014) A new species of Scinax Wagler, 1830 (Anura: Hylidae) from the Middle Amazon River Basin, Brazil. Papéis Avulsos de Zoologia, 54, Valdujo, P.H., Silvano, D.L., Colli, G. & Martins, M. (2012) Anuran species composition and distribution patterns in Brazilian Cerrado, a neotropical hotspot. South Amerinca Journal of Herpetology, 7, Vanzolini, P.E. (1981) Introduction. The scientific and political contexts of the bavarian expedition to Brazil. In: von Spix, J.B. &Wagler, J.G. (Eds.), Herpetology of Brazil. Society for the Study of Amphibians and Reptiles, Lawrence, pp. ix xxix. APPENDIX 1. Specimens examined. Scinax baumgardneri: KU (paratype). S. cabralensis: MNRJ 42884, (paratypes). S. danae: KU (paratypes). S. dolloi: IRSNB (syntypes, 2 specimens). S. exiguus: KU , (paratypes). S. karenanneae: UTA-A (paratypes). S. nasicus: MACN ; ; ; S. lindsayi: UTA-A 4301, 4303 (paratypes). S. rupestris: MZUSP (holotype); MZUSP , , (paratypes); CHUNB ; (paratypes). Scinax similis: USNM 97319, 97324, (paratypes). S. wandae: KU (paratype). NEW SPECIES OF THE SCINAX RUBER CLADE Zootaxa 4061 (3) 2016 Magnolia Press 273