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Araujo-Vieira, Katyuscia, Caramaschi, Ulisses, Novaes-e-Fagundes, Gabriel, Orrico, Victor G.D., Faivovich, Julián (2020): A neotype for Hyla x-signata Spix, 1824 (Amphibia, Anura, Hylidae). Papéis Avulsos de Zoologia 60: 1-30, DOI: 10.11606/1807-0205/2020.60.56

Scinax x���signatus (Spix, 1824) Hyla x���signata Spix, 1824. Hyla affinis Spix, 1824 ��� Considered a synonym of Hyla x���signata by Hoogmoed & Gruber (1983). Sturaro & Peloso (2014) questioned this association based on the description and figure provided by Spix (1824). Our study of photographs of the holotype (ZSM 2945) indicates that the situation is uncertain. Only a study of the taxonomy of amazonian populations associated with Scinax x���signatus would allow to clarify the status of this nomen. Hyla coerulea Spix, 1824 ��� Considered a synonym of Hyla x���signata by Hoogmoed & Gruber (1983). Sturaro & Peloso (2014) questioned this association based on the description and figure provided by Spix (1824). Our study of photographs of the lectotype designated by Hoogmoed & Gruber (1983) (ZSM 2710-0-1) indicates that the situation is uncertain. Only a study of the taxonomy of amazonian populations associated with Scinax x���signatus would allow to clarify the status of this nomen. Hyla rubra Daudin, 1802 (part) ��� Dum��ril & Bibron, 1841. First treatment as a synonym of Hyla rubra Laurenti, 1768 (not Daudin, 1802; see Le��n, 1969; Rivero, 1969; Duellman & Wiens, 1993). Scytopis xsignatus [sic] ��� Cope, 1870. First combination with Scytopis Cope, 1862. Hyla rubra var. x���signata ��� Peters 1872. First treatment as a variety of Hyla rubra Laurenti. Hyla rubra x���signata ��� M��ller, 1927. First treatment as a subspecies of Hyla rubra Laurenti, 1768. Hyla x���signata x���signata ��� Lutz, 1973. First use as nominal subspecies. Ololygon x���signata ��� Fouquette & Delahoussaye, 1977. First combination with Ololygon Fitzinger, 1843. Scinax x���signata ��� Duellman & Wiens, 1992. First combination with Scinax Wagler, 1830. Scinaxx���signatus ��� K��hler & B��hme,1996.Gender change. Neotype CFBH 44688, adult male, campus of the Universidade Estadual de Santa Cruz ��� UESC, Salobrinho, Ilh��us, State of Bahia, Brazil [39��10���24���W, 14��47���52���S; about 30 m above sea level (a.s.l.)], collected 10 April 2018 by G. Novaes-e-Fagundes. urn:lsid:zoobank. org:act: F357D8CC-446B-4F2C-9B53-190AFEEFD533 Referred specimens Fifteen adults (12 males and three females) from eight localities in the State of Bahia, Brazil. CFBH 21071 (male), Povoado Senote, Caetit�� (42��28���48���W, 14��04���55���S); MHNJCH 1014 (male), Floresta Nacional Contendas do Sincor��, Contendas do Sincor�� (41��07���04���W, 13��55���20���S); MZUESC 20683 (male), Condom��nio Parque Universit��rio, Salobrinho, Ilh��us (39��10���43���W, 14��47���44���S); CFBH 44687 (male), Fazenda Lagoa Nova, Irajuba (39��59���57���W, 13��12���19���S); CFBH 18797 (male), Fazendas Santo Onofre and Cana Brava, Marac��s (approx. 40��25���23.58���W, 13��23���29.30���S); MHNJCH 1701 (male), near Fazendas Santo Onofre and Cana Brava, Marac��s (40��26���26.23���W, 13��21���59.10���S); MHNJCH 1698-1700 (males), Assentamento do Cumbe, Marac��s (40��27���38���W, 13��26���40���S); MZUESC 14890, 14893 (males), and 14891 (female), Companhia de Pesquisa de Recursos Minerais ��� CPRM, Morro do Chap��u (41��09���28���W, 11��32���56���S or 41��06���26���W, 11��29���34���S); MZUESC 15894 and 17503 (females), Pra��a Municipal, Potiragu�� (39��57���30���W, 15��37���07���S); and UFMG 4787 (male), Route Sebasti��o Laranjeiras-Candiba, Sebasti��o Laranjeiras (approx. 42��56���21.34���W, 14��33���13.74���S). Diagnosis (based on neotype and referred specimens) Scinax x���signatus is a species of Scinax, as it shares three synapomorphies of this genus: webbing between toes I and II that does not extend beyond the subarticular tubercle of toe I; origin of the m. pectoralis abdominalis through well-defined tendons; and m. pectoralis abdominalis overlapping m. obliquus externus (da Silva, 1998; Faivovich, 2002; Faivovich et al., 2005). A single synapomorphy is known for the S.ruber Clade: tadpoles with the vent tube above the margin of the lower fin (Faivovich, 2002; Faivovich et al., 2005). While tadpoles unequivocally associated to S. x���signatus remain unknown, this species was associated to the S. ruber Clade by having the unique combination of external vocal sac and presence of pectoral fold [internal vocal sac and pectoral fold absent in most species of the S. catharinae Clade; in few species where the vocal sac is external, the pectoral fold is absent (e.g., S. garibaldiae, S. rizibilis); otherwise, in the two cases where the pectoral fold is present, the vocal sac is internal (S.agilis and S.melanodactylus); J.Faivovich & K. Araujo-Vieira, pers. obs.; see also Bokermann, 1964; Cruz & Peixoto, 1982; Faivovich, 2002; Louren��o et al., 2014, 2019]. Scinax x���signatus can be differentiated from all other species of the S. ruber Clade by the combination of the following characters: (1) male SVL 34.5-38.4 mm, n = 13; (2) snout rounded in dorsal view and profile; (3) pointed tubercles on lower jaw absent; (4) vocal sac subgular, weakly bilobate; (5) spicule-shaped papillary epidermal projections on the nuptial pads and pectoral region present in males; (6) pectoral glands present in males; (7) dorsal color pattern with large irregular dark blotches, commonly with dark X-shaped mark composed of one or two pairs of inverted parenthesis-like blotches; (8) hidden surfaces of thighs dark with irregular pale blotches, yellow in living specimens; (9) iris yellowish golden or bronze with a median black streak; (10) physiological chlorosis absent; and (11) advertisement call composed of a single, multipulsed note, 0.11- 0.25 s duration, 6-14 pulses/note, 52-64 pulses/s. Comparisons with other species of Scinax ruber Clade The SVL in males of Scinax x���signatus (34.5-38.4 mm, n = 13) distinguishes it from the larger species S. castroviejoi and S.eurydice (SVL males 44.0-52.0 mm; De la Riva, 1993; Bokermann, 1968), and from the smaller species S. altae, S. auratus, S. cabralensis, S. caldarum, S. cruentomma, S. danae, S. exiguus, S. fuscomarginatus, S. juncae, S. karenanneae, S. lindsayi, S. madeirae, S. maracaya, S. ruberoculatus, S. rupestris, S. staufferi, S. strussmannae, S. tymbamirim, S. villasboasi, and S. wandae (SVL males 15.7-29.0 mm; Lutz, 1968, 1973; Duellman, 1970, 1986; Pyburn & Fouquette, 1971; Cardoso & Sazima, 1980; Duellman & Wiens, 1993; Pyburn, 1992, 1993; Drummond et al., 2007; Nunes & Pombal, 2010, 2011; Nunes et al., 2012; Brusquetti et al., 2014; Araujo-Vieira et al., 2015; Ferr��o et al., 2018a, b). The snout rounded in dorsal view and profile differentiates Scinax x���signatus from S. alter, S. auratus, S. cretatus, S. crospedospilus, S. imbegue, S. juncae, and S. tymbamirim (sub-elliptical with a pointed tip in dorsal view and slightly acute in profile), S.fuscovarius (roundly acute in dorsal view and protruding in profile), S. caldarum, S. curicica, S. duartei, S. maracaya, S. rossaferesae, and S. tigrinus (sub-elliptical or subovoid in dorsal view and slightly acute in profile), S. squalirostris (pointed in dorsal view and acute in profile), and species of the S. rostratus group (elongate pointed in dorsal view and acute with or without a fleshy proboscis in profile). Furthermore, the absence of pointed tubercles on the lower jaw differentiates S. x���signatus from almost all species of the S. rostratus Group; exceptions are S.kennedyi and S.rostratus (e.g., Duellman, 1972 a, 1973; Pyburn, 1973; Lescure & Marty, 2000; Lima et al., 2005; this study). The presence of a weakly bilobate subgular vocal sac in Scinax x���signatus distinguishes it from S.camposseabrai (bilobate subgular vocal sac; see also Caramaschi & Cardoso, 2006: fig. 1) and from the remaining species of the S. ruber Clade with single subgular vocal sac;exceptions are S.acuminatus, S.dolloi, S.funereus, S.fuscovarius, S.hayii, S.karenanneae, S.montivagus, S.onca, S.oreites, S.pachycrus, S.perereca, S. ruberoculatus, and S.tsachila, that have a weakly bilobate subgular vocal sac (e.g., Cei, 1980; Duellman & Wiens, 1993; Pyburn, 1993;Ferr��o et al., 2017, 2018a; this study). The presence of spicule-shaped papillary epidermal projections on the nuptial pad and pectoral region in males differentiates Scinax x���signatus from all other species of the S. ruber Clade, except for S. fuscovarius (see also Luna et al., 2018: fig. 10A, C). The presence of pectoral glands in males differentiates S. x���signatus from most species of the S. ruber Clade, except for S. funereus, S.fuscovarius, S.nasicus, S.onca, and S.similis, and species of the S. uruguayus Group (e.g., M��ller & Hellmich, 1936; Lutz, 1973; Cei, 1980; this study). The dorsal pattern with large irregular dark blotches, commonly with dark X-shaped marks composed of one or two pairs of inverted parenthesis-like blotches, distinguishes Scinax x���signatus from S. altae, S. alter, S. auratus, S. boesemani, S. caldarum, S. cretatus, S. crospedospilus, S. curicica, S. cuspidatus, S. duartei, S. exiguus, S. fuscomarginatus, S. imbegue, S. juncae, S. madeirae, S. oreites, S. pachycrus, S. quinquefasciatus, S. ruber, S. squalirostris, S. staufferi, S. tsachila, S. tymbamirim, and S. villasboasi (variable number of dorsal and/or lateral stripes; e.g., Duellman, 1970; Lutz, 1973; Duellman & Wiens, 1993; Pugliese et al., 2004; Nunes et al., 2012; Brusquetti et al., 2014; Ron et al., 2018; this study), and S. blairi, S. cabralensis, S. chiquitanus, S. danae, S. iquitorum, S. lindsayi, S. maracaya, and S. strussmannae (scattered or homogeneously distributed spots and/or irregular blotches; e.g., Fouquette & Pyburn, 1972; Cardoso & Sazima, 1980; De la Riva, 1990; Drummond et al., 2007; Ferr��o et al., 2018b; this study). The hidden surfaces of thighs dark colored with light irregular pale blotches, yellow in living specimens differentiate Scinax x���signatus from S. altae, S. auratus, S. baumgardneri, S. boesemani, S. cretatus, S. crospedospilus, S. cruentomma, S. cuspidatus, S. danae, S. elaeochroa, S. exiguus, S. fuscomarginatus, S. ictericus, S. iquitorum, S. madeirae, S. manriquei, S. pachycrus, S. ruberoculatus, S. staufferi, S. strussmannae, S. squalirostris, S. tsachila, S. villasboasi, S. wandae, and species of the S. uruguayus Group (hidden surfaces of thighs uniform, light or dark colored; e.g., Rivero, 1961; Duellman, 1970, 1986; Lutz, 1973; De la Riva, 1990; Duellman & Wiens, 1993; Barrio-Amor��s et al., 2004; Nunes & Pombal, 2011; Brusquetti et al., 2014; Ferr��o et al., 2018a, b; Ron et al., 2018; Baldo et al., 2019; this study), S. funereus, S. onca, and S. iquitorum (hidden surfaces of thighs with horizontal or irregular dark blotches; Duellman, 1971; Ferr��o et al., 2017; Moravec et al., 2009; this study), and from species of the S. rostratus Group (hidden surfaces of thighs uniform light or marked with bold dark and light mottling or broad vertical bars; Duellman, 1972 a, 1973; Henle, 1991; Lescure & Marty, 2000; Lima et al., 2005; this study). The yellowish golden or bronze iris, with a median black streak, distinguishes Scinax x���signatus from S. cruentomma (silvery bronze iris, with a median red streak; Duellman et al., 1972b), S. ruberoculatus (bicolored, reddish upper half and grey lower half; Ferr��o et al., 2018a), and species of the S. uruguayus Group (bicolored, golden upper half and dark brown to black lower half; Baldo et al., 2019). The absence of physiological chlorosis in S. x���signatus distinguishes it from S. boesemani, S. caprarius, S. cruentomma, S. cuspidatus, S. elaeochroa, S. funereus, S. ictericus, S. iquitorum, S. karenanneae, S. manriquei, S. onca, S. strussmannae, and S. tsachila (present in these species; Le��n, 1969; Lutz, 1973; Pyburn, 1993; La Marca, 2004;Moravec et al., 2009;Cole et al., 2013; Melo-Sampaio & Souza, 2015; Ferr��o et al., 2017, 2018b; Acosta-Galvis, 2018; Ron et al., 2018;Taboada et al., 2020). The advertisement call composed of a single multipulsed short note (0.11- 0.25 s), with 6-14 pulses/note, and pulse rate of 52-64 pulses/s differentiates Scinax x���signatus from S. castroviejoi and S. eurydice (two or three multipulsed notes; De la Riva, 1993; De la Riva et al., 1994; Pombal et al., 1995a; Magrini et al., 2011; M��ngia et al., 2017), S. alter, S. curicica, and S. perereca (note duration 0.28- 4.5 s and 21-152 pulses/note; Pombal et al., 1995a, b; Pugliese et al., 2004), S. cruentomma, S. fuscomarginatus, and S. strussmannae (17-90 pulses/note and 113-272 pulses/s; De la Riva et al., 1994; Duellman, 1972b; Brusquetti et al., 2014; Carvalho et al., 2015;Ferr��o et al., 2018b), S. exiguus (23-90 pulses/note; Carvalho et al., 2017), S. madeirae (note duration 0.72- 1.16 s and 104-145 pulses/s; Brusquetti et al., 2014), S. staufferi (100-130 pulses/s; Le��n, 1969), S. wandae (note duration 0.44- 0.69 s and 70-108 pulses/note; Pyburn & Fouquette, 1971; Duellman, 1986; Pombal et al., 2011), and from some large species of the S. rostratus Group: S. boulengeri and S. proboscideus (80-230 pulses/s; Le��n, 1969; Duellman, 1972a), S. jolyi (note duration 2.5 s and 180 pulses/note; Lescure & Marty,2000), S.kennedyi (note duration 0.66- 2.9 s; Pyburn, 1973),and S.sugillatus (note duration 0.28- 0.60 s and 110-140 pulses/s; Duellman, 1973). Description of the neotype Head as wide as long, HL 35.1% and HW 33.2% of SVL (Fig. 2). Snout rounded in dorsal view and profile, with a low protuberance on the tip (Fig. 3A, B). Nostrils dorsolateral, elliptical, protruded; IND 39.7% of IOD. Canthus rostralis marked, convex. Loreal region slightly concave. Eyes large, protuberant, ED 94.7% of IOD and 92.3% of END. Pupil horizontal, subelliptical. Tympanum rounded, separated from eye by a distance almost half TD; TD 75.0% of ED. Tympanic annulus rounded, with the posterior upper portion hidden by the supratympanic fold. Supratympanic fold evident, from the posterior upper portion of the tympanum to the insertion of the forearm. Vocal sac subgular, weakly bilobate, externally evident by the loose skin, not occupying space between head and body, and ventrally not reaching the pectoral fold (Fig. 2B). Pectoral fold present, with pre- and postaxillar elements. Vocal slits present, nearly parallel to the mandible, originating laterally to the tongue and running towards the corner of the mouth. Tongue ovoid, free laterally and posteriorly, slightly notched posteriorly. Vomerine teeth in two slightly separated convex series, bearing five (right) and four (left) teeth. Choanae oval. Axillary membrane absent. Upper arm more slender than forearm. A series of small, flat, ulnar tubercles on the forearm. Fingers short and slender, fringed (Fig. 3C). Relative finger length IIMeasurements (mm): SVL 36.7; HL 12.9; HW 12.2; IND 2.5; IOD 3.8; ED 3.6; END 3.9; TD 2.7; FL 14.6; TL 18.0; 3FD 1.6; 4TD 1.6. Coloration in life: The description is based on the freshly euthanized specimen (Fig. 5). Dorsal color dark brown, with two pairs of large, irregular, black blotches on the suprascapular and sacral regions, and scattered, small, round or irregular, light blotches; interocular region with an inverted triangle-shaped, black marking (Fig. 5A). Upper lip light with diffuse brown blotches anteriorly, and a white stripe on the infraorbital region extending to posterior margin of the tympanum. Loreal region brown with small, irregular, black dots; dark brown canthal line. Post-orbital dark brown line from anterior corner of the eyes, upper margin of tympanum, to the middle of the flanks. Flanks light with irregular, dark brown blotches. Dorsal surfaces of discs, fingers, toes, forearms, and tarsus brownish gray with transverse, brown bars; upper arms uniform; shanks and thighs with large dark brown blotches.Toe webbing covered by brown melanophores. Iris grayish bronze with thin black reticulations, thin yellow halo bordering the pupil, and a, Published as part of Araujo-Vieira, Katyuscia, Caramaschi, Ulisses, Novaes-e-Fagundes, Gabriel, Orrico, Victor G. 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E. 1971. A taxonomic review of South American hylid frogs, genus Phrynohyas. Occasional Papers of the Museum of Natural History, University of Kansas, 4: 1 - 21.","Henle, K. 1991. Ololygon pedromedinae sp. nov., ein neuer Knickzehenlaubfrosch (Hylidae) aus Peru. Salamandra, 27 (1 - 2): 76 - 82.","Duellman, W. E. 1972 b. A New Species of Hyla from Amazonian Ecuador. Copeia, 1972: 265 - 271.","La Marca, E. 2004. Descripcion de dos nuevos anfibios del Piedemonte Andino de Venezuela. Herpetotropicos, 1 (1): 1 - 9.","Melo-Sampaio, P. R. & Souza, M. B. 2015. New and noteworthy distributional records of treefrogs (Anura) from southwestern Amazonia. Check List, 11: 1 - 7.","Acosta-Galvis, A. R. 2018. Una nueva rana de huesos verdes del genero Scinax (Anura: Hylidae) asociada a los bosques subandinos de la cuenca del rio Magdalena, Colombia. Biota Colombiana, 19 (Supp. 1): 131 - 159.","Duellman, W. E. 1973. Descriptions of new hylid frogs from Colombia and Ecuador. Herpetologica, 29: 219 - 227.","Vacher, J. - P.; Chave, J.; Ficetola, F.; Sommeria-Klein, G.; Tao, S.; Thebaud, C.; Blanc, M.; Camacho, A.; Cassimiro, J.; Colston, T. J.; Dewynter, M.; Ernst, R.; Gaucher, P.; Gomes, J. O.; Jairam, R.; Kok, P. J. R.; Dias Lima, J.; Martinez, Q.; Marty, C.; Noonan, B. P.; Nunes, P. M. S.; Ouboter P.; Recoder, R.; Rodrigues, M. T.; Snyder, A.; Souza, S. M. & Fouquet, A. 2020. Large-scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism. Journal of Biogeography, 2020, 00: 1 - 11."]}

Although studies of anuran hand and foot musculature began in the first half of the 19th century, all studies to date have been taxonomically or anatomically restricted in scope, and none has considered the diversity of autopodial myology in Anura as a whole. As a model for future comparisons, we thoroughly describe the hand and foot musculature of an arboreal species (the hylid Triprion petasatus), define the layers in which these muscles are arranged, and attribute presumed functions. On the basis of our myological analysis of 155 species representing 46 of the 54 currently recognized families and main clades of anurans, we describe 20 characters related to hand and foot muscles. Optimization of these characters on the most recent and inclusive phylogenetic hypothesis of Anura results in synapomorphies for several major clades (Bombinatoridae, Alytidae, Xenoanura + Acosmanura, Xenoanura, Pipidae, Acosmanura, Anomocoela, Scaphiopodidae, Pelodytidae + Pelobatidae + Megophryidae, Megophryidae, Neobatrachia, Heleophrynidae, Sooglossidae, Laurentobatrachia, Calyptocephalellidae, Myobatrachoidea, and Nobleobatrachia), including new, nonhomoplastic synapomorphies for clades previously supported only by molecular evidence and a few conflicting phenotypic characters (e.g., Acosmanura, Anomocoela, Neobatrachia). Additionally, we (1) address controversies regarding the homology of anuran and caudate muscles in the context of putative synapomorphies for Ascaphidae + Leiopelmatidae and its sister clade Lalagobatrachia; (2) evaluate a recently proposed terminology for anuran hand and foot musculature; (3) discuss the identities of several hand and foot muscles with problematic homologies; (4) establish a unified terminology for anuran hand and foot muscles, including a list of synonyms for all names employed in the literature; and (5) propose hypotheses for the origin of several myological novelties (neomorphs). Fil: Blotto Acuña, Boris Leonardo. Universidade Estadual Paulista Julio de Mesquita Filho; Brasil. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Pereyra, Martín Oscar. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Posadas | Universidad Nacional de Misiones. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Posadas; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Grant, Taran. Universidade de Sao Paulo; Brasil Fil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

Fil: Vera Candioti, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico (Tucumán). Unidad Ejecutora Lillo. Instituto de Herpetología; Argentina. Fil: Vera Candioti, María Florencia. Fundación Miguel Lillo. Unidad Ejecutora Lillo. Instituto de Herpetología; Argentina. Fil: Taboada, Carlos Alberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia". Área Zoología de Vertebrados. División Herpetología; Argentina. Fil: Salica, María José. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Investigaciones y Transferencia (Jujuy); Argentina. Fil: Salica, María José. Universidad Nacional de Jujuy. Centro de Investigaciones y Transferencia (Jujuy); Argentina. Fil: Baldo, Juan Diego. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico (Nordeste). Instituto de Biología Subtropical; Argentina. Fil: Baldo, Juan Diego. Universidad Nacional de Misiones. Facultad de Ciencias Exactas, Química y Naturales. Instituto de Biología Subtropical; Argentina. Fil: Baldo, Juan Diego. Universidad Nacional de Misiones. Facultad de Ciencias Exactas, Química y Naturales. Departamento de Genética. Laboratorio de Genética Evolutiva; Argentina. Fil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia". Área Zoología de Vertebrados. División Herpetología; Argentina. Fil: Faivovich, Julián. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Biodiversidad y Biología Experimental; Argentina. Fil: Pontes Baêta Da Costa, Délio. Universidad Estatal Paulista Julio de Mesquita Filho. Instituto de Biociencias. Departamento de Zoología. Laboratorio de Herpetología; Brasil. Fil: Pontes Baêta Da Costa, Délio. Universidad Federal de Rio de Janeiro. Museo Nacional. Departamentos de Vertebrados. Herpetología; Brasil. Entre las estructuras embrionarias de los anuros, las glándulas adhesivas (o de cemento) aparecen posterolaterales al estomodeo y producen una secreción mucosa que adhiere los embriones a las superficies dentro y fuera del huevo. En este trabajo estudiamos la ontogenia de las glándulas adhesivas de cinco especies de Phyllomedusa, representantes de los dos principales clados reconocidos en el género, y de embriones de Agalychnis aspera y Phasmahyla cochranae. Las puestas se colectaron en el campo, los embriones fueron fijados periódicamente para obtener series de desarrollo completas y luego estudiados con lupa estereoscópica, microscopía electrónica de barrido y técnicas histológicas de rutina. Las variaciones estructurales incluyen glándulas ausentes (en P. cochranae y Phyllomedusa boliviana), glándulas cónicas funcionales (Tipo morfogenético C, en Phyllomedusa sauvagii, Phyllomedusa iheringii y Phyllomedusa tetraploidea), y un patrón similar al Tipo C inusual en P. azurea, caracterizado por ser glándulas grandes, oblongas, con una disposición en herradura en torno al disco oral. Este último patrón es comparable al observado en A. aspera. Las variaciones interespecíficas también conciernen al arreglo y patrón de regresión de la región secretora, a su vez diferentes del de glándulas Tipo C de otros clados. Aún se requiere información sobre la ocurrencia y desarrollo de las glándulas en géneros basales de Phyllomedusinae (Phrynomedusa y Cruziohyla), y en los taxones basales de los dos clados principales de Phyllomedusa, a fin de interpretar el origen y evolución de los patrones morfogenéticos de las glándulas adhesivas en el grupo. Among anuran embryonic structures, the adhesive (cement) glands appear posterolaterally to the stomodeum and produce a mucous secretion that adheres embryos to surfaces in and out of the egg. In this paper, we study the ontogeny of the adhesive glands in five species of Phyllomedusa representing the two main clades recognized in the genus, plus embryos of Agalychnis aspera and Phasmahyla cochranae. Clutches were collected in the field, and embryos were periodically fixed to obtain complete developmental series and then studied with a stereomicroscope, scanning electron microscopy and routine histological techniques. Structural variations include glands absent (in P. cochranae and Phyllomedusa boliviana), functional club-shaped glands (morphogenetic Type C in Phyllomedusa sauvagii, Phyllomedusa iheringii, and Phyllomedusa tetraploidea), and an unusual Type C-like pattern in Phyllomedusa azurea, characterized by large, oblong glands in a horseshoe-like disposition around the oral disc. This latter gland configuration is similar to that of A. aspera. Interspecific variations also include the arrangement and regression pattern of the secretory region, which are in turn different from those of Type C glands in other clades. To interpret the origin and evolution of gland developmental patterns in the group, we still need information on gland occurrence and development in the basal genera of Phyllomedusinae (Phrynomedusa and Cruziohyla) and in the basal taxa of the two major clades of Phyllomedusa

Osteocephalus helenae: BOLIVIA: Santa Cruz: Buena Vista: MZUSP 95529. BRAZIL: Acre: Catuaba (S 10 ��04'00''; W 63 �� 37 '00''): SMNS 14190 -1, 17502, 17520. Amap��: Rio Marac��, boca Igarap�� Camaip��: MZUSP 99990-99999; Seringal Boa Fortuna: MZUSP 74253. Amazonas: Reserva Adolpho Ducke: MZUSP 59516, 74253, 75648, 84581; Reserva INPA- WWF (Rio Preto): MZUSP 58055. Rond��nia: Cachoeira de Nazar��, Rio Machado: MZUSP 63805. UHE Samuel: MZUSP 76921. GUYANA: Cuyuni-Mazaruni: Kartabo: AMNH H- 13492. Demerara-Mahaica: Marudi Creek: AMNH A- 46233. Essequibo: Shudikar-wau: AMNH A- 49252. Essequibo Islands-West Demerara: Dunoon: UMMZ 52681 (holotype). Potaro-Siparuni: Iwokrama Forest: MTD 48360 - 2; Magdalen's Creek, NW bank of Konawaruk River, ca. 40 km WSW Mabura Hill: AMNH A- 166220 -4, 166229. VENEZUELA: Amazonas: SW sector of Cerro Yapacana: AMNH-A 100598., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

A new species of treefrog of the genus Osteocephalus is described from the Rio Abacaxis, a southern tributary of the Amazonas in the state of Amazonas, Brazil. This member of the O. buckleyi group is characterized by green dorsal colouration with irregular blotches of various shades of brown, light venter with tan spots and bold dark markings on the posterior surfaces of the thighs. It can be distinguished from its closest relative, O. helenae from the same general area, by the lack of an axillary membrane, a few indistinct tubercles on the proximal segment of Finger IV and single ulnar tubercles. Fil: Jungfer, Karl Heinz. Universitat Koblenz. Landau Institut Fur Biologie; Alemania Fil: Verdade, Vanessa K.. Universidade Federal do ABC; Brasil Fil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales ; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Biodiversidad y Biología Experimental; Argentina Fil: Rodrigues, Miguel Trefaut. Universidade de Sao Paulo; Brasil

Osteocephalus verruciger: COLOMBIA: Caquet��: Municipio de Florencia: Escuela Tarqui, carretera Altamira-Florencia km 48-49: ICN 23648. Municipio de Florencia: 35.2 km. arriba de Florencia: ICN 23943. Municipio de Florencia: Vereda Tarqui 38.8 -39.0 km: ICN 23944 - 5, ICN 23948, ICN 23952, ICN 23954. Municipio de Florencia: Vereda Tarqui 13.2 km arriba de Florencia: ICN 23946. Municipio de Florencia: 39.3 km arriba de Florencia: ICN 23947. Huila: Acevedo, R��o Suaza, R��o Aguas Claras near San Adolfo, 1400 m: FMNH 69709 - 10. Palestina: ICN 0 1542. Putumayo: 10.3 km W El Pepino, 1440 m: KU 169586 -7, 169589-93, 169595-6,169599, 169601 -4, 169606- 7. ECUADOR: No specific locality: ZMH-A 946 (syntype). Cotopaxi: Las Pampas (in error): MHNG 2259.20, 2560.62, 2560.64 - 68. Napo: El Reventador: MHNG 2259.18, 2273.28, 2485.65 - 70, 2560.63. 2 km SSW R��o Reventador, 1700 m: KU 164408, 164414, 164416 -9, 164421, 164423 -4, 164426. 3.2 km NNE Oritoyacu, 1910 m: KU 178839 - 44. R��o Azuela, 1740 m: KU 143210 -2, 143215- 7, 143219-24, 164434, SMNS 14197. R��o Azuela, 9.5 km W of Reventador, 1630 m: KU 217750 - 1. R��o Salado, 1 km upstream from R��o Coca, 1420 m: KU 164437, 164442, 178844, 178846- 7. 0.7 km NE R��o Salado bridge on Lago Agrio road, 1380 m: KU 190054. San Rafael: MHNG 2259.19, 2272.98 - 99. S slope Cordillera del Due, 1150 m: KU 123181, 123186. 11.1 km NE Santa Rosa, 1900 m: KU 19004953. 16.5 km NNE Santa Rosa, 1700 m: KU 143209. Pastaza: 9.5 km NW Mera, 1270 m: KU 178848. Tungurahua: 11 km E R��o Negro, 1170 m: KU 146469 - 70., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

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 29.4–35.4 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. Fil: de Araujo Vieira, Katyuscia. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Valdujo, Paula Hanna. World Wildlife Fund. Laboratório de Ecologia da Paisagem; Brasil Fil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Biodiversidad y Biología Experimental; Argentina

Osteocephalus camufatus sp. nov. (Figs. 1–4) Osteocephalus buckleyi [Ca 1 _ MTR 12779 _ 2748]— Jungfer et al. 2013 Holotype: MZUSP 142389 (field number MTR 12779), an adult male of 39.5 mm SVL with well-developed nuptial pads from Brazil: Amazonas: Igarapé-açu (04° 20 ' 40 ''S 58 ° 38 '06''W), right bank of Rio Abacaxis, 30 m asl., collected by Miguel T. Rodrigues, Sergio Marques de Souza, José Cassimiro and José Mário Guellere, on 9 January 2007 (Figs. 1–4). Paratype: MZUSP 157020 (field number MTR 13147), an adult male with well-developed nuptial pads from Brazil: Amazonas: Areal, left bank of Rio Abacaxis (04° 35 ' 49 ''S 58 ° 13 ' 14 ''W), 39 m asl., collected by Miguel T. Rodrigues, Sergio Marques de Souza, José Cassimiro and José Mário Guellere, on 20 January 2007. Diagnosis and comparisons. Osteocephalus camufatus may be diagnosed as (1) a small to medium sized species (as defined by Jungfer 2010: 29) in males (females unknown); (2) skin on dorsum of males shagreen with a few irregular small tubercles, only some bearing keratinized tips; (3) skin on flanks coarsely areolate between limb insertions; (4) rounded, tuberculate canthus rostralis curved inward; (5) frontoparietal ridges not visible externally; (6) dentigerous processes of vomers angular; (7) thick tuberculate supratympanic fold from the posterior edge of the orbit sloping in an arch towards the arm insertion, not reaching below tympanum posteroventrally; (8) webbing on inner edge of third finger extending slightly beyond penultimate subarticular tubercle; (9) distal subarticular tubercle on Finger IV bifid; (10) dorsum boldly blotched irregularly; (11) throat, chest and venter creamy tan to creamy white with numerous irregular tan spots; (12) supralabial area with irregular light and dark markings; (13) flanks light with irregular dark markings; (14) vocal sacs paired, protruding ventral to angles of jaws; (15) juvenile colouration unknown; (16) tadpole habitat (most likely in pools along streams) and labial tooth row formula unknown; (17) colour of tibiofibular bones white in preservative. O. camufatus is a member of the O. buckleyi species group as revealed by molecular data. A phylogenetic analysis using both maximum parsimony (MP) and likelihood (ML) inference resulted in trees that showed that O. camufatus was basal to a large and well-supported clade containing frogs from large areas of Amazonia and Guiana. The oldest available name for this clade is O. helenae (Jungfer et al. 2013). Uncorrected p distances of the mitochondrial 16 S gene between O. camufatus and O. helenae from various localities were 1.3–2.6, while genetic divergences within O. helenae, whose subclade relationships are not yet sufficiently resolved, were 0.0– 2.2 (Jungfer et al. 2013: Appendix S 4 d). The new species shares with several other species of the group the following character states: 1) greenish ground colour, 2) strongly tuberculate canthus rostralis and upper eyelids and 3) posterior surfaces of thighs light with large dark brown spots in preservative. These three characters distinguish it from all frogs in the O. alboguttatus, O. leprieurii, O. planiceps and O. taurinus groups (Jungfer et al. 2013) in which the ground colours are shades of tan or brown, the canthus rostralis is smooth to granulate and the posterior thigh surfaces are uniform. From other species in the O. buckleyi group it differs as follows (characters of O. camufatus in parentheses): Six upland species from the eastern Andean slopes between 600 and 2300 m differ as follows: in Osteocephalus carri, a predominantly brown species (bold blotches of green and brown), the canthus rostralis and upper eyelids are smooth (strongly tuberculate) and the iris is black with golden spots in life (light); O. duellmani is a tan species with or without dark brown blotches (bold blotches of green and brown), with the posterior surfaces of the thighs uniform tan (marbled) and the canthus and upper eyelid bearing a few low tubercles (strongly tuberculate); O. festae exhibits brown ground colour (bold blotches of green and brown), a dark brown iris in life (light, Fig. 4) and uniform tan posterior thigh surfaces (marbled). Males of Osteocephalus mimeticus, O. mutabor (the latter also occurring as low as 150 m asl.) and O. verruciger have a brown ground color lacking green (blotches in shades of green and brown), uniform posterior thigh surfaces (marbled dark brown), and heavily tuberculate dorsa with numerous spinous tubercles bearing keratinized tips during breeding (a few irregular small tubercles, few of them with keratinization). Species of the O. buckleyi group from the lowlands (O. camufatus, differ as follows: Osteocephalus buckleyi has low tubercles or is smooth on the upper eyelid (strongly tuberculate), and has low or lacks tarsal tubercles (prominent). Breeding males of O. buckleyi exhibit dorsal tubercles, many with keratinized tips (few keratinized tips). Osteocephalus cabrerai has a row of tubercles on the lower jaw (absent) and an irregular, deep fringe on the outer edge of Finger IV (low tubercles on proximal segment of Finger IV). Osteocephalus cannatellai from lowlands, but also up to 1290 m asl. (Ron et al. 2012), appears to be larger (38.5–57.2 mm SVL in males, mean, 46.8 mm, Ron et al. 2012) (to 40.4 mm, but note that the sample size is n= 2), and differs by a longer supratympanic fold that reaches the arm insertion (not reaching lower level of tympanum); axillary membrane present (absent); usually a dark venter varying from light grey to brown with or without dark markings (Ron et al. 2012) (yellowish white with small irregular brown spots). Osteocephalus helenae (Ruthven) is a variable frog throughout its range. Specimens from localities closest to the two sites known for O. camufatus, from about 230 km northwest of the type locality, near Manaus and other sites just north and south of the Amazonas in Amazonas, Brazil, exhibit an axillary membrane (absent), a scalloped fringe on the outer edge of Finger IV continued to the ulna (low, indistinct tubercles on proximal segment of Finger IV and single tubercles on ulna), and more webbing on the hand, reaching beyond the distal subarticular tubercle on Finger IV (to distal subarticular tubercle, Fig. 4 a). Specimens from near Rio Branco, Acre, Brazil, roughly 1100 km WSW of the type locality of O. camufatus, have posterior thigh surfaces brown, finely mottled with cream (bold dark brown markings on light ground) and lack tubercles on the eyelids (present). Frogs in the closely related genus Dryaderces are similar in general appearance to some frogs of the O. buckleyi group, but can be distinguished as follows: They differ in exhibiting bold dark ventral reticulation (dark spotting) and uniform dark iris (light iris) in D. pearsoni (Gaige), a longer snout of EN/HL 0.27–0.33 (shorter, EN/ HL 0.25), non-tuberculate canthus rostralis (tuberculate) and light, uniform venter (venter with small dark spots) in Dryaderces sp. (Dryaderces pearsoni [Ca 1 _ MTR 13158 _ 2768] of Jungfer et al. 2013) (KHJ, unpublished data). The inspection of a fresh specimen of “ Osteocephalus ” inframaculatus (Boulenger), a species until recently only known from the holotype and tentatively placed with the O. buckleyi group by Jungfer (2010), was placed in Dryaderces by Hoogmoed (2013). It is easily distinguishable from O. camufatus by its bold dark markings on throat and chest and has a longer snout with EN/HL 0.33 (0.25) and larger tympanum with TD/HL 0.26 (0.22– 0.23). Description of holotype. Snout short (EN/HL 0.245) and blunt, head as long as wide. Nostrils raised, opening laterally. Snout truncate in dorsal and lateral outline. Canthus rostralis well-developed, rounded, describing an inwardly curved line, bearing large tubercles. Skin of head coarsely shagreen, orbits with a few additional small tubercles, some bearing small keratinized tips. A few more scattered keratinized tips on some tubercles of the head and anterior part of the body. No frontoparietal crests evident externally. Loreal region strongly concave bearing low tubercles. Lips flared below the eye. Choanae oblique, elliptical, though slightly angular. Dentigerous processes of vomers short, considerably shorter than choanae width, angular, not in contact with each other, bearing 4 teeth on the left and 3 on the right. The anterior edges of the dentigerous processes are in line with the posterior third of the choanae, their posterior edges well behind the posterior margins of the choanae. Tongue elliptical, slightly wider (by 6 %) than long. The vocal sacs are paired, subgular and protrude ventral to the jaw articulation. A thick glandular supratympanic fold from the posterior part of the orbit medially, covering tympanic annulus dorsally from “ 11 ” (when tympanum is thought to be a clock face) to “ 3 ”, then sloping in a curve towards the arm insertion, reaching no further than the lower one third level of the tympanum. The latter is conspicuous, rounded (barely wider than high), with about 66 % the diameter of the eye. Skin on dorsum coarsely shagreen with a few low, irregularly spaced tubercles. Transversal furrows on the shoulders are absent. Skin smooth between eye and tympanum and areolate posterior to the tympanum above the arm insertion, very coarsely areolate laterally between the limb insertions. Throat and belly areolate. Cloacal opening situated at about half the level of thighs, surrounded by large, irregular warts (except on the cloacal flap, which is smooth). Skin on arms shagreen with a few faint tubercles dorsally, hidden surfaces finely shagreen. A row of low ulnar tubercles also encompasses the proximal segment of Finger IV. Axillary membrane absent. Dorsal surfaces of hind legs shagreen, posterior surfaces of thighs and ventral parts of shank smooth, proximal three fourths of thigh coarsely areolate ventrally. A row of low outer tarsal tubercles from the heel to the proximal subarticular tubercle of Toe V. On the heel, three and four low tubercles, respectively, directed dorsally and laterally. Finger and toe discs elliptical and wider than long. Diameter of disc on Finger III is 74 % of tympanum diameter. Relative finger length is I I basal II 1 ½— 2 ¾ III 2 ⅓— 2 IV. On the foot a large elliptical inner metatarsal tubercle and a small rounded plantar tubercle. A larger rounded conical outer metatarsal tubercle in line with the row of tarsal tubercles. The latter form a shallow fringe on the distal part of the proximal toe segment that is continued to the toe disc. A few indistinct supernumerary tubercles on the proximal segments of Toes III–V. Subarticular tubercles single and conical. Relative lengths of appressed toes I I 1—2 - II 1—2 III 1 — 1 + IV 1 +— 1 V. Measurements and proportions. SVL 39.5; HL 14.3; HW 14.3; TL 22.2; FL 26.8; ED 4.7; TD 3.1; FD 2.3; EN 3.5; IN 3.4; TE 2.0. HL/SVL 0.36; HW/SVL 0.36; TL/SVL 0.56; FL/SVL 0.68; TD/ED 0.66; TD/FD 1.35; EN/ HL 0.25; EN/SVL 0.09; HL/HW 1.00; TE/TD 0.65; TD/HL 0.22. Colour in preservative. The ground colour of head and dorsum is light (green in life) with numerous, more or less irregular blotches in tan and dark brown (various shades of brown in life). This pattern extends to the lateral sides of the head and body. One ill-defined dark brown mark somewhat more conspicuous than others has two arms between the eyes (including the orbits), two between the tympana and on midbody the posterior two arms diverging like the lower part of an X and are continued on the flanks. Two additional ill-defined dark brown marks posteriorly to the proximal part of the sacrum, continued to the flanks. Throat and belly with small irregular brown spots. Although there are light areas on the upper lip, there are no discrete labial marks distinguishable from the rest of the pattern. Tympanum tan. Arms dorsally and laterally with irregular dark brown crossbars, with irregular brown spotting ventrally. Legs dorsally with irregular dark brown crossbars, posterior surfaces of thigh and calf marbled dark brown. Ventral surfaces of thigh tan with lighter areolae, of femur tan peppered with darker markings. Webbing tan, slightly marbled. Subcloacal warts are in creamy white and different shades of tan. Colour in life. Colours were taken from photographs (Fig. 4). Dorsal surfaces covered by bold blotches of different shades of green and brown. A light mark posteroventral to the eye is light green, the posterior part of the upper lip dark brown, of the lower lip white. The tympanum is tan. The iris is light tan in its upper half and creamy white in its lower one, the latter bearing a diffuse dark brown median vertical streak. Upper and lower halves are separated by a weakly demarcated reddish brown horizontal streak and are both finely reticulated black. Some light blue coloration can be seen on the hidden surface of the thigh. Outer tarsal and heel as well as ulnar tubercles are tipped white. Variation in the paratype. An adult male slightly larger than the holotype (40.4 mm SVL) with welldeveloped nuptial pads. The snout is bluntly rounded in dorsal aspect. The dentigerous processes bear five teeth on each side. Keratinized tips on tubercles are lacking entirely. Coloration is similar, but the large tan dorsal mark is less conspicuously outlined. The belly is spotted evenly. There is slightly more webbing on the hand with I basal II 1 2 / 3 — 2 2 / 3 III 2 — 2 IV and little variation on the foot: I 1 +— 2 II 1—2 - III 1 — 1 1 / 3 IV 1 1 / 3 — 1 V. There is also little variation in proportions. Measurements and proportions. SVL 40.4; HL 15.0; HW 14.9; TL 21.2; FL 25.9; ED 5.0; TD 3.4; FD 2.1; EN 3.7; IN 3.3; TE 1.9. HL/SVL 0.37; HW/SVL 0.37; TL/SVL 0.53; FL/SVL 0.64; TD/ED 0.68; TD/FD 1.62; EN/ HL 0.25; EN/SVL 0.09; HL/HW 1.01; TE/TD 0.56; TD/HL 0.23. Distribution and habitat. The new species is so far only known from two sites about 54 km apart along the terra firme forests of the right bank of the Rio Abacaxis, a black water river south of the Amazonas that discharges into Paraná do Arariá, a white water channel delimiting the southern border of Ilha de Tupinambarana (Fig. 5). This is a huge island (about 300 km long) extending between Nova Olinda do Norte on the lower Rio Madeira to Parintins on the Amazonas. Habitat at Igarapé-açu consisted of typical terra firme forest with higher trees reaching up to 40 m and a diameter at breast height of around 80–100 cm. The understory was particularly rich in low palm trees and the leaf litter was dense. The specimen was obtained close to a small stream perched on a tree about 1 m high. At Areal, a site further upstream (also referred to as Paca or Pacamiri locally) the local habitat consisted of an extensive white sand area (campinarana) with scattered vegetation near the margin of the river with abundant clumps of ground bromeliads, gradually replaced by denser areas covered by palm trees and then terra firme forest. The terra firme forest with large trees became more evident farther from the river margin, where the second specimen was obtained. Etymology. The species name camufatus is the Latinized past participle of Italian camuffare, to disguise or mask, from which the word camouflaged is derived, in allusion to the shape-dissolving pattern of greens and browns of the new species.

Osteocephalus camufatus: BRAZIL: Amazonas: Igarap��-a��u, right bank of Rio Abacaxis, 30 m (04�� 20 ' 40 ''S; 58 �� 38 '06''W): MZUSP 142389. Areal, left bank of Rio Abacaxis, 39 m (04�� 35 ' 49 ''S; 58 �� 13 ' 14 ''W). MZUSP 157020., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 180, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624