Search

Araneus coccinella Pocock, 1898 is a South African endemic described from a female collected in Estcourt in KwaZulu-Natal, South Africa. Prior to this paper, the species was only known from the type specimen. It is a rare spider and is now known from a few localities in Gauteng, KwaZulu-Natal and Limpopo in South Africa. The aim of this paper is to add notes on the morphology of A. coccinella based on images of living spiders, and provide information on its behaviour and distribution.

Pseudopaludicola coracoralinae sp. nov. urn:lsid:zoobank.org:act: FAB2ABCB-37D5-429C-9628-91BB62B185B6 Figs 4���5; Tables 3���4 Pseudopaludicola facureae from Palmeiras de Goi��s, GO ��� Carvalho et al. 2015a: 267, 271, table 4, appendix 1���2. Diagnosis Pseudopaludicola coracoralinae sp. nov. is assigned to Pseudopaludicola by having a hypertrophied antebrachial tubercle (see Lynch 1989; Lobo 1995) and by its phylogenetic position within the genus. The new species is characterized by the following combination of characters: (1) upper eyelids smooth, without enlarged palpebral tubercles; (2) heel smooth, without conical tubercle; (3) single, subgular vocal sac, cream-colored with white or off-white warts; (4) terminal phalanges knobbed, without T-shaped terminal phalanges or expanded toe tips; (5) relative short hind limbs (tibio-tarsal articulation just reaching the corner of the mouth); (6) trilled advertisement call pattern, composed of 2���6 welldefined series of tonal notes, having each series of 7���116 notes, emitted at rates of 1485���2077 notes per minute. Etymology The specific name honors Anna Lins dos Guimar��es Peixoto Bretas, better known by her pseudonym Cora Coralina. She was a simple woman, a Brazilian candy maker, writer and poetess. She was born and raised on the banks of the Vermelho River, in the municipality of Goi��s, GO, and lived apart from urban centers. Cora Coralina studied until the third year of elementary school and did a typing course at the age of 70, due to a requirement of the publisher that would publish her first book. She is considered one of the most influential Brazilian writers. Although Cora Coralina wrote her first verses during her adolescence, she had her first book (Poemas dos Becos de Goi��s e Est��rias Mais) published in June 1965, when she was 75 years old. In 1984, the Brazilian Union of Writers awarded her the ���literary personality of the year���. Following that honor, Carlos Drummond de Andrade, another distinguished Brazilian poet, said: ���I admire Cora Coralina and her mastery of living in a state of grace with her poetry. Her verse is like running waters, her lyricism has the power and delicacy of the natural world.��� Type material Holotype BRAZIL ��� adult ♂; state of Goi��s, municipality of Palmeiras de Goi��s; 16��46���59��� S, 49��52���2��� W; 652 m a.s.l. (Fig. 1); 14 Mar. 2019; F.S. Andrade and I.A. Haga leg.; GenBank: MT385245; ZUEC 24704 (Figs 4, 5A). Paratypes BRAZIL ��� 6 adult ♂♂; same data as for holotype; GenBank: MT385243, MT385244; ZUEC 24701 to 24703, 24707 to 24709 ��� 5 adult ♀♀; same data as for holotype; ZUEC 24705, 24706, 24710 to 24712 ��� 4 adult ♂♂; state of Goi��s, municipality of Palmeiras de Goi��s; 16��50���48��� S, 49��51���51��� W; 611 m a.s.l.; 18 Dec. 2013; T.R. de Carvalho and L.B. Martins leg; GenBank: MT385241, MT385242; AAG-UFU 3393 to 3396. Type locality Brazil, GO, municipality of Palmeiras de Goi��s (16��46���59��� S, 49��52���2��� W; 652 m a.s.l.; Fig. 1). Description of the holotype Body elliptic and broad (Fig. 4 A���B; Table 3). Head elliptical, slightly wider than long. Snout subovoid in dorsal view and rounded in profile (Fig. 4 C���D). Eye not protuberant. Eye diameter almost equal to interorbital distance. Interorbital area flat. Pupil rounded. Upper eyelid without tubercles. Nostril not protuberant and closer to snout tip than to eye. Canthus rostralis rounded, smooth. Loreal region slightly concave. Single subgular vocal sac, externally expanded with warty texture. Choanae rounded, well separated from each other. Vocal slits present. Tympanum membrane and annulus absent. Discrete tympanic ridge from behind eye to proximal portion of arm insertion. Mouth opening ventral. Vomerine teeth absent. Tongue ovoid, longer than wide, free posteriorly, without pigmentation at its base. Lateral of head and flanks with discrete granules. One ovoid antebrachial tubercle presents in first quarter of forearm. Finger and toe tips not expanded. Outer and inner metacarpal tubercles welldefined; outer metacarpal tubercle rounded and inner metacarpal tubercle ovoid. Fingers with single and rounded subarticular tubercles. Supernumerary tubercles absent on palm of hand. Thumb with discrete, keratinized, light brown nuptial pad, extending from base of hand to proximal limit of terminal phalanx, covering almost entire external portion of finger. Webbing absent between fingers. Relative finger lengths, when adpressed one to another: IP. coracoralinae sp. nov. were note duration, notes per minute and all spectral traits. The other traits were classified as dynamic. Call quantitative traits and CV values are summarized in Table 4. Differential diagnosis Pseudopaludicola coracoralinae sp. nov. is promptly diagnosed from the P. pusilla species group (sensu Lynch 1989), which includes P. boliviana, P. ceratophyes Rivero & Serna, 1985, P. llanera, P. pusilla and P. motorzinho, by the absence of either T-shaped terminal phalanges or expanded toe tips (discs or pads). The new species has terminal phalanges knobbed, similar in shape to those of P. falcipes (Cardozo & Su��rez 2012: fig. 2B). The new species is also distinguished from P. ceratophyes by having upper eyelids smooth; P. ceratophyes has upper eyelids with an enlarged palpebral tubercle (Lynch 1989). The new species also differs from P. boliviana, P. ceratophyes, P. llanera and P. motorzinho by having a smooth heel, without enlarged, conical tubercle (Lynch 1989; Pansonato et al. 2016). Pseudopaludicola coracoralinae sp. nov. is promptly distinguished from the P. saltica species group that includes P. saltica, P. murundu and P. jaredi, by having short hind limbs (tibiotarsal articulation reaching near the corner of the mouth), whereas all three above-mentioned species have long hind limbs (tibiotarsal articulation extending beyond the tip of snout; Andrade et al. 2016). The color and skin texture of the vocal sac of the P. coracoralinae sp. nov. is whitish cream with white or off-white warts (Fig. 4B), thereby distinguishing it from all congeners, except from P. facureae. Pseudopaludicola ameghini, P. ternetzi, P. falcipes, P. giarettai, P. hyleaustralis Pansonato, Morais, ��vila, Kawashita-Ribeiro, Strussmann & Martins, 2012, P. canga, P. florencei, P. pocoto, P. mineira, P. restinga, P. matuta, P. mystacalis, P. ceratophyes, P. llanera, P. boliviana, P. motorzinho, P. ibisoroca Pansonato, Veiga-Menoncello, Mudrek, Jansen, Recco-Pimentel, Martins & Str ��ssmann, 2016 and P. saltica have vocal sacs that are whitish, yellowish, or light cream with no warty texture (combined characters of the vocal sac of all above-mentioned species: Miranda-Ribeiro 1937; Ruthven 1916; Rivero & Serna 1985; Haddad & Cardoso 1987; Lynch 1989; Lobo 1994; Giaretta & Kokubum 2003; Carvalho 2012; Pansonato et al. 2012, 2013, 2016; Roberto et al. 2013; Magalh���es et al. 2014; Carvalho et al. 2015b, Andrade et al. 2017a, 2018a, 2018b; Cardozo et al. 2018); P. jazmynmcdonaldae has a dark and smooth vocal sac with no warty texture (Andrade et al. 2019); and P. atragula has a white vocal sac with warty texture and dark-colored reticulations (Pansonato et al. 2014a). The trilled pattern of its advertisement call (presence of non-pulsed notes) promptly distinguishes the new species from all species of Pseudopaludicola that have notes with pulsatile structure (pulses separated by silence intervals or not): P. ameghini, P. atragula, P. boliviana, P. falcipes, P. florencei, P. ibisoroca, P. jaredi, P. jazmynmcdonaldae, P. matuta, P. mineira, P. motorzinho, P. murundu, P. mystacalis, P. pocoto, P. restinga, P. saltica and P. ternetzi (Haddad & Cardoso 1987; Dur�� et al. 2004; Pereira & Nascimento 2004; Pansonato et al. 2013, 2014 a, 2014b, 2016; Magalh���es et al. 2014; Andrade et al. 2016, 2017a, 2017b, 2018a, 2018b, 2019; Cardozo et al. 2018). Acoustic comparison with its sister species Pseudopaludicola coracoralinae sp. nov. and P. facureae are indistinguishable in external morphology, but the new species was recovered as a sister species of P. facureae + P. atragula (Fig. 3). Furthermore, the RF multivariate approach applied to morphometric data indicated a broad overlap between the two partitions (Fig. 7 A���B), with a considerable classification error (Table 5). In relation to three species of Pseudopaludicola that share the trilled advertisement call pattern (P. hyleaustralis, P. facureae and P. canga), P. facureae is the one with the most similar call to that of P. coracoralinae sp. nov. The trait of notes per minute distinguishes the new species (1485���2077 notes per minute) from P. canga and P. hyleaustralis (368���1286 notes per minute; combined values, Table 1; see Carvalho et al. 2015a). The RF multivariate analysis on acoustic data indicated a complete segregation between P. coracoralinae sp. nov. and P. facureae, without any classification error (Table 5; Fig. 7C). Notes per minute (P. coracoralinae sp. nov. 1796 �� 123 (1485���2077) vs P. facureae 1383 �� 192 (512���1843)), number of series per call (P. coracoralinae sp. nov. 3 �� 1 (2���6) vs P. facureae 10 �� 6 (4���20)) and number of notes per series (P. coracoralinae sp. nov. 29 �� 16 (7���116) vs P. facureae 17 �� 18 (2���93)) were the main sources of variation in both variable importance measurements (Fig. 7D). In addition to these abovementioned traits, we found differences (P ��0.01) between these two species in note duration, internote interval, series of notes duration, interseries interval and dominant frequency. Phylogenetic inference and genetic distances of the new species Pseudopaludicola coracoralinae sp. nov. was recovered as a sister species of the P. atragula + P. facureae clade (Fig. 3). Uncorrected genetic distance between the P. coracoralinae sp. nov. and P. atragula was 4.5% (mean value), and from P. facureae, it was 4.9% (mean value). The maximum intraspecific distance was 0.4% (Supplementary file 2). No molecular data are available for P. ceratophyes, P. hyleaustralis and P. ibisoroca; however, the new species is easily diagnosed from these species by morphology and acoustics (see further details in Differential diagnosis section). Natural history notes Males of the new species were found calling in a partially flooded open area surrounded by a newly planted cornfield (corn stalk Leptodactylus fuscus (Schneider, 1799) and Physalaemus marmoratus (Reinhardt & L��tken, 1862) at its type locality. Curiously, the congener Pseudopaludicola mystacalis was observed about 50 meters in a similar partially flooded open area surrounded by the same cornfield. We heard and observed only P. mystacalis at this site, not P. coracoralinae sp. nov. Distribution Pseudopaludicola coracoralinae sp. nov. is known only from the type locality (Fig. 1). However, we are aware of other populations that have trilled advertisement calls similar to those of P. coracoralinae sp. nov. and P. facureae. These populations occur in Limeira do Oeste, MG (Andrade & Carvalho 2013); Goian��sia, Piracanjuba and in the Altamiro de Moura Pacheco State Park, all in GO, Brazil (Guimar���es et al. 2001; Carvalho et al. 2015a; Ramalho et al. 2018). Goian��sia is about 180 km northeast from the type locality of P. coracoralinae sp. nov., Piracanjuba is about 100 km southeast and the Altamiro de Moura Pacheco State Park is about 80 km northeast. Limeira do Oeste is closer to the type locality of P. facureae, about 250 km east. However, the specific identities of these populations will only be confirmed when their genetic information is available because they are morphologically and acoustically cryptic., Published as part of Andrade, Felipe Silva de, Haga, Isabelle Aquemi, Lyra, Mariana L��cio, Carvalho, Thiago Ribeiro de, Haddad, C��lio Fernando Baptista, Giaretta, Ariovaldo Antonio & Toledo, Lu��s Felipe, 2020, Reassessment of the taxonomic status of Pseudopaludicola parnaiba (Anura, Leptodactylidae, Leiuperinae), with the description of a new cryptic species from the Brazilian Cerrado, pp. 1-36 in European Journal of Taxonomy 679 on pages 13-20, DOI: 10.5852/ejt.2020.679, http://zenodo.org/record/3934484, {"references":["Carvalho T. R., Teixeira B. F. V., Martins L. B. & Giaretta A. A. 2015 a. Intraspecific variation and new distributional records for Pseudopaludicola species (Anura, Leptodactylidae, Leiuperinae) with trilled advertisement call pattern: diagnostic characters revisited and taxonomic implications. North-Western Journal of Zoology 11: 262 - 273.","Lynch J. D. 1989. A review of leptodactylid frogs of the genus Pseudopaludicola in northern South America. Copeia 3: 577 - 588. https: // doi. org / 10.2307 / 1445483","Lobo F. 1995. Analisis filogenetico del genero Pseudopaludicola (Anura: Leptodactylidae). Cuadernos de Herpetologia 9: 21 - 43.","Rivero J. A. & Serna M. A. 1985. Una nueva Pseudopaludicola (Amphibia: Leptodactylidae) cornuda del sureste de Colombia. Caribbean Journal of Science 20: 169 - 171.","Cardozo D. & Suarez P. 2012. Osteological description of Pseudopaludicola canga with implications for the taxonomic position of this taxon. Zootaxa 3515: 75 - 82. https: // doi. org / 10.11646 / zootaxa. 3515.1.6","Pansonato A., Veiga-Menoncello A. C. P., Mudrek J. R., Jasen M., Recco-Pimentel S. M., Martins I. A. & Str ʾ ssmann C. 2016. Two new species of Pseudopaludicola (Anura: Leptodactylidae: Leiuperinae) from eastern Bolivia and western Brazil. Herpetologica 72: 235 - 255. https: // doi. org / 10.1655 / Herpetologica-D- 14 - 00047.1","Andrade F. S., Magalh ues F. M., Nunes-de-Almeida C. H. L., Veiga-Menoncello A. C. P., Santana D. J., Garda A. A., Loebmann D., Recco-Pimentel S. M., Giaretta A. A. & Toledo L. F. 2016. A new species of long-legged Pseudopaludicola from northeastern Brazil (Anura, Leptodactylidae, Leiuperinae). Salamandra 52: 107 - 124.","Miranda-Ribeiro A. 1937. Alguns batrachios novos das collecies do Museo Nacional. O Campo 8: 66 - 69.","Ruthven A. G. 1916. A new species of Paludicola from Colombia. Occasional Papers of the Museum of Zoology, University of Michigan 30: 1 - 3.","Haddad C. F. B. & Cardoso A. J. 1987. Taxonomia de tres especies de Pseudopaludicola (Anura, Leptodactylidae). Papeis Avulsos de Zoologia 36: 287 - 300.","Lobo F. 1994. Descripcion de una nueva especie de Pseudopaludicola (Anura: Leptodactylidae), redescripcion de P. falcipes (Hensel, 1867) y P. saltica (Cope, 1887). Cuadernos de Herpetologia 8: 177 - 199.","Giaretta A. A. & Kokubum M. N. C. 2003. A new species of Pseudopaludicola (Anura, Leptodactylidae) from northern Brazil. Zootaxa 383: 1 - 8. https: // doi. org / 10.11646 / zootaxa. 383.1.1","Carvalho T. R. 2012. A new species of Pseudopaludicola Miranda-Ribeiro (Leiuperinae: Leptodactylidae: Anura) from the Cerrado of southeastern Brazil with a distinctive advertisement call pattern. Zootaxa 3328: 47 - 54. https: // doi. org / 10.11646 / zootaxa. 3328.1.4","Pansonato A., Morais D. H., Avila R. W., Kawashita-Ribeiro R. A., Str ʾ ssmann C. & Martins I. A. 2012. A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leiuperidae) from the state of Mato","Pansonato A., Str ʾ ssmann C., Mudrek J. R. & Martins I. A. 2013. Morphometric and bioacoustic data on three species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) described from Chapada dos Guimar ues, Mato Grosso, Brazil, with the revalidation of Pseudopaludicola ameghini (Cope, 1887). Zootaxa 3620: 147 - 162. https: // doi. org / 10.11646 / zootaxa. 3620.1.7","Roberto I. J., Cardozo D. & Avila R. W. 2013. A new species of Pseudopaludicola (Anura, Leiuperidae) from western Piaui State, Northeast Brazil. Zootaxa 3636: 348 - 360. https: // doi. org / 10.11646 / zootaxa. 3636.2.6","Magalh ues F. M., Loebmann D., Kokubum M. N. C., Haddad C. F. B. & Garda A. A. 2014. A new species of Pseudopaludicola (Anura: Leptodactylidae: Leiuperinae) from northeastern Brazil. Herpetologica 70: 77 - 88. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00054","Carvalho T. R., Borges-Martins M., Teixeira B. F. V., Godinho L. B. & Giaretta A. A. 2015 b. Intraspecific variation in acoustic traits and body size, and new distributional records for Pseudopaludicola giarettai Carvalho, 2012 (Anura, Leptodactylidae, Leiuperinae): implications for its congeneric diagnosis. Papeis Avulsos de Zoologia 55: 245 - 254. https: // doi. org / 10.1590 / 0031 - 1049.2015.55.17","Andrade F. S., Haga I. A., Bang D. L. & Giaretta A. A. 2017 a. The differential acoustic diagnosis between two Pseudopaludicola sister species (Anura, Leptodactylidae, Leiuperinae). Zootaxa 4319 (2): 391 - 400. https: // doi. org / 10.11646 / zootaxa. 4319.2.12","Andrade F. S., Haga I. A., Lyra M. L., Leite F. S. F., Kwet A., Haddad C. F. B., Toledo L. F. & Giaretta A. A. 2018 a. A new species of Pseudopaludicola Miranda-Ribeiro (Anura: Leptodactylidae: Leiuperinae) from eastern Brazil, with novel data on the advertisement call of Pseudopaludicola falcipes (Hensel). Zootaxa 4433 (1): 71 - 100. https: // doi. org / 10.11646 / zootaxa. 4433.1.4","Andrade F. S., Haga I. A., Lyra M. L., Carvalho T. R., Haddad C. F. B., Giaretta A. A. & Toledo L. F. 2018 b. A new species of Pseudopaludicola (Anura, Leptodactylidae, Leiuperinae) from the state of Minas Gerais, Brazil. European Journal of Taxonomy 480: 1 - 25. https: // doi. org / 10.5852 / ejt. 2018.480","Cardozo D. E., Baldo D., Pupin N., Gasparini J. L. & Haddad C. F. B. 2018. A new species of Pseudopaludicola (Anura, Leiuperinae) from Espirito Santo, Brazil. PeerJ 6: e 4766. https: // doi. org / 10.7717 / peerj. 4766","Andrade F. S., Silva L. A., Koroiva R., Fadel R. M. & Santana D. J. 2019. A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) from an Amazonia-Cerrado transitional zone, state of Tocantins, Brazil. Journal of Herpetology 53 (1): 68 - 80. https: // doi. org / 10.1670 / 18 - 125","Pansonato A., Mudrek J. R., Veiga-Menoncello A. C. P., Rossa-Feres D. C., Martins I. A. & Str ʾ ssmann C. 2014 a. A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) from northwestern state of S uo Paulo, Brazil. Zootaxa 3861: 249 - 264. https: // doi. org / 10.11646 / zootaxa. 3861.3.3","Andrade F. S. & Carvalho T. R. 2013. A new species of Pseudopaludicola Miranda-Ribeiro (Leiuperinae: Leptodactylidae: Anura) from the Cerrado of southeastern Brazil. Zootaxa 3608 (5): 389 - 397. https: // doi. org / 10.11646 / zootaxa. 3608.5.7","Dure M. I., Schaefer E. F., Hamann M. I. & Kehr A. I. 2004. Consideraciones ecologicas sobre la dieta, la reproduccion y el parasitismo de Pseudopaludicola boliviana (Anura, Leptodactylidae) de Corrientes, Argentina. Phyllomedusa 3: 121 - 131. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 3 i 2 p 121 - 131","Pereira E. G. & Nascimento L. B. 2004. Descric uo da vocalizac uo e do girino de Pseudopaludicola mineira Lobo, 1994, com notas sobre a morfologia de adultos (Amphibia, Anura, Leptodactylidae). Arquivos do Museu nacional 62: 233 - 240.","Pansonato A., Mudrek J. R., Simioni F., Martins I. A. & Str ʾ ssmann C. 2014 b. Geographical variation in morphological and bioacoustic traits of Pseudopaludicola mystacalis (Cope, 1887) and a reassessment of the taxonomic status of Pseudopaludicola serrana Toledo, 2010 (Anura: Leptodactylidae: Leiuperinae). Advances in Zoology 2014: 1 - 13. https: // doi. org / 10.1155 / 2014 / 563165","Andrade F. S., Leite F. S. F., Carvalho T. R., Bernardes C. S. & Giaretta A. A. 2017 b. First record of Pseudopaludicola pocoto Magalh ues, Loebmann, Kokubum, Haddad & Garda, 2014 (Anura, Leptodactylidae, Leiuperinae) in Bahia state, northeastern Brazil, with further data on its advertisement call. Check List 13 (1): 2047. https: // doi. org / 10.15560 / 13.1.2047","Guimar ues L. D., Lima L. P., Juliano R. F. & Bastos R. P. 2001. Vocalizacies de especies de anuros (Amphibia) no Brasil Central. Boletim do Museu Nacional, Zoologia (Nova Serie) 474: 1 - 15. https: // doi. org / 10.1590 / S 0073 - 47212003000200005","Ramalho W. P., Franca D. P. F., Guerra V., Marciano R., Vale N. C. & Silva H. L. R. 2018. Herpetofauna of Parque Estadual Altamiro de Moura Pacheco: one of the last remnants of seasonal forest in the core region of the Brazilian Cerrado. Papeis Avulsos de Zoologia 58: e 20185851. http: // doi. org / 10.11606 / 1807 - 0205 / 2018.58.51"]}

Leptodactylus watu sp. nov. (Figs. 2, 4a, 4c, 5h, 6h, 10) [http://zoobank.org/ urn:lsid:zoobank.org:act: 5D79A081-C50C-463A-8101-04351E87AD23] Leptodactylus aff. spixi: Guimar��es et al. (2019) Holotype. UFMG 21332, an adult male from Parque Estadual do Rio Doce (19.753922��S, 42.633047��W, approximately 280 m a.s.l.), in Marli��ria, Minas Gerais State, southeastern Brazil, collected by H. Thomassen on 21 November 2017. Paratypes. Seven adult males (UFMG 21327���21331 and 21337���21338) and five adult females (UFMG 21333��� 21336 and 21339) collected from the type locality by H. Thomassen from November 2017 to February 2018. Additional material. One genetic voucher (UFMG 16995) from Santa B��rbara do Leste, Minas Gerais State (Fig. 1). Etymology. The word watu, a noun in apposition, is derived from the dialect of the Bor��n indigenous people and is a reference to the Doce River (Costa-Reis & Genovez 2013). In 2015, the Doce River experienced the worst environmental disaster that ever took place in Brazil, the collapse of a mining tailings dam owned by Samarco (and co-owned by the Brazilian Vale and Australian BHP Billiton). The catastrophic dam failure released around 60 million cubic meters of iron ore tailings (toxic slurry) directly into the Doce River watershed, killing 20 people and affecting biodiversity across hundreds of kilometers around the river drainage, riparian lands, and the Atlantic coast, in addition to a severe contamination of the soil and water table within the affected region. The epithet is a tribute to the resistance of the Bor��n people and to the Watu (Doce River) in southeastern Brazil. Diagnosis. Leptodactylus watu is characterized by the following combination of character states: (1) small to moderate size, male SVL = 38.5���42.2 mm, female SVL = 40.5���47.9 mm; (2) sole of foot with few, non-obvious tubercles; (3) advertisement call composed of single, pulsed notes; (4) note duration varying from 27���70 ms; (5) note dominant frequency varying from 495���1098 Hz; (6) frequency upsweep varying from 0���560 Hz. Comparisons with members of the L. mystaceus complex. Leptodactylus watu is distinguished from other species belonging to the L. mystaceus complex, except L. barrioi, L. kilombo and L. notoaktites, by having the sole of the foot covered with few, non-obvious tubercles, whereas L. didymus, L. elenae, L. mystaceus, and L. spixi have prominent tubercles (Heyer 1978, 1983; Heyer et al. 1996). The new species can be further distinguished from some species by a smaller size (Table 2): L. didymus (male SVL = 45.9���52.2 mm, mean = 49.3; female SVL = 45.8���53.5 mm, mean = 49.8; Heyer et al. 1996), L. mystaceus (male SVL = 42.4���52.2 mm, mean = 47.4; female SVL = 44.8��� 56.1 mm, mean = 50.0; Heyer et al. 1996), and L. notoaktites (male SVL = 42.5���54.2 mm, mean = 47.5; female SVL = 43.4���55.8 mm, mean = 48.0; de S�� et al. 2014). Members of the L. mystaceus complex are mainly distinguished from each other by their distinct advertisement calls (Fig. 6). Acoustic comparisons in the L. fuscus group. The advertisement call of Leptodactylus watu is composed of single, pulsed notes (Table 3; Fig. 6h), differing from species with trill calls (L. cunicularius, L. cupreus, L. oreomantis, and L. plaumanni; Fig. 7; Carvalho et al. 2013) and species with nonpulsed calls (L. albilabris, L. bufonius, L. camaquara, L. didymus, L. elenae, L. furnarius, L. fuscus, L. gracilis, L. jolyi, L. kilombo sp. nov., L. laticeps, L. latinasus, L. longirostris, L. marambaiae, L. mystacinus, L. notoaktites, L. poecilochilus, L. sertanejo, L. spixi, L. syphax, L. tapiti, and L. troglodytes). From other species with pulsed calls, L. watu is distinguished by having its call formed by partly fused pulses, whereas the calls of L. barrioi, L. mystaceus and L. cf. mystaceus are formed by complete pulses (Fig. 6). From species also having calls with partly fused pulses (L. caatingae, L. fragilis, and L. labrosus), L. watu is distinguished by a shorter note (27���70 ms) from L. fragilis (> 100 ms; Heyer et al. 2006). The calls of L. watu, L. caatingae, and L. labrosus are the most similar to each other in the L. fuscus group, but these species can be distinguished, although value ranges partially overlap, by differences in the dominant frequency (L. watu: 495���1098 Hz, mean = 763; L. caatingae: 1148���1430 Hz, mean = 1315, in our sample; 943���1620 Hz, mean = 1423 Hz, in the original description; Heyer & Junc�� 2003) or duration (L. watu: 27���70 ms, mean = 40; L. labrosus: 64���133 ms, mean = 100; Carvalho & Ron 2011) of their calls. Still, these species differ markedly in morphology and size (see previous paragraph; Heyer 1978; Heyer & Junc�� 2003). Description of holotype. Body robust. Snout sub-elliptical in dorsal and ventral views (Fig. 10 a���b), acuminate in lateral view (Fig. 10e). Canthus rostralis rounded; loreal region flat; tympanic annulus well-defined, circular (TD = 62.4% ED); supratympanic fold from the posterior corner of the eye, passing over the dorsal edge of the tympanic annulus, and ending at the base of the arm; vocal sac subgular; vocal slits present; vomerine teeth in two nearly straight rows, almost connected, medial and posterior to choanae and almost parallel to sagittal plane. Tongue rounded, free at its posterior third. Relative finger lengths II = IV Colors. In life, dorsal surfaces (body and limbs) light brown with dark gray blotches poorly delimited (Figs. 4a, c, 5h). Upper region of the loreal region black, lower region light cream, lips dark brown. Black stripe on the supratympanic fold, delimited dorsally by a white line. Light cream stripe on the dorsolateral fold, more conspicuous posteriorly. Flank light gray. Tympanic membrane dark brown on the center and upper rim, light brown laterally and on ventral rim. Throat immaculate white with nonpigmented portions near the jaw. Ventral surface of arms, legs (except tarsus), and belly nonpigmented; tarsus with scattered dark brown stains. Weakly defined dark brown transversal bars on the upper surface of hindlimbs and forearm. Posterior surface of thigh with a yellow longitudinal stripe bordered by two black stains; upper portion of with black spots and stains. Ventral surface of hand, foot, and tarsus dark brown interspersed with nonpigmented areas. In preservative, dorsal surfaces (body and hindlimbs) light gray with dark gray blotches poorly delimited; forearms light cream with scattered dark gray dots and spots (Fig. 10). Upper portion of the loreal region black, lower half cream-colored, lips blackish gray. Black stripe on the supratympanic fold. White stripe on the dorsolateral fold, more conspicuous posteriorly. Flank light gray. Tympanic membrane light brown with the center and upper edge dark gray. Ventral surfaces cream-colored. Throat with few gray dots and stains near the jaw laterally. Ventral surface of hindlimbs with few dark gray spots. Dark brown transversal bars on the dorsal surface of hindlimbs and forearm. Posterior surface of thigh with a white longitudinal stripe bordered by two black stains. Ventral surface of hand, foot, and tarsus dark brown interspersed with nonpigmented areas. Intraspecific variation. In life, dorsal coloration varies from brown, sometimes conspicuous reddish shades, or dark gray. The dorsal light brown coloration in life becomes light gray in preservative. Dark brown blotches on the dorsum can be present, more conspicuous in preserved specimens. Dark brown transversal bars on the upper surface of thigh and tibia are more evident in preserved species. Lateral line of tubercles is poorly evident in most specimens. Inner metacarpal tubercle is rounded in UFMG 21329. Advertisement call. Description based on 129 calls recorded from two males (Table 3). The call (Fig. 6h) consists of single, pulsed notes given at a rate of 124���275 per minute. Note duration varies from 27���70 ms. Rise time varies from 37���80% of note duration. Notes are formed by 2���7 partly fused pulses emitted at a rate of 57���153 per second. The dominant frequency coincides with the fundamental harmonic at 495���1098 Hz. Notes have no frequency modulation or a modest frequency upsweep throughout their duration, varying from 0���560 Hz. Habitat and natural history. At the type locality, Leptodactylus watu was found calling during the rainy season in the region from late October to late February. Calling is strongly associated with rainy days or few days after heavy pours. Males were found calling around flooded areas or close to moist ground in both open areas (transitional marshlands between the forest and lakes) and inside the forest. All calling males were hidden either under dense vegetation, ground leaf litter or inside underground chambers. They were never found calling in pastures. The congeneric species found calling in syntopy with L. watu were L. fuscus, L. latrans, and L. labyrinthicus. Distribution. Leptodactylus watu is only known from two localities in a region characterized by lowland interior Atlantic forests associated with the upper-middle Doce River drainage, in the Brazilian state of Minas Gerais (Fig. 1): (1) the type locality, at Parque Estadual do Rio Doce, and (2) Santa B��rbara do Leste, approximately 70 km southeast from the type locality. Remarks. The identity of populations assigned to Leptodactylus aff. mystaceus (Santana et al. 2010), L. aff. spixi (Ferreira et al. 2019), and L. spixi (R��dder et al. 2007; Silva-Soares & Scherrer 2013; Pereira-Ribeiro et al. 2019) from different localities in southeastern and northeastern Brazil should be addressed using calls and/or DNA sequences, as these records could correspond to L. barrioi or L. watu, or additional candidate new species., Published as part of Da Silva, Leandro A., Magalh��es, Felipe M., Thomassen, Hans, Leite, Felipe S. F., Garda, Adrian A., Brand��o, Reuber A., Haddad, C��lio F. B., Giaretta, Ariovaldo A. & De Carvalho, Thiago R., 2020, Unraveling the species diversity and relationships in the Leptodactylus mystaceus complex (Anura: Leptodactylidae), with the description of three new Brazilian species, pp. 151-189 in Zootaxa 4779 (2) on pages 167-170, DOI: 10.11646/zootaxa.4779.2.1, http://zenodo.org/record/3833431, {"references":["Guimaraes, C. S., Assis, C. L., Thomassen, H., Leite, F. S. F. & Feio, R. N. (2019) Anfibios do Parque Estadual do Rio Doce, Minas Gerais. Universidade Federal de Vicosa, Departamento de Biologia Animal, Museu de Zoologia Joao Moojen, Vicosa, Minas Gerais, 57 pp.","Costa-Reis, R. & Genovez, P. F. (2013) Territorio sagrado: exilio, diaspora e reconquista Krenak no vale do Rio Doce, Resplendor, MG. Boletim Goiano de Geografia, 33, 1 - 15. https: // doi. org / 10.5216 / bgg. v 33 i 1.23628","Heyer, W. R. (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Science Bulletin, Natural History Museum of Los Angeles County, 29, 1 - 85.","Heyer, W. R. (1983) Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978, and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae). Proceedings of the Biological Society of Washington, 96, 270 - 272.","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. https: // doi. org / 10.1163 / 156853896 X 00252","de Sa, R. O., Grant, T., Camargo, A., Heyer, W. R., Ponssa, M. L. & Stanley, E. (2014) Systematics of the Neotropical genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the relevance of non-molecular evidence, and species accounts. South American Journal of Herpetology, 9, S 1 - S 128. https: // doi. org / 10.2994 / SAJH-D- 13 - 00022.1","Carvalho, T. R., Leite, F. S. F. & Pezzuti, T. L. (2013) A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae, Leptodactylinae) from montane rock fields of the Chapada Diamantina, northeastern Brazil. Zootaxa, 3701 (3), 349 - 364. https: // doi. org / 10.11646 / zootaxa. 3701.3.5","Heyer, W. R., Heyer, M. M. & de Sa, R. O. (2006) Leptodactylus fragilis. Catalogue of American Amphibians and Reptiles, 830, 1 - 26.","Heyer, W. R. & Junca, F. A. (2003) Leptodactylus caatingae, a new species of frog from eastern Brazil (Amphibia: Anura: Leptodactylidae). Proceedings of the Biological Society of Washington, 116, 317 - 329.","Carvalho, T. R. & Ron, S. R. (2011) Advertisement call of Leptodactylus labrosus Jimenez de la Espada, 1875 (Anura, Leptodactylidae): an unusual advertisement call within the L. fuscus group. Herpetology Notes, 4, 325 - 326.","Santana, D. J., Sao Pedro, V. A., Hote, P. S., Roberti, H. M., Sant'Anna, A. C., Figueiredo-de-Andrade, C. A. & Feio, R. N. (2010) Anurans in the region of the High Muriae River, state of Minas Gerais, Brazil. Herpetology Notes, 3, 1 - 10.","Ferreira, R. B., Monico, A. T., Silva, E. T., Lirio, F. C. F., Zocca, C., Mageski, M. M., Tonini, J. F. R., Beard, K. H., Duca, C. & Silva- Soares, T. (2019) Amphibians of Santa Teresa, Brazil: the hotspot further evaluated. ZooKeys, 857, 139 - 162. https: // doi. org / 10.3897 / zookeys. 857.30302","Rodder, D., Teixeira, R. L., Ferreira, R. B., Dantas, R. B., Pertel, W. & Guarniere, G. J. (2007) Anuran hotspots: the municipality of Santa Teresa, Espirito Santo, southeastern Brazil. Salamandra, 43, 91 - 110.","Silva-Soares, T. & Scherrer, P. V. (2013) Amphibians of Parque Estadual do Forno Grande, State of Espirito Santo, Southeastern Brazil: Species composition and conservation. North-Western Journal of Zoology, 9, 113 - 120.","Pereira-Ribeiro, J., Ferreguetti, A. C., Linause, T. M., Cozer, J. S., Bergallo, H. G. & Rocha, C. F. D. (2019) Diversity and distribution of anurans from Mata das Flores State Park, Espirito Santo, southeastern Brazil. Oecologia Australis, 23, 292 - 300. https: // doi. org / 10.4257 / oeco. 2019.2302.08"]}

Adenomera phonotriccus, n. sp. Figures 1���3 REFERRED MATERIAL: Voucher specimens with DNA sequenced by Fouquet et al. (2014) and assigned to Adenomera sp. F (in part). SUGGESTED VERNACULAR NAME: Tody-tyrant-voiced nest-building frog. HOLOTYPE: MPEG 41155 (field number TRC 135), adult male, from the municipality of Palestina do Par�� (5.70228�� S, 48.24949�� W; 166 m), on the western margin of the lower Araguaia River, state of Par��, northern Brazil, collected by P.L. V. Peloso, M.J. Sturaro, P. V. Cerqueira, G. Gon��alves, A.A. Giaretta, P. Marinho, and T. R. de Carvalho on January 9, 2018 (Zoobank LSID registration: urn:lsid:zoobank.org:act: 330AE4AB-1ABB-4B90-9245-1DBC2A1F661B). SOUND RECORDING: Adenomera _phonotriccusPalestinaPA2bAAGm661 MK 2. PARATOPOTYPES: MPEG 41156 and CFBH 43130���43131, all adult males, collected (5.70300�� S, 48.24833�� W; 176 m) on January 9���10, 2018, by the same collectors. ETYMOLOGY: The epithet phonotriccus is the combination of Greek phono - (from phon��, ���sound, voice���) and triccus (a small bird whose species is not identifiable, though in modern times the name is applied to tyrant flycatchers; Jobling, 2010). This name is to be treated as a noun in apposition and is an allusion to the similarity between the vocalization of the new species and those of tody-tyrants. Tody-tyrants of the Neotropical genus Hemitriccus have peculiar vocalizations, which are reminiscent of the vocalization of Adenomera phonotriccus, especially the trilled song of H. cohnhafli (see Zimmer et al., 2013). Interestingly, several Hemitriccus species are morphologically very similar, and consequently best differentiated by their vocalizations, a pattern also observed in Adenomera. DIAGNOSIS: Adenomera phonotriccus is differentiated from its congeners by the following combination of characters: (1) small size (adult male SVL 19.8���21.6 mm; table 1); (2) robust body shape; (3) toe tips unexpanded to slightly swollen (character states B, C; Heyer, 1973); (4) distal antebrachial tubercle on underside of forearm; (5) throat and belly cream colored, mottled white and gray in some parts; (6) two color morphotypes (presence/absence of dorsolateral stripes); (7) advertisement call consisting of a single type of pulsed note, emitted regularly, not in calling bouts; (8) advertisement notes composed of complete pulses (pulses with periods of silence in between); (9) long-lasting call duration (213���433 ms). COMPARISONS: Adenomera phonotriccus has adult males (SVL 19.8���21.6 mm; table 1) smaller than those of A. coca (23.6���25.6 mm; Angulo and Reichle, 2008), A. lutzi (25.7���33.5 mm; Kok et al., 2007), and A. simonstuarti (SVL 25.9���26.2 mm; Angulo and Icochea, 2010). Adenomera phonotriccus has a robust body shape (figs. 1���2), whereas A. diptyx, A. martinezi, and A. saci have a slender body (Carvalho and Giaretta, 2013a). Adenomera phonotriccus has toe tips unexpanded or slightly swollen (character states B, C), but not expanded into flattened discs (character state D) as in A. ajurauna, A. andreae, A. marmorata, A. lutzi, A. nana, and A. simonstuarti (Angulo et al., 2003; Kok et al., 2007; Kwet, 2007; Berneck et al., 2008; Angulo and Icochea, 2010; appendix 1). Adenomera phonotriccus is distinguished from congeners (except A. cotuba and A. lutzi; Kok et al., 2007; Carvalho and Giaretta, 2013b) by having antebrachial tubercles on underside of forearm (fig. 3), presumably absent in all remaining species. Adenomera phonotriccus differs from A. heyeri and A. lutzi by having ventral surfaces cream colored (mottled white and gray in some parts), yellow in the latter two species (Boistel et al., 2006; Kok et al., 2007). From A. engelsi, which does not have distinctive dorsolateral stripes and pigmentation on throat (Kwet et al., 2009), Adenomera phonotriccus differs by the occurrence of both color features (fig. 1). Adenomera cotuba does not have dorsolateral stripes (Carvalho and Giaretta, 2013b), whereas A. phonotriccus has two color morphotypes (presence/ absence of dorsolateral stripes). These species are better distinguished by their distinct vocalizations (see below; fig. 4). The advertisement call of Adenomera phonotriccu s (fig. 4; table 2) distinguishes it from all congeners by having pulsed notes composed of complete pulses (periods of silence in between), whereas the other species��� notes are formed by incomplete pulses, i.e., not separated one from the next by periods of silence (fig. 4). From the remaining species of Adenomera, the new spe- cies differs by having pulsed advertisement notes, whereas those species��� calls consist of nonpulsed notes (table 3). The advertisement call of A. phonotriccus is distinguished from that of A. cotuba by being composed of single notes emitted regularly (table 2), whereas A. cotuba��� s call is made up of many notes emitted as calling bouts, i.e., multinote advertisement call (Carvalho and Giaretta, 2013b: fig. 4A). In addition, to date, Adenomera phonotriccus has the longest note duration in the genus, whose range does not overlap with note-duration values for any congeners (table 3). DESCRIPTION OF HOLOTYPE: MPEG 41155 (figs. 1A, 2, 3A). Adult male. Body robust (figs. 1A, 2A). Snout subovoid in dorsal view (fig. 2A), acuminate in lateral view (fig. 2D). Nostrils closer to the snout tip than to the eyes; fleshy ridge on snout tip; canthus rostralis not marked; loreal region slightly concave; supratympanic fold from the posterior corner of the eye to the base of the arm; oval postcommissural gland; vocal sac subgular with a fold from jaw extending to forearm on each side, vocal slits present; vomerine teeth in two straight rows medial and posterior to choanae and oblique to sagittal plane. Tongue elongated, free behind. Relative finger lengths IV Adenomera, based on (quasi-) periodic sinusoidal FMs (sometimes there is strong modulation unaccompanied by distinct pulses though). b Based on the call of Adenomera sp. 2 from Joinville. c See Materials and Methods for a detailed explanation. d Frequencies lower than 1 kH do not pertain to this species��� call (appendix 2). COLOR OF HOLOTYPE IN PRESERVATIVE (fig. 2): Snout tip with a faded white coloration (coincident with the fleshy ridge). Dorsum marble gray with a few darker specks and spots. Sacral stripe on the posterior third of body length, cream-colored. Black lumbar glands. Dorsal surface of limbs with dark brown stripes/blotches on a slightly lighter brown background. Dorsolateral, broad stripe from the posterior corner of the eye, passing over the flanks, predominantly cream-colored to a somewhat orange hue (faded coloration; orange in life). Upper and lower jaws covered with white-colored spots/blotches, whitish postcommissural gland, outlined at the bottom by a dark coloration. Tympanum reddish brown. Throat, belly, and ventral surface of limbs cream, with melanophores, throat with a black-dotted pattern, solid dark-colored laterally, coinciding with the expanded vocal sac. Posterior surface of thighs with a few dark brown spots on a light brown background, and paracloacal glands cream colored. COLOR OF HOLOTYPE IN LIFE (fig. 1A): Dorsum covered with black specks and spots irregularly distributed on medium brown background. Arms and legs reddish brown. Sacral and dorsolateral stripe, and postcommissural gland orange. Iris copper. Tympanum brown. Ventral surfaces cream colored and with mottling on throat and belly (laterally). Groin yellow. VARIATION IN THE TYPE SERIES: Despite minor variation in dorsal coloration and distribution of spots, specks, and warts on dorsal surfaces, characteristics of the three paratypes largely agree with the description of the holotype. Dorsolateral stripe is absent in two paratypes (MPEG 41156 and CFBH 43130). Shape and size of antebrachial tubercle varies among the type series; some of them are very discrete (fig. 2). ADVERTISEMENT CALL: The description is based on recordings of three males (appendix 3). Sample sizes for each acoustic trait and descriptive statistics, i.e., means and standard deviations, are provided in table 2. The call (fig. 4) consists of a single type of pulsed note, which is emitted at a rate of 26���35 calls per minute. Note duration varies from 213���433 msec. Notes have a smooth attack and decay, reaching the amplitude peak at 43%���85% of their duration. Pulse number varies from 14���26; pulse duration from 4���26 msec, emitted at a rate of 51���67 pulses per second; silence between pulses varies from nearly zero to 13 msec. The fundamental frequency of the note occupies a narrow bandwidth harmonic, with a peak frequency of 1857��� 1997 Hz; the dominant frequency varies from 3639���4113 Hz and is contained within the second harmonic. Notes usually have a slight positive frequency modulation along their duration, but a few notes do not have any modulation, and a single note has negative modulation. Linear frequency modulation varies from -43���861 Hz. HABITAT AND NATURAL HISTORY: Adenomera phonotriccus is associated with forest habitats in Brazil���s eastern Amazonia in the state of Par��. The type locality is located on the west margin of the lower Araguaia River. This region is also occupied by savanna landscapes or ecotones between the Cerrado and Amazonian rainforest. The species was heard along an open area transect (by recent logging activity), especially at the forest edge, and inside the forest remnant. Males called while exposed or under leaf litter, and increased their calling activity during and shortly after rainfalls. The vocalization of Adenomera phonotriccus is very peculiar when heard in the field. Our first impression was that the calls might not be produced by a leptodactylid frog, or even an anuran species. Indeed, the vocalization is similar to that of some Neotropical suboscine birds (e.g., Rhynchocyclidae, Tyrannidae). There were two partially syntopic species of Adenomera also in calling activity at the study site: Adenomera sp. (A. heyeri clade) and A. aff. hylaedactyla. We heard the three species calling at the same period (late afternoon), even though A. phonotriccus and Adenomera sp. ceased most calling activity at dusk, whereas A. aff. hylaedactyla continued calling through the first hours of the night. The last species occupied open areas, e.g. pasture. In contrast, A. phonotriccus and Adenomera sp., are associated with forest habitats. DISTRIBUTION (fig. 5): Adenomera phonotriccus is known with certainty only from the type locality and Marab��. However, specimens from other regions in the state of Par�� were also assigned to this lineage, referred as Adenomera sp. F, by Fouquet et al. (2014). Given the high levels of cryptic species diversity in Adenomera and complex genetic structure within the lineages identified by Fouquet et al. (2014), we conservatively restrict the species��� distribution to the type locality region until additional data, especially vocalizations, are obtained for the other populations assigned to Adenomera sp. F, a lineage widely distributed in central-east Par��, northern Brazil. CONSERVATION STATUS: Because the species is apparently restricted to forested habitats, or its immediate margins (forest edge), we infer that the current population trend for this species is ���declining.��� Major threats are deforestation (destruction of habitats) due to conversion of large areas where the species occur into pasture and agricultural land. The species appears to be locally abundant (although difficult to observe), which means that as long as its habitat is preserved the species should have its future secured. The known range of the species is hard to determine with current evidence. It may be much wider, if all the clade F specimens refer to A. phonotriccus, in which case the species would be found from the western margin of the Tocantins River to the western margins of the Tapaj��s River (Fouquet et al., 2014: fig. 2). Alternatively, it could have a much narrower distribution (southern Par��) if only specimens in subclade F3 correspond to A. phonotriccus ���in the latter case, the species would be probably restricted to small forest fragments in one of the most highly threatened portions of the Amazonia (the so-called Arc of Deforestation). This is a typical example of the issues raised by Peloso (2010) and Angulo and Icochea (2010), where lack of taxonomic resolution hinders a proper assessment of threats to cryptic species. For now, we must unfortunately suggest this new species is yet another case of data-deficient taxa���a lot more studies and data are needed to define the conservation status of A. phonotriccus. REMARKS: Calls of the voucher PV 2597 (specimen no. DT 2123; recordings PLPDR 100; see appendix 3) were unambiguously assigned to A. phonotriccus (���F3��� in fig. 6). This voucher was cited as collected in Araguatins (state of Tocantins), on the east margin of Araguaia River. However, this specimen was actually collected in Palestina do Par��, state of Par�����the animal was collected by P.L.V.P. in 2010 (coordinates given by Fouquet et al.���s appendix 1 are correct). The basecamp for this collection site was in Araguatins, and this may have been the source of confusion with the sample provenance (P.L.V.P., personal obs.). Another voucher specimen used in the phylogeny (PV 2412, specimen no. DT 2016), also assigned here to A. phonotriccus (���F3��� in fig. 6) was collected in Marab�����this municipality is located in the state of Par��, not Amazonas (AM), as mentioned in appendix 1 of Fouquet et al. (2014). Of special interest is the acquisition of additional information for the subclade F1, which occurs in partial sympatry with A. phonotriccus (fig. 6). Meanwhile, we conservatively assign only subclade F3 (sensu Fouquet et al., 2014) to A. phonotriccus, and this species is an endemic of the central-east part of the state of Par�� in association with the west margin of the lower Araguaia River (fig. 6). With respect to subclade F2, we suspect that it may not be conspecific with A. phonotriccus, given its distribution range associated with another region in the state of Par�� (Xingu-Tapaj��s interfluve; fig. 6). Therefore, we maintain the original status of unconfirmed candidate species of both subclades F1 and F2 for the moment., Published as part of De Carvalho, Thiago R., Giaretta, Ariovaldo A., Angulo, Ariadne, Haddad, C��lio F. B. & Peloso, Pedro L. V., 2019, A new Amazonian species of Adenomera (Anura: Leptodactylidae) from the Brazilian state of Par��: a tody-tyrant voice in a frog, pp. 1-24 in American Museum Novitates 2019 (3919) on pages 5-14, DOI: 10.1206/3919.1, http://zenodo.org/record/4585412, {"references":["Fouquet, A., C. S. Cassini, C. F. B. Haddad, N. Pech, and M. T. Rodrigues. 2014. Species delimitation, patterns of diversification and historical biogeography of the Neotropical frog genus Adenomera (Anura, Leptodactylidae). Journal of Biogeography 41: 855 - 870.","Jobling, J. A. (editor). 2010. Helm dictionary of scientific bird names. London: A & C Black Publishers Ltd.","Zimmer, K. J., A. Whittaker, C. H. Sardelli, E. Guilherme, and A. Aleixo. 2013. A new species of Hemitriccus tody-tyrant from the state of Acre, Brazil. In J. Del Hoyo, A. Elliott, and D. Christie (editors), Handbook of the birds of the world, special volume: new species and global index: 292 - 296. Barcelona: Lynx Edicions.","Heyer, W. R. 1973. Systematics of the marmoratus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Contributions in Science, Los Angeles County Natural History Museum 251: 1 - 50.","Angulo, A., and S. Reichle. 2008. Acoustic signals, species diagnosis, and species concepts: the case of a new cryptic species of Leptodactylus (Amphibia, Anura, Leptodactylidae) from the Chapare Region, Bolivia. Zoological Journal of the Linnean Society 152: 59 - 77.","Kok, P. J. R., M. N. C. Kokubum, R. D. MacCulloch, and M. Lathrop. 2007. Morphological variation in Leptodactylus lutzi (Anura, Leptodactylidae) with description of its advertisement call and notes on its courtship behavior. Phyllomedusa 6: 45 - 60.","Angulo, A., and J. Icochea. 2010. Cryptic species complexes, widespread species and conservation: lessons from Amazonian frogs of the Leptodactylus marmoratus group (Anura: Leptodactylidae). Systematics and Biodiversity 8: 357 - 370.","Carvalho, T. R., and A. A. Giaretta. 2013 a. Taxonomic circumscription of Adenomera martinezi (Bokermann, 1956) (Anura: Leptodactylidae: Leptodactylinae) with the recognition of a new cryptic taxon though a bioacoustic approach. Zootaxa 3701: 207 - 237.","Angulo, A., R. B. Cocroft, and S. Reichle. 2003. Species identity in the genus Adenomera (Anura: Leptodactylidae) in southeastern Peru. Herpetologica 59: 490 - 504.","Kwet, A. 2007. Bioacoustic variation in the genus Adenomera in southern Brazil, with revalidation of Leptodactylus nanus Muller, 1922 (Anura, Leptodactylidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe Supplement 83: 56 - 68.","Berneck, B. V. M., C. O. R. Costa, and P. C. A. Garcia. 2008. A new species of Leptodactylus (Anura: Leptodactylidae) from the Atlantic Forest of Sao Paulo State, Brazil. Zootaxa 1795: 46 - 56.","Carvalho, T. R., and A. A. Giaretta. 2013 b. Bioacoustics reveals two new syntopic species of Adenomera Steindachner (Anura: Leptodactylidae: Leptodactylinae) in the Cerrado of central Brazil. Zootaxa 3731: 533 - 551.","Boistel, R., G. C. Massary, and A. Angulo. 2006. Description of a new species of the genus Adenomera (Amphibia, Anura, Leptodactylidae) from French Guyana. Acta Herpetologica 1: 1 - 14.","Kwet, A., J. Steiner, and A. Zillikens. 2009. A new species of Adenomera (Amphibia: Anura: Leptodactylidae) from the Atlantic rain forest in Santa Catarina, southern Brazil. Studies on Neotropical Fauna and Environment 44: 93 - 107.","Kwet, A., and A. Angulo. 2002. A new species of Adenomera (Anura, Leptodactylidae) from the Araucaria forest of Rio Grande do Sul (Brazil), with comments on the systematic status of southern populations of the genus. Alytes 20: 28 - 43.","Marquez, R., I. de la Riva, and J. Bosch. 1995. Advertisement calls of Bolivian Leptodactylidae (Amphibia, Anura). Journal of Zoology 237: 313 - 336.","Straughan, I. R., and W. R. Heyer. 1976. A functional analysis of the mating calls of the Neotropical frog genera of the Leptodactylus complex (Amphibia, Leptodactylidae). Papeis Avulsos de Zoologia, Sao Paulo 29: 221 - 245.","Almeida, A. P., and A. Angulo. 2006. A new species of Leptodactylus (Anura: Leptodactylidae) from the state of Espirito Santo, with remarks on the systematics of associated populations. Zootaxa 1334: 1 - 25.","Peloso, P. L. C. 2010. A safe place for amphibians? A cautionary tale on the taxonomy and conservation of frogs, caecilians, and salamanders in the Brazilian Amazonia. Zoologia 27: 667 - 673."]}

Pseudopaludicola matuta sp. nov. urn:lsid:zoobank.org:act: DF080E14-2F88-45CC-876B-147D2D06C6D9 Figs 2���5, Tables 1���4 Pseudopaludicola sp. ��� Carvalho (2012): 52, Natural History Section. Pseudopaludicola mineira from Pampulha, Belo Horizonte, Minas Gerais ��� Andrade et al. 2018: 87, Discussion Section. Diagnosis Pseudopaludicola matuta sp. nov. is assigned to the genus Pseudopaludicola by having a hypertrophied antebrachial tubercle (see Lynch 1989, Lobo 1995). The new species is characterized by the following combination of characters: (1) small size (SVL 12.8���14.2 mm in adult males); (2) absence of both, enlarged palpebral tubercles and an enlarged conical tubercle on heel; (3) relatively short hind limbs (tibiotarsal articulation only reaching the posterior margin of the eye); and (4) advertisement call composed of regular series of stereotyped two-pulsed notes, with notes having a short duration and emitted at a higher rate (notes per minute). Differential diagnosis Pseudopaludicola matuta sp. nov. is promptly distinguished from the species of the P. saltica group (P. saltica, P. murundu, and P. jaredi) by having relatively short hind limbs; i.e., the tibiotarsal articulation reaches the posterior margin of the eye in the new species, whereas it reaches the tip of the snout in species of the P. saltica group (Andrade et al. 2016). Pseudopaludicola matuta sp. nov. is distinguished from species of the P. pusilla group (P. boliviana, P. ceratophyes, P. llanera, P. pusilla, and P. motorzinho) by the absence of either T-shaped terminal phalanges or expanded toe tips (disks or pads). The tips of the phalanges of the new species are not T-shaped; they are similar in shape to those of P. falcipes (fig. 2B in Cardozo & Su��rez 2012). The new species is also distinguished from P. ceratophyes by the absence of an enlarged palpebral tubercle (Lynch 1989). Pseudopaludicola matuta sp. nov. differs from P. boliviana and P. motorzinho by the absence of an enlarged, conical tubercle on the heel (Pansonato et al. 2016). Pseudopaludicola matuta sp. nov. has an advertisement call composed of series of notes with two nonconcatenated pulses each (Fig. 4 A���B); therefore, it is promptly distinguished from all species with advertisement call composed of non-pulsed notes: P. canga Giaretta & Kokubum, 2003 (Giaretta & Kokubum 2003; Pansonato et al. 2012; Carvalho et al. 2015a), P. giarettai (Carvalho 2012; Carvalho et al. 2015b), P. hyleaustralis Pansonato, Morais, ��vila, Kawashita-Ribeiro, Str��ssmann & Martins, 2012 (Pansonato et al. 2012), P. facureae (Andrade & Carvalho 2013; Carvalho et al. 2015a), and P. parnaiba Roberto, Cardozo & ��vila, 2013 (Roberto et al. 2013; Carvalho et al. 2015a); and from all species with notes with concatenated pulses (= lack of interpulse interval; sensu Magalh��es et al. 2014): P. mystacalis [12���14 concatenated pulses; Pansonato et al. 2013], P. boliviana [3���6; Dur�� et al. 2004], P. ibisoroca Pansonato, Veiga-Menoncello, Mudrek, Jansen, Recco-Pimentel, Martins & Str��ssmann, 2016 [3���12; Pansonato et al. 2016], and P. motorzinho [2���6; Pansonato et al. 2016]. Pseudopaludicola matuta sp. nov. is distinguished from other congeners with notes with non-concatenated pulses (values within square brackets ���[]���) by the following acoustic traits: P. ameghini (Cope, 1887) has longer note duration (27���52 [60���121] ms), higher number of pulses per note (2 [3���5]), and lower note rate (542���1101 [348���452] notes per minute) (Andrade et al. 2017a); P. ternetzi Miranda-Ribeiro, 1937 has higher number of pulses per note [3���6] (Andrade et al. 2017a); P. atragula Pansonato, Mudrek, Veiga-Menoncello, Rossa-Feres, Martins & Str��ssmann, 2014 has longer note duration [300���700 ms], higher number of pulses per note [9���36], and lower note rate [42���98 notes per minute] (Pansonato et al. 2014). The three species of the P. saltica species group have lower note rate [180���480 notes per minute, combined values], and vary highly the number of pulses in their notes (2���7 pulses per note, combined values; Andrade et al. 2016). Pseudopaludicola falcipes and P. restinga Cardozo, Baldo, Pupin, Gasparini & Haddad, 2018 have lower note rates [238���535 notes per minute] (Andrade et al. 2018; Cardozo et al. 2018). Pseudopaludicola pocoto Magalh��es, Loebmann, Kokubum, Haddad & Garda, 2014 and P. florencei Andrade, Haga, Lyra, Leite, Kwet, Haddad, Toledo & Giaretta, 2018 have stereotyped three-pulsed notes along their series of notes, longer note duration [108���397 ms, combined values], and lower note rate [100���297 notes per minute, combined values] (Magalh��es et al. 2014; Andrade et al. 2017b, 2018). In comparison with the phylogenetically close related species, the new species is distinguished from P. mineira by having shorter note duration [50���114 ms] and lower note rate [282���485 notes per minute] (Table 2). The advertisement call of P. ceratophyes, P. llanera, and P. pusilla are unknown; however, there are strong morphological differences among them and P. matuta sp. nov. In relation to the most closely related species, P. mineira, the randomForest multivariate approach applied to morphometric data indicated a broad overlap between the two partitions (Fig. 5 A���B), with a considerable classification error (Table 3). We found no statistical significance in any trait. Therefore, we were unable to distinguish P. matuta sp. nov. from P. mineira based on morphometric or any other feature of external morphology or coloration. In contrast, the RandomForest multivariate approach on acoustic data highlighted a full segregation between P. matuta sp. nov. and P. mineira (Table 3, Fig. 5C), without any classification error. Pulse rate, note duration, notes per minute, and interpulse interval are the main sources of variation in both variable importance measurements (Fig. 5D). The trait of notes per minute for P. mineira did not overlap the values for the new species either. It is noteworthy that the air temperature overlapped at the time of the field recordings for both species (Table 2), and is among the variables that least explained the variation of the dataset, with low importance measures as produced by randomForest (Fig. 5D). Therefore, we do not attribute this found acoustic difference to the influence of the air temperature. In addition, there was only a slight overlap between both species in note duration (see Table 2). The new species can be significantly differentiated from P. mineira in the following traits: note duration, internote interval, interpulse interval, pulse rate, and dominant frequency (for all these traits, Wilcoxon-Mann-Whitney Tests had P matuta in Brazilian Portuguese means rustic, provincial, related to those who live in the countryside. Type material Holotype BRAZIL: adult ��, Minas Gerais, municipality of Curvelo, S��tio Mato do Engenho, 18��46���07.6��� S, 44��26���50.7��� W, 620 m a.s.l., 29 Nov. 2017, F.S. Andrade & I.A. Haga leg. (ZUEC 24302; Figs 2���3, call voucher). Paratopotypes BRAZIL: 12 adult ����, same collection data as the holotype (ZUEC 24303���6, 24308���10, 24313, 24315��� 8); 3 adult ����, same locality as holotype, 21 Feb. 2011, T. R. Carvalho leg. (AAG-UFU 0308, 0386���7); 5 adult ♀♀, same collection data as the holotype (ZUEC 24307, 24311���2, 24314, 24323). Type locality BRAZIL: Minas Gerais, municipality of Curvelo, S��tio Mato do Engenho, 18��46���07.6��� S, 44��26���50.7��� W, 620 m a.s.l. Other material examined BRAZIL: 3 adult ����, Minas Gerais, municipality of Santana do Riacho, district of Serra do Cip��, southern foothills of Serra do Cip�� National Park, near Lagoa da Capivara (19��20���46.30��� S, 43��36���59.62��� W, 797 m a.s.l.), 2���3 Dec. 2017, F.S. Andrade & I.A. Haga leg. (ZUEC 24324, 24327, 24329); 3 adult ♀♀, same collection data as previous (ZUEC 24325���6, 24328). Description of the holotype Body elliptic and broad (Table 1, Fig. 2 A���B). Head elliptical, slightly wider than long. Snout subovoid in dorsal view and rounded in profile (Fig. 2 C���D). Eye not protuberant. Eye diameter almost equal to the interorbital distance. Interorbital area flat. Pupil rounded. Upper eyelid without tubercles. Nostril not protuberant and closer to the snout tip than to the eye. Canthus rostralis rounded, smooth. Loreal region slightly concave. Single subgular vocal sac, externally expanded and with discrete longitudinal folds. Choanae rounded, well separated from each other. Vocal slits present. Tympanum indistinct. A discrete tympanic ridge from behind the eye to the proximal portion of the arm. Mouth opening ventral. Vomerine teeth absent. Tongue elliptical, longer than wide, posteriorly free, without pigmentation at its base. Flank with discrete granules. One ovoid antebrachial tubercle present in the first quarter of the forearm and a second ovoid tubercle closer to elbow. Finger and toe tips not expanded. Outer and inner metacarpal tubercles well-defined, outer metacarpal tubercle ovoid and inner metacarpal tubercle rounded. Fingers with single and rounded subarticular tubercles. Supernumerary tubercles absent on palm of hand. Thumb with a keratinized, light brown nuptial pad, extending from the base of the hand to the proximal limit of the terminal phalanx, covering almost the entire external portion of the finger. Webbing absent between fingers. Relative finger lengths, when adpressed one to another: I Pseudopaludicola matuta sp. nov. emits the advertisement call with highly variable duration (3.6��� 66.2 s), consisting of series of stereotyped two-pulsed notes (1���9 series of two-pulsed notes per call) that lasts 0.2��� 34.0 s, separated by intervals of 0.2��� 1.9 s (Fig. 4A). Before the emission of the series of stereotyped two-pulsed notes, 1���10 (mean = 4.4, SD = 3.3) isolated pulsed notes with irregular structure, duration, interval, and number of pulses are emitted, herein referred to as introductory notes (Fig. 4A). Introductory notes last 29���117 ms (mean 64, SD = 24), separated by intervals of 37���495 ms (mean = 213, SD = 111), and are composed of 2���5 non-concatenated pulses (mean = 3.3, SD = 0.7). These pulses vary from 4���16 ms (mean = 9, SD = 2), separated by intervals of 1���36 ms (mean = 16, SD = 6). Dominant frequency peaks are between 4.1���4.5 kHz (mean = 4.4, SD = 0.2). On the other hand, within the series of stereotyped two-pulsed notes, the notes have regular structure, duration, interval, and number of pulses. These notes last 27���52 ms, separated by intervals of 17���76 ms, and are released at a rate of 542���1,101 notes per minute; notes have a slightly increase in amplitude from the first to the second pulse (see oscillogram in Fig. 4B). Pulses vary from 4���15 ms, separated by intervals of 11���40 ms, and are released at a rate of 43���74 pulses per second (Fig. 2B). Dominant (= fundamental) frequency peaks are between 3.9���5.1 kHz; the minimum frequency ranges between 1.8���4.6 kHz and the maximum frequency ranges from 4.4���5.5 kHz. Notes have up to three harmonics; the second ranging from 6.9���9.8 kHz (mean = 8.7; SD = 0.4). Air temperature of recorded calls varied from 24.0 to 25.2��C. Call quantitative traits are summarized in Table 2. Traits that were classified as static (within-male CV P. matuta sp. nov. were note duration (mean = 4.2%, SD = 1.8 [1.6���7.4]), number of pulses per note (mean = 0%), pulse rate (mean = 4.2%, SD = 1.8 [1.6���7.4]), and dominant peak (mean = 0.9%, SD = 0.7 [0.0���3.0]). The other traits were classified as dynamic (CV P. matuta sp. nov., in which the three males of the population from the southern foothills of the Serra do Cip�� National Park have a lower note rate (529���737 notes per minute) than those of male types (738���1101 notes per minute). It is noteworthy that the air temperatures measured during the recordings in these two localities did not overlap, 24.0���25.2��C at type locality, and 20.5���23.0��C in lowlands of the Serra do Cip�� National Park. Phylogenetic inferences and mitochondrial DNA divergences The inferred tree topology agreed with previous phylogenetic analyses of Pseudopaludicola (Veiga- Menoncello et al. 2014; Andrade et al. 2016, 2018), and was very similar between BI and ML inferences (Fig. 6). The new species was found as sister clade of P. mineira (Fig. 6). Uncorrected genetic distance between the new species and P. mineira was 3.5% (mean value) and maximum intraspecific distance was 0.1% (Table 4). No molecular data are available for P. ceratophyes, P. llanera, P. pusilla, P. hyleaustralis, P. parnaiba, P. giarettai, P. ibisoroca; however, the new species is strongly diagnosed from these species by morphology and acoustics (see further details in Differential diagnosis section). Natural history notes We recorded specimens of P. matuta sp. nov. in an area with relatively dense vegetation. In this site, there were grasses, shrub vegetation, and some spaced trees of 5���7 m in height, with the soil well soaked. Males of other Pseudopaludicola species usually call exposed in open areas. Curiously, two other specimens of P. matuta sp. nov. were recorded a few meters from this above-mentioned site in an open and waterlogged area, where P. giarettai also occurs syntopically. In both places, we observed dozens of specimens of P. matuta sp. nov. However, during the fieldwork in the lowlands of the Serra do Cip�� National Park, we were able to find a single small population near Lagoa da Capivara and Cip�� River. During the two sampling nights, it was possible to hear only eight males, which were recorded. They were not excited and were vocalizing wellspaced from each other. This social condition may have influenced the intraspecific acoustic variation described by us. Besides that, air temperature was lower (without overlap) during the fieldwork in the lowlands of the Serra do Cip�� National Park. Distribution Pseudopaludicola matuta sp. nov. is known from its type locality, lowlands of the Serra do Cip�� National Park (southern foothills), district of Serra do Cip��, municipality of Santana do Riacho, and in the municipality of Belo Horizonte from a report in the 1960s (Fig. 1). However, it is not possible to find this species nowadays in the municipality of Belo Horizonte (F.S.F. Leite, pers. obs.). Lobo (1994) reported the occurrence of P. mineira in the Serra do Cip�� National Park (at 1264 m a.s.l., type locality) and in Serra do Cabral (at 1104 m a.s.l.), a nearby mountainous area but isolated from the main Espinha��o mountain range (Fig. 1). The altitudes we found P. matuta sp. nov. vary between 620 and 883 m a.s.l. In addition, in the type locality and where we collected P. mineira during our fieldwork are only about 14 km northeast from the site where we recorded P. matuta sp. nov. (797 m a.s.l., Fig. 1); however, the elevational difference between these two sites is about 460 m. We did not find P. matuta sp. nov. during our fieldwork in the rupestrian grasslands of the Serra do Cip�� National Park. Therefore, it seems that P. mineira is restricted to the higher rupestrian grasslands of the Serra do Cip�� National Park and Serra do Cabral (Lobo 1994), whereas P. matuta sp. nov. occurs in adjacent lowl, Published as part of Andrade, Felipe Silva de, Haga, Isabelle Aquemi, Lyra, Mariana L��cio, Carvalho, Thiago Ribeiro de, Haddad, C��lio Fernando Baptista, Giaretta, Ariovaldo Antonio & Toledo, Lu��s Felipe, 2018, A new species of Pseudopaludicola (Anura, Leptodactylidae, Leiuperinae) from the state of Minas Gerais, Brazil, pp. 1-25 in European Journal of Taxonomy 480 on pages 6-18, DOI: 10.5852/ejt.2018.480, http://zenodo.org/record/3825293, {"references":["Carvalho T. R. 2012. A new species of Pseudopaludicola Miranda-Ribeiro (Leiuperinae: Leptodactylidae: Anura) from the Cerrado of southeastern Brazil with a distinctive advertisement call pattern. Zootaxa 3328: 47 - 54.","Andrade F. S., Haga I. A., Lyra M. L., Leite F. S. F., Kwet A., Haddad C. F. B., Toledo L. F. & Giaretta A. A. 2018. A new species of Pseudopaludicola Miranda-Ribeiro (Anura: Leptodactylidae: Leiuperinae) from eastern Brazil, with novel data on the advertisement call of Pseudopaludicola falcipes (Hensel). Zootaxa 4433 (1): 71 - 100. https: // doi. org / 10.11646 / zootaxa. 4433.1.4","Lynch J. D. 1989. A review of leptodactylid frogs of the genus Pseudopaludicola in northern South America. Copeia 1989 (3): 577 - 588. https: // doi. org / 10.2307 / 1445483","Lobo F. 1995. Analisis filogenetico del genero Pseudopaludicola (Anura: Leptodactylidae). Cuadernos de Herpetologia 9: 21 - 43.","Andrade F. S., Magalhaes F. M., Nunes-de-Almeida C. H. L., Veiga-Menoncello A. C. P., Santana D. J., Garda A. A., Loebmann D., Recco-Pimentel S. M., Giaretta A. A. & Toledo L. F. 2016. A new species of long-legged Pseudopaludicola from northeastern Brazil (Anura, Leptodactylidae, Leiuperinae). Salamandra 52: 107 - 124.","Cardozo D. & Suarez P. 2012. Osteological description of Pseudopaludicola canga with implications for the taxonomic position of this taxon. Zootaxa 3515: 75 - 82.","Pansonato A., Veiga-Menoncello A. C. P., Mudrek J. R., Jansen M., Recco-Pimentel S. M., Martins I. A. & Strussmann C. 2016. Two new species of Pseudopaludicola (Anura: Leptodactylidae: Leiuperinae) from eastern Bolivia and western Brazil. Herpetologica 72: 235 - 255. https: // doi. org / 10.1655 / Herpetologica-D- 14 - 00047.1","Giaretta A. A. & Kokubum M. N. C. 2003. A new species of Pseudopaludicola (Anura, Leptodactylidae) from northern Brazil. Zootaxa 383: 1 - 8. https: // doi. org / 10.11646 / zootaxa. 383.1.1","Pansonato A., Morais D. H., Avila R. W., Kawashita-Ribeiro R. A., Strussmann C. & Martins I. A. 2012. A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leiuperidae) from the state of Mato Grosso, Brazil, with comments on the geographic distribution of Pseudopaludicola canga Giaretta & Kokubum, 2003. Zootaxa 3523: 49 - 58.","Carvalho T. R., Teixeira B. F. V., Martins L. B. & Giaretta A. A. 2015 a. Intraspecific variation and new distributional records for Pseudopaludicola species (Anura, Leptodactylidae, Leiuperinae) with trilled advertisement call pattern: diagnostic characters revisited and taxonomic implications. North-Western Journal of Zoology 11: 262 - 273.","Carvalho T. R., Borges-Martins M., Teixeira B. F. V., Godinho L. B. & Giaretta A. A. 2015 b. Intraspecific variation in acoustic traits and body size, and new distributional records for Pseudopaludicola giarettai Carvalho, 2012 (Anura, Leptodactylidae, Leiuperinae): implications for its congeneric diagnosis. Papeis Avulsos de Zoologia 55: 245 - 254. https: // doi. org / 10.1590 / 0031 - 1049.2015.55.17","Andrade F. S. & Carvalho T. R. 2013. A new species of Pseudopaludicola Miranda-Ribeiro (Leiuperinae: Leptodactylidae: Anura) from the Cerrado of southeastern Brazil. Zootaxa 3608 (5): 389 - 397. https: // doi. org / 10.11646 / zootaxa. 3608.5.7","Roberto I. J., Cardozo D. & Avila R. W. 2013. A new species of Pseudopaludicola (Anura, Leiuperidae) from western Piaui State, Northeast Brazil. Zootaxa 3636: 348 - 360. https: // doi. org / 10.11646 / zootaxa. 3636.2.6","Magalhaes F. M., Loebmann D., Kokubum M. N. C., Haddad C. F. B. & Garda A. A. 2014. A new species of Pseudopaludicola (Anura: Leptodactylidae: Leiuperinae) from northeastern Brazil. Herpetologica 70: 77 - 88. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00054","Pansonato A., Strussmann C., Mudrek J. R. & Martins I. A. 2013. Morphometric and bioacoustic data on three species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) described from Chapada dos Guimaraes, Mato Grosso, Brazil, with the revalidation of Pseudopaludicola ameghini (Cope, 1887). Zootaxa 3620 (1): 147 - 162. https: // doi. org / 10.11646 / zootaxa. 3620.1.7","Dure M. I., Schaefer E. F., Hamann M. I. & Kehr A. I. 2004. Consideraciones ecologicas sobre la dieta, la reproduccion y el parasitismo de Pseudopaludicola boliviana (Anura, Leptodactylidae) de Corrientes, Argentina. Phyllomedusa 3: 121 - 131. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 3 i 2 p 121 - 131","Andrade F. S., Haga I. A., Bang D. L. & Giaretta A. A. 2017 a. The differential acoustic diagnosis between two Pseudopaludicola sister species (Anura, Leptodactylidae, Leiuperinae). Zootaxa 4319 (2): 391 - 400. https: // doi. org / 10.11646 / Zootaxa. 4319.2.12","Pansonato A., Mudrek J. R., Veiga-Menoncello A. C. P., Rossa-Feres D. C., Martins I. A. & Strussmann C. 2014. A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) from northwestern state of Sao Paulo, Brazil. Zootaxa 3861: 249 - 264. https: // doi. org / 10.11646 / zootaxa. 3861.3.3","Cardozo D. E., Baldo D., Pupin N., Gasparini J. L. & Haddad C. F. B. 2018. A new species of Pseudopaludicola (Anura, Leiuperinae) from Espirito Santo, Brazil. PeerJ 6: e 4766. https: // doi. org / 10.7717 / peerj. 4766","Andrade F. S., Leite F. S. F., Carvalho T. R., Bernardes C. S. & Giaretta A. A. 2017 b. First record of Pseudopaludicola pocoto Magalhaes, Loebmann, Kokubum, Haddad & Garda, 2014 (Anura, Leptodactylidae, Leiuperinae) in Bahia state, northeastern Brazil, with further data on its advertisement call. Check List 13 (1): 2047. https: // doi. org / 10.15560 / 13.1.2047","Lobo F. 1994. Descripcion de una nueva especie de Pseudopaludicola (Anura: Leptodactylidae), redescripcion de P. falcipes (Hensel, 1867) y P. saltica (Cope, 1887). Cuadernos de Herpetologia 8: 177 - 199."]}

Brachycephalus actaeus sp. nov. Figures 1, 2, 6, 7, and 8 Holotype. CFBH 39850, adult male, collected at Serra da Palha, Laranjeiras, Ilha de S��o Francisco do Sul, municipality of S��o Francisco do Sul, state of Santa Catarina, Brazil (26��17'50"S; 48 �� 40'28"W, Datum WGS 84, ca 60 meters above sea level), on 19 May 2015, by C.F.B. Haddad, J.P.C. Monteiro, and E.C. Nardin (Figures 1 and 2). Paratopotypes. CFBH 39851, adult female, collected with the holotype; CFBH 39872, 39873, and UFMG 18973, adult males, collected on 25 November 2015, by J.P.C. Monteiro, T.H. Condez, and E.C. Nardin; CFBH 39876, adult male, and CFBH 39877, adult female, collected on 0 2 December 2015, by J.P.C. Monteiro and E.C. Nardin. Paratypes. CFBH 39846, adult male, cleared and double-stained, collected at Fazenda Morro Grande, Morro Grande, Ilha de S��o Francisco do Sul, municipality of S��o Francisco do Sul, state of Santa Catarina, Brazil (26 �� 17'47"S; 48��37'10"W, Datum WGS 84, ca 60 meters above sea level), on 14 November 2014, by E.J. Comitti. UFMG 18970, adult female, collected on 15 November 2014, by J.P.C. Monteiro, T.H. Condez, and E.J. Comitti; CFBH 39849, sub-adult female, collected on 21 November 2014, by J.P.C. Monteiro and E.C. Nardin; CFBH 39848, adult male, cleared and double-stained, collected on 21 January 2015, by C.F.B. Haddad, J.P.C. Monteiro, T.H. Condez, and E.J. Comitti; UFMG 18971, adult male, collected on 0 1 August 2015, by J.P.C. Monteiro and E.C. Nardin; CFBH 39855���39858 and 39861, adult males, CFBH 39860, adult female, CFBH 39859, juvenile, collected on 23 November 2015, by J.P.C. Monteiro, T.H. Condez, and E.C. Nardin; all collected at Centro de Estudos e Pesquisas Ambientais da Univille (CEPA), Vila da Gl��ria, Distrito do Sa��, municipality of S��o Francisco do Sul, state of Santa Catarina, Brazil (26 �� 13'39"S; 48 �� 41'31"W, Datum WGS 84, ca 120 meters above sea level). CFBH 39853 and 39854, adult males, collected on 27 August 2015, by J.P.C. Monteiro and E.C. Nardin; CFBH 39862, adult female, collected on 23 November 2015, by J.P.C. Monteiro, T.H. Condez, and E.C. Nardin; at Estrada do Sa��, Distrito do Sa��, municipality of S��o Francisco do Sul, state of Santa Catarina, Brazil (26��12'06"S; 48 �� 41'37"W, Datum WGS 84, ca 80 meters above sea level). CFBH 39863, 39864, 39867, 39868, and UFMG 18972 adult females, CFBH 39865 and 39870, adult males, collected on 24 November 2015, by J.P.C. Monteiro, T.H. Condez, and E.C. Nardin; CFBH 39875, adult female, collected on 30 November 2015, by J.P.C. Monteiro and E.C. Nardin; at Bra��o do Norte, municipality of Itapo��, state of Santa Catarina, Brazil (26��07'29"S; 48 �� 43'48"W, Datum WGS 84, ca 220 meters above sea level). CFBH 42005���42008, adult females, collected on 17 September 2016, by J.P.C. Monteiro and E.C. Nardin; Fazenda Palmito Juriti, municipality of S��o Francisco do Sul, state of Santa Catarina, Brazil (26��08'09"S; 48 �� 43'54"W, Datum WGS 84, 125���170 meters above sea level). Diagnosis. Brachycephalus actaeus sp. nov. is a new species of the B. pernix group, distinguished from all its congeners by the following combination of characters: (1) ���bufoniform��� body; (2) general dorsal body color dark green with a dark brown vertebral stripe, and orange background more evident in ventral view; (3) absence of hyperossification of the skull and skeleton; (4) pectoral girdle arciferal and robust, with small ovoid fenestra, distant from the epicoracoid; (5) radius and ulna fused; (6) finger IV greatly reduced, almost not visible externally; (7) manus with two prepollical elements; (8) tips of terminal phalangeal elements of fingers I and IV pointed, II and III arrow-shaped; (9) tibiale and fibulare completely fused; (10) toes I and V present but externally indistinguishable, toe II greatly reduced, toe III short and distinct, and toe IV larger and robust; (11) pes with distal tarsal element I present; (12) pes with phalangeal formula 1���2���3���4���0; (13) tips of terminal phalangeal elements of toes I and II pointed, and of toes III and IV arrow-shaped; (14) arytenoid cartilages not mineralized; (15) body size (SVL of adults: 9.2���10.8 mm for males and 11.1���12.4 mm for females); (16) proportional measurements HL/SVL 18���24% and ED/HL 52���73%; (17) rounded snout in dorsal and lateral views; (18) protuberant nostrils; (19) skin texture rough; and (20) advertisement call short (0.02���0.03 seconds), composed of one high-frequency note (dominant frequency 6.6���7.3 kHz). Comparisons with other species. Brachycephalus actaeus sp. nov. exhibits a ���bufoniform��� body and an orange background color, both characteristics that clearly differentiate it from the ���leptodactyliform��� species: B. didactylus, B. hermogenesi, B. pulex, and B. sulfuratus (Izecksohn 1971; Giaretta & Sawaya 1998; Napoli et al. 2011; Condez et al. 2016). These species are generally smaller, exhibit ���leptodactyliform��� bodies, and always exhibit a brown background color (Napoli et al. 2011; Condez et al. 2016). The absence of hyperossification of the skull and skeleton distinguishes Brachycephalus actaeus sp. nov. from B. darkside, B. ephippium, B. garbeanus, and B. margaritatus, which exhibit the extreme condition of hyperossification within Brachycephalus, including a dorsal bony shield (Clemente-Carvalho et al. 2009; Guimar��es et al. 2017). The absence of hyperossification also distinguishes the new species from B. alipioi, B. atelopoide, B. bufonoides, B. crispus, B. guarani, B. nodoterga, B. pitanga, B. toby, and B. vertebralis, which exhibit the intermediate condition of hyperossification within Brachycephalus (Clemente-Carvalho et al. 2009; Haddad et al. 2010; Pombal 2010; Clemente-Carvalho et al. 2011; Condez et al. 2014; Condez et al. 2016). Brachycephalus actaeus sp. nov. differs from B. brunneus, B. coloratus, B. ferruginus, B. izecksohni, and B. pombali, which possess pectoral girdles with larger fenestra, disposed closer to the epicoracoid (small fenestra distant from the epicoracoid in the new species). Likewise, B. brunneus, B. ferruginus, B. izecksohni, and B. pombali differ in having the radius and ulna not fused, pes with distal tarsal element I absent, and toe V reduced (Ribeiro et al. 2005; Alves et al. 2006); the new species has the radius and ulna fused, pes with distal tarsal element I present, and toe V externally not visible. Also, B. albolineatus , B. coloratus, B. curupira, B. ferruginus, and B. pombali have just one prepollical element, and B. izecksohni has no prepollical element (Ribeiro et al. 2005; Alves et al. 2006; Bornschein et al. 2016b; Ribeiro et al. 2017); the new species has two prepollical elements. The phalangeal formula for the pes of B. brunneus , B. izecksohni, and B. pombali is 0���2���3���4���0 (Ribeiro et al. 2005; Alves et al. 2006), and for B. curupira it is 0���1���3���4���0 (Ribeiro et al. 2017); the phalangeal formula of the pes of the new species is 1���2���3���4���0. In B. albolineatus, B. coloratus, and B. curupira the fibulare and tibiale are not completely fused; tips of terminal phalangeal elements of toes II���IV are arrow-shaped (fibulare and tibiale completely fused; tips of terminal phalangeal elements of toes I and II pointed, III and IV arrow-shaped in the new species). Finally, the arytenoid cartilages are mineralized in B. albolineatus and B. coloratus (arytenoid cartilages not mineralized in the new species). Although the osteology of B. pernix was not studied in detail (Pombal et al. 1998), on the basis of the available information its osteology is quite similar to that of the new species. See Figure 3 for osteological details of the new species. Body size (males SVL = 9.2���10.8 mm; females SVL = 11.1���12.4 mm) distinguishes the new species from Brachycephalus ferruginus (males SVL = 11.6���12.5 mm; females SVL = 13.0��� 14.5 mm; Alves et al. 2006), B. pernix (males SVL = 12.0��� 13.3 mm; females SVL = 14.1���15.8 mm; Pombal et al. 1998), and B. pombali (males SVL = 12.6���13.9 mm; females SVL= 14.6���15.3 mm; Alves et al. 2006). Also, in B. coloratus (males SVL = 10.3��� 10.6 mm; females SVL = 12.2���13.3 mm; Ribeiro et al. 2017), B. izecksohni (males SVL = 10.3���12.1 mm; females SVL = 12.5���13.1 mm; Ribeiro et al. 2005), and B. tridactylus (males SVL = 10.6���11.6; females SVL = 13.5���13.8 mm; Garey et al. 2012) males and/or females are slightly larger than in the new species. Brachycephalus actaeus sp. nov. is distinguishable from B. albolineatus, B. auroguttatus, B. boticario, B. fuscolineatus, B. leopardus, B. mariaeterezae, B. olivaceus, B. quiririensis, and B. verrucosus by a proportionally shorter head relative to body length (HL/SVL) and by a proportionally larger eye diameter related to head length (ED/HL). In B. actaeus sp. nov., HL/ SVL is 18���24% (x��= 20, SD = 1) and ED/HL is 52���73% (x��= 63, SD = 4), for the 32 adult specimens of the type series, without sexual distinction. In B. albolineatus, HL/SVL is 28���34% (x��= 31, SD = 4) and ED/HL is 36���42% (x��= 38, SD = 4); in B. auroguttatus, HL/ SVL is 29���38% (x��= 33, SD = 2) and ED/HL is 30���44% (x��= 33, SD = 3); in B. boticario, HL/ SVL is 31���36% (x��= 34, SD = 2) and ED/HL is 30���34% (x��= 32, SD = 1); in B. fuscolineatus, HL/ SVL is 29���34% (x��= 31, SD = 1) and ED/HL is 36���41% (x��= 39, SD = 1); in B. leopardus, HL/ SVL is 31���35% (x��= 33, SD = 1) and ED/HL is 34���43% (x��= 38, SD = 3); in B. mariaeterezae, HL/ SVL is 29���36% (x��= 33, SD = 2) and ED/HL is 36���42% (x��= 39, SD = 2); in B. olivaceus, HL/ SVL is 32���36% (x��= 34, SD = 1) and ED/HL is 26���36% (x��= 32, SD = 3); in B. quiririensis, HL/ SVL is 31���36% (x��= 34, SD = 1) and ED/HL is 28���34% (x��= 32, SD = 2); and in B. verrucosus, HL/ SVL is 30���36% (x��= 33, SD = 2) and ED/HL is 30���40% (x��= 34, SD = 3) (Pie & Ribeiro 2015; Ribeiro et al. 2015; Bornschein et al. 2016b). Additionally, Brachycephalus actaeus sp. nov. exhibits a rounded snout in dorsal and lateral views, which distinguishes it from B. brunneus, which has a slightly mucronate snout in dorsal view (Ribeiro et al. 2005); from B. leopardus, which has a slightly truncate snout in dorsal and lateral views (Ribeiro et al. 2015); and from B. quiririensis, which has a mucronate snout in dorsal view (Pie & Ribeiro 2015). Nostrils are not protuberant in B. auroguttatus, B. pernix, and B. fuscolineatus (Pombal et al. 1998; Ribeiro et al. 2015), which differ from the protuberant nostrils of the new species. In Brachycephalus tridactylus, finger IV is not externally visible (Garey et al. 2012), while in the new species it is reduced but externally distinct. In B. actaeus sp. nov., fingertips I, II, and IV are rounded, differing from B. pernix in which these fingertips are pointed (Pombal et al. 1998) and from B. brunneus, B. izecksohni, and B. leopardus, which have the tip of finger II pointed (Ribeiro et al. 2005; Ribeiro et al. 2015). The texture of the skin on the dorsum of Brachycephalus actaeus sp. nov. is rough; this characteristic distinguishes the new species from B. albolineatus, B. brunneus, B. coloratus, B. curupira, B. ferruginus, B. izecksohni, B. leopardus, B. pernix, B. pombali, and B. tridactylus, which have a smooth dorsum (Pombal et al. 1998; Ribeiro et al. 2005; Alves et al. 2006; Garey et al. 2012; Ribeiro et al. 2015; Bornschein et al. 2016b; Ribeiro et al. 2017). The dark green general color of Brachycephalus actaeus sp. nov. in life is very distinct from B. boticario, B. coloratus, B. ferruginus, B. fuscolineatus, B. izecksohni, B. leopardus, B. mariaeterezae, B. quiririensis, B. pernix, B. pombali, B. tridactylus, and B. verrucosus, which exhibit a bright yellow or orange general dorsal body color (Pombal et al. 1998; Ribeiro et al. 2005; Alves et al. 2006; Garey et al. 2012; Pie & Ribeiro 2015; Ribeiro et al. 2015; Ribeiro et al. 2017). The body color of the new species is similar to B. albolineatus, which has a greenish general coloration (Bornschein et al. 2016b), and B. olivaceus, which is predominantly dark green to brown (Ribeiro et al. 2015). The advertisement call of Brachycephalus actaeus sp. nov. differs in structure and frequency from all known advertisement calls within Brachycephalus. It is clearly distinct from B. crispus, B. darkside , B. ephippium, and B. pitanga by having higher frequencies together with a different call structure (Pombal et al. 1994; Ara��jo et al. 2012; Condez et al. 2014; Guimar��es et al. 2017). In these species, the advertisement call is characterized by the regular repetition of one low-frequency note with several pulses, while in B. actaeus sp. nov. the high-frequency notes are composed of just two pulses. In B. crispus, the notes generally last 0.28 seconds and are composed of 10 pulses; the frequency range is 3.5���5.7 kHz, while the dominant frequency is 4.6 kHz (Condez et al. 2014). In B. darkside, the average note duration is 0.11 seconds and notes are composed of six pulses; the frequency range is 2.5���5.8 kHz [considered dominant frequency in Guimar��es et al. (2017)], while the dominant frequency is 3.4 kHz [considered peak frequency in Guimar��es et al. (2017)]. In B. ephippium, notes typically last 0.12 seconds and are also composed of 12 pulses; the minimum and maximum frequencies are 3.4���5.3 kHz (Pombal et al. 1994). In B. pitanga, notes last 0.17 seconds and are composed of 11 pulses; the dominant frequency is 4.9 kHz (Ara��jo et al. 2012). The high frequency of the advertisement call of the new species is comparable to that described for B. hermogenesi and B. sulfuratus, the latter having the highest dominant frequency known for the genus (Condez et al. 2016). Nevertheless, the advertisement calls of these species differ from the new species in general structure, which is long and composed of a set of high-frequency notes. In B. hermogenesi, the call is composed of 1���5 notes; call lasts 0.2 seconds, with 1���5 pulses (Verdade et al. 2008). The dominant frequency in B. hermogenesi is 6.8 kHz (Verdade et al. 2008). In B. sulfuratus, the call is composed of 4���7 notes, each one lasting 0.19 seconds, with 9 pulses (Condez et al. 2016). The frequency range is 4.9���9.3 kHz and the dominant frequency is 6.7 kHz (Condez et al. 2016). When compared to the advertisement calls of its most closely related species, B. pernix and B. tridactylus, the new species call has shorter notes and higher frequencies. In B. pernix, notes last 0.03���0.06 seconds and are composed of three pulses; the frequency range is 4.5 kHz���6.7 kHz (Wistuba 1998). In B. tridactylus, the frequency range is 3.2���6.4 kHz and dominant frequency is 4.8 kHz (Garey et al. 2012). Description of holotype. Body robust, bufoniform; head wider than long; head length 19% of SVL; snout short, rounded in lateral and dorsal views (Figures 1A and 1B); nostrils protuberant; canthus rostralis indistinct; loreal region slightly concave; eyes slightly protruding laterally and dorsally; eye diameter 72% of head length; tympanum absent; lips nearly sigmoid; vocal sac not expanded externally; vocal slits present; tongue longer than wide, with the posterior half not adherent to floor of mouth; vomerine teeth absent; choanae small and ovoid, anterior to eyes. Arm and forearm moderately slender; hands with fingers I and IV reduced; finger II short but distinct; finger III large and robust; fingertips I, II, and IV rounded, fingertip III pointed; finger lengths IV Measurements of holotype (in mm). SVL 9.4; HL 1.8; HW 3.0; ND 0.4; IND 1.1; ED 1.3; IOD 2.1; END 0.6; AL 2.0; FAL 2.1; HAL 1.4; THL 3.8; TBL 3.3; FL 4.6. Color in life. (Figure 6) Iris black. General body color orange, dorsal surface of body covered by dark green blotches; in dorsal view, a poorly defined dark brown stripe extends from the interorbital region to the posterior end of the vertebral column; arms, legs, fingers III and IV and toe IV dark green, other fing, Published as part of Monteiro, Juliane Petry De Carli, Condez, Thais Helena, Garcia, Paulo Christiano De Anchietta, Comitti, Estev��o Jasper, Amaral, Ivan Borel & Haddad, C��lio Fernando Baptista, 2018, A new species of Brachycephalus (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil, pp. 483-505 in Zootaxa 4407 (4) on pages 486-500, DOI: 10.11646/zootaxa.4407.4.2, http://zenodo.org/record/1221060, {"references":["Izecksohn, E. (1971) Novo genero e nova especie de Brachycephalidae do estado do Rio de Janeiro, Brasil. Boletim do Museu Nacional, Zoologia, 280, 1 - 12.","Giaretta, A. A. & Sawaya, R. J. (1998) Second species of Psyllophryne (Anura: Brachycephalidae). Copeia, 1998, 985 - 987. https: // doi. org / 10.2307 / 1447345","Napoli, M. F., Caramaschi, U., Cruz, C. A. G. & Dias, I. R. (2011) A new species of flea-toad, genus Brachycephalus Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil. Zootaxa, 2739 (1), 33 - 40. https: // doi. org / 10.11646 / zootaxa. 2739.1.3","Condez, T. H., Monteiro, J. P. C., Comitti, E. J., Garcia, P. C. A., Amaral, I. B. & Haddad, C. F. B. (2016) A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa, 4083 (1), 40 - 56. https: // doi. org / 10.11646 / zootaxa. 4083.1.2","Clemente-Carvalho, R. B. G., Antoniazzi, M. M., Jared, C., Haddad, C. F. B., Alves, A. C. R., Rocha, H. S., Pereira, G. R., Oliveira, D. F., Lopes, R. T. & Reis, S. F. (2009) Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): Microscopic structure and macroscopic patterns of variation. Journal of Morphology, 270 (11), 1285 - 1295. https: // doi. org / 10.1002 / jmor. 10755","Guimaraes, C. S., Luz, S., Rocha, P. C. & Feio, R. N. (2017) The dark side of pumpkin toadlet: a new species of Brachycephalus (Anura: Brachycephalidae) from Serra do Brigadeiro, southeastern Brazil. Zootaxa, 4258 (4), 327 - 344. https: // doi. org / 10.11646 / zootaxa. 4258.4.2","Haddad, C. F. B., Alves, A. C. R., Clemente-Carvalho, R. B. G. & Reis, S. F. (2010) A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae). Copeia, 2010, 410 - 420. https: // doi. org / 10.1643 / CH- 09 - 102","Pombal, J. P. (2010) A posicao taxonomica das \" variedades \" de Brachycephalus ephippium (Spix, 1824) descritas por Miranda- Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae). Boletim do Museu Nacional, Zoologia, 526, 1 - 12.","Clemente-Carvalho, R. G. B., Klaczo, J., Perez, I., Alves, A. C. R., Haddad, C. F. B. & Reis, S. F. (2011) Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 61, 79 - 89. https: // doi. org / 10.1016 / j. ympev. 2011.05.017","Condez, T. H., Clemente-Carvalho, R. B. G., Haddad, C. F. B. & Reis, S. F. (2014) A new species of Brachycephalus (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, southeastern Brazil. Herpetologica, 70 (1), 89 - 99. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00044","Ribeiro, L. F., Alves, A. C. R., Haddad, C. F. B. & Reis, S. F. (2005) Two new species of Brachycephalus Gunther, 1985 from the state of Parana, southern Brazil (Amphibia, Anura, Brachycephalidae). Boletim do Museu Nacional, Zoologia, 519, 1 - 18.","Alves, A. C. R., Ribeiro, L. F., Haddad, C. F. B. & Reis, S. F. (2006) Two new species of Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest in Parana state, southern Brazil. Herpetologica, 62 (2), 221 - 233. https: // doi. org / 10.1655 / 05 - 41.1","Bornschein, M. R., Ribeiro, L. F., Blackburn, D. C., Stanley, E. L. & Pie, M. R. (2016 b). A new species of Brachycephalus (Anura: Brachycephalidae) from Santa Catarina, southern Brazil. PeerJ, 4 (e 2629), 1 - 19. https: // doi. org / 10.7717 / peerj. 2629","Ribeiro, L. F., Blackburn, D. C., Stanley, E. L., Pie, M. R. & Bornschein M. R. (2017) Two new species of the Brachycephalus pernix group (Anura: Brachycephalidae) from the state of Parana, southern Brazil. PeerJ, 5 (e 3603), 1 - 28. https: // doi. org / 10.7717 / peerj. 3603","Pombal, J. P., Wistuba, E. M. & Bornschein, M. R. (1998) A new species of brachycephalid (Anura) from the Atlantic Rain Forest of Brazil. Journal of Herpetology, 32 (1), 70 - 74. https: // doi. org / 10.2307 / 1565481","Garey, M. V., Lima, A. M. X., Hartmann, M. T. & Haddad, C. F. B. (2012) A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from southern Brazil. Herpetologica, 68 (2), 266 - 271. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 11 - 00074.1.","Araujo, C. B., Guerra, T. J., Amatuzzi, M. C. O. & Campos, L. A. (2012) Advertisement and territorial calls of Brachycephalus pitanga (Anura: Brachycephalidae). Zootaxa, 3302, 66 - 67.","Verdade, V. K., Rodrigues, M. T., Cassimiro, J., Pavan, D., Liou, N. & Lange, M. (2008) Advertisement call, vocal activity, and geographic distribution of Brachycephalus hermogenesi (Giaretta and Sawaya, 1998) (Anura, Brachycephalidae). Journal of Herpetology, 42 (3), 542 - 549. https: // doi. org / 10.1670 / 07 - 287.1","Goutte, S., Mason, M. J., Christensen-Dalsgaard, J., Montealegre-Z, F., Chivers, B. D., Sarria-S, F. A., Antoniazzi, M. M., Jared, C., Sato, L. A. & Toledo, L. F. (2017) Evidence of auditory insensitivity to vocalization frequencies in two frogs. Scientific Reports, 7, 12121. https: // doi. org / 10.1038 / s 41598 - 017 - 12145 - 5","Pie, M. R., Meyer, A. L. S., Firkowski, C. R., Ribeiro, L. F. & Bornschein, M. R. (2013) Understanding the mechanisms underlying the distribution of microendemic montane frogs (Brachycephalus spp., Terrarana: Brachycephalidae) in the Brazilian Atlantic Rainforest. Ecological Modelling, 250, 165 - 176. https: // doi. org / 10.1016 / j. ecolmodel. 2012.10.019","Bornschein, M. R., Firkowski, C. R., Belmonte-Lopes, R., Correa, L., Ribeiro, L. F., Morato, S. A. A., Antoniazzi-Junior, R. L., Reinert, B. L., Meyer, A. L. S., Cini, F. A. & Pie, M. R. (2016 a). Geographical and altitudinal distribution of Brachycephalus (Anura: Brachycephalidae) endemic to the Brazilian Atlantic Rainforest. PeerJ, 4 (e 2490), 1 - 41. https: // doi. org / 10.7717 / peerj. 2490","Oliveira, J. C. F., Coco, L., Pagotto, R. V., Pralon, E., Vrcibradic, D., Pombal, J. P. & Rocha, C. F. D. (2012) Amphibia, Anura, Brachycephalus didactylus (Izecksohn, 1971) and Zachaenus parvulus (Girard, 1853): Distribution extension. Check List, 8 (2), 242 - 244. https: // doi. org / 10.15560 / 8.2.242","Clemente-Carvalho, R. B. G., Monteiro, L., Bonato, V., Rocha, H., Pereira, G., Oliveira, D., Lopes, R., Haddad, C. F. B., Martins, E. & Reis, S. F. (2008) Geographic variation in cranial shape in the pumpkin toadlet (Brachycephalus ephippium): a geometric analysis. Journal of Herpetology, 42 (1), 176 - 185. https: // doi. org / 10.1670 / 07 - 141 R 1.1","Pimenta, B. V. S., Bernils, R. S. & Pombal, J. P. (2007) Amphibia, Anura, Brachycephalidae, Brachycephalus hermogenesi: filling gap and geographic distribution map. Check List, 3 (3), 277 - 279. https: // doi. org / 10.15560 / 3.3.277","Pombal, J. P. & Izecksohn, E. (2011) Uma nova especie de Brachycephalus (Anura: Brachycephalidae) do estado do Rio de Janeiro. Papeis Avulsos de Zoologia, 51 (28), 443 - 451. https: // doi. org / 10.1590 / S 0031 - 10492011002800001","Clemente-Carvalho, R. B. G., Perez, S. I., Tonhati, C. H., Condez, T. H., Sawaya, R. J., Haddad, C. F. B. & Reis, S. F. (2015) Boundaries of morphological and molecular variation and the distribution range of a miniaturized froglet, Brachycephalus nodoterga (Anura: Brachycephalidae). Journal of Herpetology, 50 (1), 169 - 178. https: // doi. org / 10.1670 / 14 - 119","Abegg, A. D., Ortiz, F. R., Rocha, B. & Condez, T. H. (2015) A new record for Brachycephalus nodoterga (Amphibia, Anura, Brachycephalidae) in the state of Sao Paulo. Brazil. Check List, 11 (1769), 1 - 3. https: // doi. org / 10.15560 / 11.5.1769"]}

Scinax caissara sp. nov. (Figs. 1���2) Holotype. CFBH 19412, an adult male collected in Escola Agr��cola Engenheiro Agr��nomo Narciso de Medeiros (24�� 40' 20.88" S, 47�� 32' 48.22" W; sea level), Municipality of Iguape, State of S��o Paulo, Brazil, 7 July, 2007, by Juliana Zina. Paratypes. All specimens were collected in the State of S��o Paulo: CFBH 19411 (adult female) collected with the holotype; CFBH 17051 (adult male) collected in Municipality of Iguape; CFBH 17079, 17087, 18523���18526, 18930, 19094, 19095, 19097, and 19398���19400 (adult males), CFBH 18522 and 19401 (adult females) collected in Municipality of Pariquera-A��u; CFBH 17864 and 19413 (adult males) collected in Municipality of Ilha Comprida; CFBH 18916���18917, 19058, 19059, 19402-19408, and 19410 (adult males) and CFBH 19409 (adult female) collected in Municipality of Canan��ia. All paratypes were collected by Juliana Zina, between July 2007 and January 2008. Referred specimens. CFBH 19545 and 19546 collected in Municipality of Canan��ia, State of S��o Paulo, Brazil, 12 January, 2008, by Juliana Zina. Diagnosis and comparison with other species. The new species is assigned to the genus Scinax based on the identification of the morphological synapomorphies suggested by Faivovich (2002): webbing between Toes I and II that does not extend beyond the subarticular tubercle of Toe I, origin of the m. pectoralis abdominalis at well defined tendons, ability to bend backwards Finger II and Toe I, and m. pectoralis abdominalis overlapping m. obliqus externus. Moreover, S. caissara is assigned to the S. catharinae species group due to the laterodistal origin of the m. extensores brevis distalis digit III, phenotypic synapomorphy pointed out by Faivovich (2002). This new species is characterized by small size (SVL of males 18.8���23.2, n = 26; SVL of females 20.1���27.5, n = 5); subovoid snout in dorsal view; canthus rostralis marked; loreal region concave and oblique; vocal slits present in males; males with vocal sac not expanded; presence of glandular acini on the mental region (Fig. 3); absence of macroscopic glandular acini on the pectoral region; pectoral fold absent; absence of macroscopic glandular acini on the medial region of the forearm; presence of macroscopic glandular acini on the dorsal region of the Fingers II and III; nuptial pad with the glandular acini diffuse over the skin and not cohesive, not forming an elevated structure on the skin; interocular region with a w-shape blotch that exceeds the posterior margin of the eyes; skin on dorsum smooth; belly and gular region cream with irregular lines and scattered dots; absence of bright coloration on inguinal region and hidden portions of the thigh and shank; presence of inguinal gland (Fig. 4); the foot webbing reaches the second subarticular tubercles or at most a half of the penultimate phalanx of Toe V. Scinax caissara is distinguishable from all species of Scinax for having glandular acini on mental region. Further, the presence of an inguinal gland distinguishes the new species from S. centralis (Pombal & Bastos, 1996) ���that has a particularly hypertrophied one���and from S. agilis (Cruz & Peixoto, 1983 "1982"), S. albicans (Bokermann, 1967), S. angrensis (Lutz, 1973), S. argyreornatus (Miranda-Ribeiro, 1926), S. aromothyella, S. berthae, S. carnevallii (Caramaschi & Kisteumacher, 1982), S. heyeri (Weygoldt, 1986), S. humilis (Lutz, 1954), S. kautskyi (Carvalho-e-Silva & Peixoto, 1991), S. littoralis (Pombal & Gordo, 1991), S. machadoi (Bokermann & Sazima, 1973), S. muriciensis Cruz, Nunes & Lima, 2011, S. pombali Louren��o, Carvalho, Ba��ta, Pezzuti & Leite, 2013, S. skuki Lima, Cruz & Azevedo, 2011, S. strigilatus (Spix, 1824), and S. trapicheiroi (Lutz, 1954) that do not have inguinal gland. Scinax caissara differs from S. angrensis, S. ariadne (Bokermann, 1967), S. brieni (De Witte, 1930), S. catharinae (Boulenger, 1888), S. humilis, S. jureia (Pombal & Gordo, 1991), S. kautskyi, S. littoralis, S. muriciensis, S. obtriangulatus (Lutz, 1973), S. skaios Pombal, Carvalho, Canelas & Bastos, 2010, S. strigilatus, and S. trapicheiroi by the smaller size of males and females (SVL of males in S. caissara 18.8���23.2; combined SVL of males in these species 23.6���42.8; SVL of female in S. caissara 20.9���27.5; combined SVL of females in these species 28.7���47.0); from S. agilis, S. argyreornatus, S. melanodactylus Louren��o, Luna & Pombal, 2014, and S. skuki by the larger size of males (combined SVL of males in these species 12.0���18.4) and from S. agilis and S. melanodactylus by the larger size of females (combined SVL of females in these species 13.2���18.9); from S. albicans, S. canastrensis (Cardoso & Haddad, 1982), S. carnevallii, S. flavoguttatus (Lutz & Lutz, 1939), S. hiemalis (Haddad & Pombal, 1987), S. pombali, and S. tripui Louren��o, Nascimento & Pires, 2009 by smaller size of females (combined SVL of females in these species 28.3���47.0). The new species differs from S. agilis, S. albicans, S. angrensis, S. aromothyella, S. carnevallii, S. catharinae, S. brieni, S. flavoguttatus, S. humilis, S. littoralis, S. luizotavioi (Caramaschi & Kisteumacher, 1989), S. obtriangulatus, S. rizibilis (Bokermann, 1964), S. trapicheiroi, and S. tripui in having nuptial pad where the glandular acini are diffuse over the skin and not cohesive, not forming an elevated structure on the skin (in these species the nuptial pad is an elevated structure with respect to the surrounding skin, and particularly enlarged in S. rizibilis). The new species differ from most species of the S. catharinae group except S. agilis, S. canastrensis, S. centralis, S. longilineus (Lutz, 1968), S. machadoi, S. melanodactylus, S. pombali, S. rizibilis, and S. skaios by its subovoid snout in dorsal view (subelliptical in S. aromothyella and S. berthae; rounded in S. ariadne, S. catharinae, S. brieni, S. obtriangulatus, and S. ranki; rounded with a mucronate tip in S. albicans, S. angrensis, S. flavoguttatus, S. hiemalis, S. heyeri, S. humilis, S. littoralis, S. muriciensis, S. strigilatus, S. trapicheiroi, and S. tripui; mucronate in S. carnevallii and S. kautskyi; sub-elliptical with acute tip in S. jureia and S. luizotavioi; mucronate or subelliptical in S. argyreornatus and S. skuki). Scinax caissara differs from S. agilis, S. albicans, S. argyreornatus, S. aromothyella, S. berthae, S. machadoi, S. melanodactylus, S. ranki, and S. rizibilis in having well marked canthus rostralis (canthus rostralis not well marked in these species). The presence of vocal slits differentiates the new species from S. ariadne and S. skaios (vocal slits absent). The new species is distinguished from S. aromothyella, S. berthae, and S. rizibilis by its vocal sac not expanded externally (notably expanded in these species). The new species differs from S. canastrensis, S. carnevallii, S. flavoguttatus, S. kautskyi, S. longilineus, S. machadoi, S. muriciensis, S. skaios, S. strigilatus, and S. tripui by the absence of glandular acini on the medial region of forearm (glandular acini present on the medial region of forearm in these species). The presence of glandular acini on the dorsal region of the Finger III in males distinguishes the new species from S. agilis, S. argyreornatus, S. ariadne, S. aromothyella, S. berthae, S. centralis, S. heyeri, S. hiemalis, S. jureia, S. melanodactylus, S. muriciensis, S. pombali, S. ranki, S. rizibilis, S. skaios, S. skuki, and S. strigilatus (glandular acini absent on Finger III in these species). Scinax caissara differs from most species of the S. catharinae group except S. aromothyella and S. berthae, in having smooth skin on dorsum (rough in S. ariadne, S. canastrensis, S. longilineus, S. pombali, and S. skaios; covered by scattered tubercles in the remaining species). Some species of the S. catharinae group have more developed foot webbing, reaching the base of the adhesive disc on the medial margin of the Toe V. This is the case of S. argyreornatus, S. aromothyella, S. catharinae, S. kautskyi, S. flavoguttatus, S. heyeri, S. luizotavioi, S. muriciensis, S. skuki, S. strigilatus, and S. tripui. Scinax caissara differs from these species because its interdigital webbing is less developed, reaching only the second subarticular tubercles or at most a half of the penultimate phalanx of Toe V. Scinax caissara differs from most species of the S. catharinae group except S. albicans, S. argyreornatus, S. carnevallii, and S. kautskyi in having a interocular W-shaped blotch that does not exceeds the posterior margin of eyes (S. flavoguttatus, S. heyeri, S. machadoi, S. muriciensis, S. rizibilis, S. strigilatus, and S. tripui, have a Wshaped blotch on the interocular region that exceeds the posterior margin of the eyes; S. agilis and S. melanodactylus have a black longitudinal line on the interocular region or they have no mark on this region; S. aromothyella and S. berthae have an inverted triangle or an irregular band on this region that does not exceeds the posterior margin of eyes; S. ariadne, S. brieni and S. catharinae have an inverted triangle or an irregular trapezoid mark on this region that exceeds the posterior border of the tympanum; S. canastrensis, S. centralis, S. hiemalis, S. jureia, S. longilineus, S. luizotavioi, S. ranki, and S. skaios have a inverted triangle on this region that exceeds the posterior border of eyes; S. angrensis, S. humilis, and S. littoralis have a complete or interrupted band on this region, which is narrow and restricted to the median interocular region; S. pombali and S. obtriangulatus have an inverted triangle on this region that exceeds the posterior border of the tympanum; S. trapicheiroi has a W-shaped blotch, which is very long, extending far beyond the interocular region, reaching half of the body). The cream belly with scattered and irregular lines and dots differentiates the new species from S. agilis, S. albicans, S. angrensis, S. ariadne, S. argyreornatus, S. aromothyella, S. berthae, S. canastrensis, S. carnevallii, S. centralis, S. flavoguttatus, S. heyeri, S. hiemalis, S. humilis, S. kautskyi, S. littoralis, S. longilineus, S. luizotavioi, S. machadoi, S. melanodactylus, S. muriciensis, S. obtriangulatus, S. pombali, S. ranki, S. rizibilis, S. skaios, S. skuki, S. strigilatus, S. trapicheiroi, and S. tripui (S. ariadne, S. flavoguttatus, and S. tripui have irregular cream spots on brown background in this region; the remaining species have fully uniform cream belly). The new species has no flash color on the inguinal region and hidden areas of the thigh and shank, unlike S. ariadne (light brown irregular blotches on violet or pink background; personal observation), S. aromothyella (dark yellow; Faivovich 2005), S. berthae (���irregular yellow-orange spots���; Barrio 1962), S. brieni (���pale bluish color on the concealed areas���; Lutz 1973), S. canastrensis (���yellow���; Cardoso & Haddad 1982), S. catharinae, S. humilis, and S. trapicheiroi (dark brown blotches on light blue or white background; personal observation), S. centralis (���yellow on dark brown background���; Pombal & Bastos 1996), S. flavoguttatus and S. heyeri (brown blotches on orange background; personal observation), S. hiemalis (���black blotches on green background���; Haddad & Pombal 1987), S. longilineus and S. machadoi (vermiculate spots on yellow or pale background; personal observation), S. obtriangulatus (���dull grayish violet���; Lutz 1973), S. pombali (irregular brown blotches on yellow background; personal observation), S. ranki (dark brown blotches on greenish background; personal observation), S. skaios (vermiculated dark brown spots on light green background; personal observation), S. strigilatus (���concealed surfaces of flanks and thighs greenish���; Pimenta et al. 2007), and S. tripui (irregular brown blotches on light green background; personal observation). Description of the holotype. SVL 19 mm. Head longer than wider (39 % of SVL). Snout is subovoid in dorsal view, protruding in profile. Nostril elliptical, located laterally, immediately before the tip of snout, opening directed dorso���laterally. Canthus rostralis marked. Loreal region oblique and concave. Eye diameter 37.4% of head width. Interorbital and internostril distance 25% and 34.1% of head width, respectively. Tympanum rounded, annulus tympanicus well defined, its diameter 45.9% of eye diameter. Curved supratympanic fold, evident, extending from posterior corner of eye to the arm insertion. Tongue large, elongated, unattached on the posterior and lateral borders. Vocal slits present, originating on the side of the tongue and running to the posterolateral corner of the mouth. Choanae elliptical. Vomerine teeth in two contiguous convex series of three teeth each, positioned between choanae. Vocal sac not expanded externally. Forearms longer than arms and enlarged. Outer margin of forearm smooth. Outer metacarpal tubercle simple and elliptical. Inner metacarpal tubercle single and elliptical. Subarticular tubercles single and rounded. Supernumerary tubercles small and rounded. Dorsal region of Fingers II and III with macroscopic glandular acini. Webbing absent between Fingers II and III and basal between other fingers. Discs on fingers elliptical, wider than long. Relative finger length IIMeasurements of the holotype. SVL 19.0; HW 6.5; HL 7.4; IND 1.4; ED 2.1; END 2.9; IOD 2.9; TD 1.2; THL 9.7; TL 10.5; FL 8.3. Color of holotype in preservative. Overall dorsal coloration brown. The upper lip is brown with a cream blotch bellow the eye. Acanthal brown line. Interocular surface with a brown blotch, bordered by a cream anterior line and with a W-shaped mark that does not exceed the posterior margin of eyes. Iris is gray. Dorsolateral region has a brown stripe that originates on the posterior margin of the eye and reaches the inguinal region. Dorsally hindlimbs are cream with brown bars. Hidden areas of thighs and inguinal region with irregular black markings on a pale background. Belly and throat are pale with small brown dots scattered throughout the region. Inguinal and mental glands have light orange color (see discussion). Color in life. (based on the type series observed in the field, including all males and females) Dorsally yellowish beige. The other color patterns are the same seen in preservative, but the overall body color is dimmer and more intense. The inguinal region shows no flash color. Variation among paratypes. Some measurements are shown on Table 1. Males are smaller than females. Females do not have mental and inguinal glands. The webbing formula varies equally among male and female specimens as follows: I���II2-(2)���3 1/ 2III 1 1/2(2, 2-)���3+(3, 3 1/2) IV3 (3+, 3-z, 3 1/2)���1 1/2(2-)V. Vomerine teeth number varies between three and five on both vomers. The specimen CFBH 19410 has a trapezoid blotch on the interocular region, instead of w-shaped mark. Four specimens have irregular spots and longitudinal lines on dorsum. In two males (CFBH 19406���19407) and one female (CFBH 19059) the spots on dorsum are dimmed and almost inconspicuous. Two females (CFBH 18916 and CFBH 19097) and two males (CFBH 18522 and CFBH 19059) have a second longitudinal band on the side of the body. In six males (CFBH 17079, CFBH 17864, CFBH 18526, CFBH 19400, 19408, and CFBH 19413) the glandular acini are present all along lateral region of the body (reaching the tympanum). Species CD (s) DF (kHz) FR (kHz) NN ND (s) NR (N/s) NI (s) Literature and original terms agilis __ 7.4)7.9 __ 1 0.3)0.4 __ __ Nunes et al. (2007), ���note a��� __ 5.6)7.9 __ 13)29 0.01)0.03 __ 0.06)0.1 Nunes et al. (2007), ���note b��� albicans 0.7 3.3)4.1 __ __ 0.03 __ __ Heyer (1980) angrensis 0.2)0.7 2.1)3.7 1.1)5.7 1)7 0.02 �� 0.01 0.07 0.02)0.07 Garey et al. (2012) argyreornatus 2.3 __ __ __ __ __ __ Bokermann (1966) 0.8 5.0)6.5 3.6)9.0 5 0.02)0.04 __ __ Pombal et al. (1995), ���short call��� 10)25 5.0)6.5 3.6)8.0 130)280 0.02)0.1 __ 0.04)0.08 Pombal et al. (1995), ���long call��� aromothyella 1.0)20.7 4.7)5.4 1.5)8.8 2)74 0.04)0.2 1.0)4.4 0.05)0.5 Pereyra et al. (2012), ���short note��� 1.0)20.7 4.8)5.5 1.5)9.1 0)3 0.3)0.6 0)0.4 0.05)0.5 Pereyra et al. (2012), ���trilleđ note��� berthae __ 3.6)5.0 __ __ 0.2 __ 0.2 Barrio (1962), ���common call, Published as part of Louren��o, Ana Carolina C., Zina, Juliana, Catroli, Gaucilene F., Kasahara, Sanae, Faivovich, Julian & Haddad, C��lio F. B., 2016, A new species of the Scinax catharinae group (Anura: Hylidae) from southeastern Brazil, pp. 415-435 in Zootaxa 4154 (4) on pages 417-426, DOI: 10.11646/zootaxa.4154.4.3, http://zenodo.org/record/266515, {"references":["Faivovich, J. (2002) A cladistic analysis of Scinax (Anura: Hylidae). Cladistics, 18, 367 - 393. http: // dx. doi: 10.1111 / j. 1096 - 0031.2002. tb 00157. x","Pombal, J. P. Jr. & Bastos, R. P. (1996) Nova especie de Scinax Wagler, 1830 do Brasil Central (Amphibia, Anura, Hylidae). Boletim do Museu Nacional Nova Serie Zoologia, 371, 1 - 11.","Bokermann, W. C. A. (1967) Dos nuevas especies de Hyla del grupo catharinae. Neotropica, 13 (41), 61 - 66.","Lutz, B. (1973) Brazilian species of Hyla. University of Texas Press, Austin and London, XVIII + 260 pp.","Miranda-Ribeiro, A. (1926) Notas para servirem ao estudo dos Gymnobatrachios (Anura) brasileiros. Arquivos do Museu Nacional, 27, 1 - 227.","Weygoldt, P. (1986) Beobachtungen zur Okologie und Biologie von Froschen an einem neotropischen Bergbach / Observations on the ecology and biology of frogs of a neotropical mountain stream. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie, 113, 429 - 454.","Lutz, B. (1954) Anfibios anuros do Distrito Federal / The frogs of the Federal District of Brazil. Memorias do Instituto Oswaldo Cruz, 52, 155 - 197 (Portuguese), 219 - 238 (English).","Pombal, J. P. Jr. & Gordo, M. (1991) Duas novas especies de Hyla da floresta atlantica no estado de Sao Paulo (Amphibia, Anura). Memorias do Instituto Butantan, 53 (1), 135 - 144.","Bokermann, W. C. A. & Sazima, I. (1973) Anfibios da Serra do Cipo, Minas Gerais, Brasil. 1: duas novas especies de Hyla (Anura, Hylidae). Revista Brasileira de Biologia, 33 (4), 457 - 472.","Cruz, C. A. G., Nunes, I. & Lima, M. G. (2011) A new Scinax Wagler belonging to the S. catharinae clade (Anura: Hylidae) from the state of Alagoas, northeastern Brazil. Zootaxa, 3096, 18 - 26.","Lourenco, A. C. C., Carvalho, A. L. G, Baeta, D., Pezzuti, T. L. & Leite, F. S. F. (2013) A new species of the Scinax catharinae group (Anura: Hylidae) from Serra da Canastra, southeastern State of Minas Gerais, Brazil. Zootaxa, 3613 (6), 573 - 588. http: // dx. doi. org / 10.11646 / zootaxa. 3613.6.4","Lima, M. G., Cruz, C. A. G. & Azevedo, S. M. Jr. (2011) A new species belonging to the Scinax catharinae group from the state of Alagoas, northeastern Brazil (Amphibia, Anura, Hylidae). Boletim do Museu Nacional Nova Serie Zoologia, 529, 1 - 12.","Spix, J. B. (1824) Animalia nova sive species novae Testudinum et Ranarum, quas in itinere per Brasiliam annis MDC- CCXVII - MDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavariae Regis, Typis Franc, Seraph, Hubschmanni, Monachii, XXXIX + 53 pp. http: // dx. doi. org / 10.5962 / bhl. title. 3665","De Witte, G. F. (1930) Liste des reptiles et batraciens recoltes au Bresil par la Mission Massart (1922 - 23) et description de sept nouvelles especes. In: Massart, J. (Ed.), Une Mission Biologique Belge au Bresil (aout 1922 - mai 1923) par Jean Massart, Raymond Bouillene, Paul Ledoux, Paul Brien, Albert Navez, 2, pp. 213 - 230.","Boulenger, G. A. (1888) A list of batrachians from the Province Santa Catharina, Brazil. Annals and Magazine of Natural History, 6, 415 - 417. http: // dx. doi. org / 10.1080 / 00222938809460758","Pombal, J. P. Jr., Carvalho, R. R. Jr., Canelas, M. A. S. & Bastos, R. P. (2010) A new Scinax of the S. catharinae species group from Central Brazil (Amphibia, Anura: Hylidae). Zoologia, 27 (5), 795 - 802. http: // dx. doi. org / 10.1590 / s 1984 - 46702010000500016","Lourenco, A. C. C., Luna, C. & Pombal, J. P. Jr. (2014) A new species of the Scinax catharinae Group (Anura: Hylidae) from Northeastern Brazil. Zootaxa, 3889 (2), 259 - 276. http: // dx. doi. org / 10.11646 / zootaxa. 3889.2.5","Cardoso, A. J. & Haddad, C. F. B. (1982) Nova especie de Hyla da Serra da Canastra (Amphibia, Anura, Hylidae). Revista Brasileira de Biologia, 42 (3), 499 - 503.","Lutz, A. & Lutz, B. (1939) New Hylidae from Brazil / Hylideos novos do Brasil. Anais da Academia Brasileira de Ciencias, 11, 67 - 89.","Haddad, C. F. B. & Pombal, J. P. Jr. (1987) Hyla hiemalis, nova especie do grupo rizibilis do estado de Sao Paulo. Revista Brasileira de Biologia, 47, 127 - 132.","Lourenco, A. C. C., Nascimento, L. B. & Pires, M. R. S. (2009) a A new species of the Scinax catharinae species group (Anura: Hylidae) from Minas Gerais, southeastern Brazil. Herpetologica, 65 (4), 468 - 479. http: // dx. doi. org / 10.1655 / 07 - 088.1","Caramaschi, U. & Kisteumacher, G. (1989) Duas novas especies de Ololygon Fitzinger, 1843, do Sudeste do Brasil (Amphibia, Anura, Hylidae). Boletim do Museu Nacional Nova Serie Zoologia, 327, 1 - 15.","Bokermann, W. C. A. (1964) Uma nova especie de Hyla da Serra do Mar em Sao Paulo (Amphibia, Salientia). Revista Brasileira de Biologia, 24 (4), 429 - 434.","Lutz, B. (1968) New Brazilian forms of Hyla. Pearce-Sellard, 10, 3 - 18.","Barrio, A. (1962) Los Hylidae de Punta Lara, Provincia de Buenos Aires. Observaciones sistematicas, ecologicas y analisis espectrografico del canto. Physis, 23 (65), 129 - 142.","Pimenta, B. V. S., Faivovich, J. & Pombal, J. P. Jr. (2007) On the identity of Hyla strigilata Spix, 1824 (Anura: Hylidae): redescription and neotype designation for a ' ghost' taxon. Zootaxa, 1441, 35 - 49.","Nunes, I., Santiago, R. S. & Junca, F. A. (2007) Advertiment calls of four hylid frogs from the State of Bahia, northeastern Brazil. South American Journal of Herpetology, 2 (2), 89 - 96. http: // dx. doi. org / 10.2994 / 1808 - 9798 (2007) 2 [89: acofhf] 2.0. co; 2","Heyer, W. R. (1980) The calls and taxonomic positions of Hyla giesleri and Ololygon opalina (Amphibia: Anura: Hylidae). Proceedings of the Biological Society of Washington, 93, 655 - 661.","Bokermann, W. C. A. (1966) Notas sobre Hylidae do Espirito Santo (Amphibia, Salientia). Revista Brasileira de Biologia, 26 (1), 29 - 37.","Pombal, J. P. Jr., Haddad, C. F. B. & Kasahara, S. (1995) A new species of Scinax (Anura: Hylidae) from southeastern Brazil, with comments on the genus. Journal of Herpetology, 29 (1), 1 - 6. http: // dx. doi. org / 10.2307 / 1565078","Pereyra, M. O., Borteiro, C., Baldo, D., Kolenc, F. & Conte, C. E. (2012) Advertisement call of the closely related species Scinax aromothyella Faivovich, 2005 and S. berthae (Barrio, 1962), with comments on the complex calls in the S. cathariae group. Herpetological Journal, 22, 133 - 137.","Bastos, R. P., Signorelli, L., Morais, A. R., Costa, T. B., Lima, L. P. & Pombal, J. P. Jr. (2011) Advertisement calls of three anuran species (Amphibia) from the Cerrado, central Brazil. South American Journal of Herpetology, 6 (2), 67 - 72. http: // dx. doi. org / 10.2994 / 057.006.0204"]}

Brachycephalus sulfuratus sp. nov. Figures 1���3 Holotype. CFBH 39137, adult female, collected at Centro de Estudos e Pesquisas Ambientais da Univille, Vila da Gl��ria, Distrito do Sa�� (26 �� 13'39"S, 48 �� 41'31"W, 123 m above sea level, Datum WGS 84), munic��pio de S��o Francisco do Sul, State of Santa Catarina, Brazil, on 14 November 2014, by T.H. Condez, J.P.C. Monteiro, and E.J. Comitti. Paratypes. CFBH 39138, cleared and double-stained adult male, and CFBH 39139, adult male, both collected with the hotolype. CFBH 39140, adult female collected on 11 September 2014, CFBH 39329, adult female, and CFBH 39331, adult male, collected on 31 July 2015, CFBH 39330 and 39332, adult females, collected on 0 1 August 2015, at the same locality as the holotype, by J.P.C. Monteiro and E.C. Nardin. CFBH 39142, adult male, CFBH 39141 and 39144, adult females, CFBH 39143, cleared and double-stained adult male, collected on 0 4 December 2013, by T.H. Condez, J.P.C. Monteiro and E.J. Comitti, UFMG 17954 and 17955, adult females, CFBH 39407���39410 and UFMG 17956 and 17957, adult males, CFBH 39406, juvenile, collected on 28 August 2015, by J.P.C. Monteiro and E.C. Nardin, at Morro do Cantagalo, Vila da Gl��ria, Distrito do Sa�� (26 �� 10'31"S, 48 �� 42'44"W, 161 m above sea level, Datum WGS 84), munic��pio de S��o Francisco do Sul, State of Santa Catarina, Brazil. CFBH 39145���39147, adult females, collected at Castelo dos Bugres, ��rea de Prote����o Ambiental Dona Francisca (26 �� 13'43"S, 49 �� 03'27"W, 835 m above sea level, Datum WGS 84), munic��pio de Joinville, State of Santa Catarina, Brazil, on 29 November 2013, by T.H. Condez, J.P.C. Monteiro, E.J. Comitti, I. Borel, P.D. Pinheiro, and P.C.A. Garcia. CFBH 39148 and 39149, adult males, collected at Parque Estadual da Ilha do Cardoso (25 �� 06'53"S, 47 �� 55'40"W, 385 m above sea level, Datum WGS 84), munic��pio de Canan��ia, State of S��o Paulo, on 19 December 2013, by T.H. Condez, L.N. Bandeira, and S. Pinheiro. CFBH 39150, adult female, collected at Morro do Anhangava (25 �� 22'51"S, 49 �� 01'26"W, 915 m above sea level, Datum WGS 84), munic��pio de Quatro Barras, State of Paran��, on 0 2 February 2012, by T.H. Condez, T.B. Rocha, and P.F. Colas-Rosas. Referred specimens. MZUSP 129855, juvenile, collected at Ilha do Cardoso (non-georeferenced data), munic��pio de Canan��ia, State of S��o Paulo, on 22 January 1979, by an unknown collector. ZUEC 16602, juvenile, collected at ��rea de Prote����o Ambiental de Guaratuba (25��47��� S, 48��54��� W, 291 m above sea level), munic��pio de S��o Jos�� dos Pinhais, State of Paran��, on 18 January 2008, by A.K. Cunha and I. Soares. Diagnosis. Brachycephalus sulfuratus sp. nov. is a new species of flea-toad, distinguished from all its congeners by the following combination of characters: (1) small body size (SVL of adults: 7.4���8.5 mm for males and 9.0��� 10.8 mm for females); (2) ���leptodactyliform��� body; (3) pectoral girdle arciferal and less robust compared to the Brachycephalus species with ���bufoniform��� body; (4) procoracoid and epicoracoid fused with coracoid but separated from the clavicle by a large fenestrae; (5) toe I externally absent; toes II, III, IV, and V distinct; phalanges of toes II and V reduced; (6) skin smooth with no dermal ossifications; (7) in life, general background color brown with small dark-brown spots; skin of throat, chest, arms, and forearms with irregular yellow blotches; in ventral view, cloacal region of alive and preserved specimens surrounded by a dark-brown inverted v-shaped mark outlined with white; (8) advertisement call long, composed of a set of 4���7 high-frequency notes (6.2���7.2 kHz) repeated regularly. Description of holotype. The holotype of Brachycephalus sulfuratus sp. nov. has a ���leptodactyliform��� body (Figure 1 and 2); head wider than long, narrower than body; head length approximately 26% of SVL; snout long, with length slightly short than the eye diameter, rounded in lateral and dorsal views (Figure 3A and 3B); nostrils protuberant, directed anterolaterally; canthus rostralis distinct and straight; loreal region weakly concave; mouth nearly sigmoid; eye slightly protruding in dorsal and lateral views, eye diameter 48% of HL; tympanum absent; tongue longer than wide, posterior half not adherent to floor of mouth; choanae relatively small and round; vomerine odontophores absent. Upper arm slightly shorter than forearm; length of upper arm plus forearm 43% of SVL; hands approximately as long as upper arm; fingers II and III thin and distinct; finger I and IV very small, vestigial; tip of fingers II and III slightly pointed; relative lengths of fingers II Measurements of holotype (in mm). SVL 9.8; HL 2.5; HW 2.7; ND 0.4; IND 1.0; ED 1.2; IOD 1.9; END 0.8; THL 4.3; TBL 4.1; FL 6.3; AL 2.0; FAL 2.3; HAL 1.8. Color of holotype in life. Dorsal coloration grey to brown covered with dark brown and red spots (Figure 1A). The dark brown spots are concentrated on the head and medial portion of the dorsum, in which a large irregular mark is distinguishable. The interorbital area exhibits a dark brown nearly v-shaped mark. Dorsal surfaces of arms and legs exhibit dark brown blotches, which on the thigh and tibia resemble incomplete stripes. Above the cloacal region, in dorsal view, there is a dark brown inverted m-shaped mark. A dark brown stripe extends laterally from the tip of the snout to the flanks and the surface of thigh (Figure 1A). In lateral view, the tip of the snout and posterior regions of maxilla and ocular globe present undefined yellow spots. The maxilla is dark brown with distinct white spots. Ventral surface of body pale-brown, slightly transparent, with small dark brown spots and white blotches (Figure 1B). Some yellow blotches are also spread among the gular region, arms, forearms, and disposed in an inverted v-shape on the chest. In ventral view, the cloacal region is surrounded by an inverted vshaped dark brown mark with white borders. The pupil is black and the iris is golden. Color of holotype in preservative. General background color is pale-brown covered with small dark brown dots (Figure 2); dorsum with dark brown marks on the ocular region, knees, and cloacal region; dark brown stripes on dorsal surfaces of thigh, tibia, and foot; ventral surfaces of hands and feet with dark brown blotches; in ventral view, cloacal region surrounded by an inverted v-shaped dark-brown mark. Osteology. The cleared and double-stained material revealed the following characters in Brachycephalus sulfuratus sp. nov. No hyperossification of the skull or skeleton. Nasals, sphenethmoid, frontoparietals, prootics, and exoccipitals fused. Premaxillae broad, not fused medially; odontoids absent; alary process of premaxillae distinct and separated from the nasal. Maxillae arched in ventral view; odontoids present. Quadratojugal and pterygoid present. Vomer not fused medially, vomerine odontophores absent. Palatine absent. Parasphenoid and sphenethmoid fused and robust. Squamosal elongated in lateral view, zygomatic ramus short and not ornamented. Tympanic annulus absent. Mandible edentate. Pectoral girdle arciferal and less robust than in other Brachycephalus; clavicle, coracoid, and scapula fused and completely ossified; procoracoid and epicoracoid fused with coracoid, but separated from the clavicle by a large fenestrae; suprascapula expanded, anterior half ossified as cleithrum; omosternum present and cartilaginous; sternum absent. Vertebral column composed of eight presacral, non-imbricate vertebrae; hyperossification absent in the spinal processes of vertebrae; all presacral vertebrae with transverse processes; transverse process of sixth presacral vertebra elongated but not ornamented. Humerus slightly shorter than forearm; radius and ulna fused but distinguishable. Manus with distal carpals (I���IV) fused with centrale; radiale and ulnare about the same size; one prepollical element; phalangeal formula 1���2���3���1; tips of terminal phalangeal element of fingers I and IV falciform and tip of finger II and III pointed. Hindlimbs with tibia and fibula fused but distinguishable, forming the tibiafibula; tibiafibula slightly shorter than femur; tibiale and fibulare fused at their distal and proximal ends (medially not fused). Pes with distal tarsal element I, II, III present and IV���V absent; centrale present. One very reduced prehallical element present; phalangeal formula 1���2���3���4���1; tips of terminal phalangeal elements of toes I and V rounded; tips of the terminal phalangeal elements of toes II, III, and IV arrow-shaped. Advertisement call. The advertisement call of the new species is long, composed by the regular repetition of five or six high-frequency pulsed notes (Figure 4). The call lasts 1.5���2.3 seconds (x��=1.8��0.2) and the interval between calls is 3.1���7.4 seconds (x��=5.1��1.4). The call is composed of 4���7 notes (x��=5.3��0.9), repeated in a rate of 0.1���0.3 notes/second (x��=0.2��0.0). Notes last 131���233 milliseconds (x��=195��13) and present 7���11 pulses (x��=8.8��1.3). Pulses last 20���30 milliseconds (x��=23.6��4.8) and are repeated at a rate of 6.1���12.3 pulses/second (x��=9.3��1.8). The minimum frequency is 4.5���5.5 kHz (x��=4.9��0.3), the maximum frequency is 8.2���10.3 kHz (x��=9.3��0.3), and the dominant frequency is 6.2���7.2 kHz (x��=6.7��0.3). Variation. Morphometric variation is given in Table 1. In our sample, females are larger than males (SVL of females x��= 9.9 mm ��0.4; SVL of males x��=8.0 mm��0.4, Welch���s t-test t = 10.2, df = 9.6, p p F = 12.86, TukeyHSD tests p = 0.02 and p p F = 31, TukeyHSD tests p B. sulfuratus sp. nov. is preserved and the range of divergent call parameters overlapped among the analyzed populations (Table 2). Call (s) Interval (s) Notes/call Notes/s Note (ms) Pulses/ note A 1.7��0.1 4.5��1.7 4.9��0.4 0.2��0.1 177��12 8.25��0.8 (1.5���1.8) (3.1���7.4) (4���6) (0.2���0.3) (131���205) (7���9) B 2.2��0.1 5.9��0.8 5.6��0.6 0.2��0.1 202��14 8.5��1.5 (2.1���2.3) (5.2���6.7) (5���7) (0.1���0.2) (180���233) (7���10) C 1.8��0.1 5.2��1.2 5.1��0.5 0.2��0.1 206��10 9.4��1.2 (1.7���2.1) (3.7���7.0) (4���6) (0.1���0.2) (171���228) (8���11) continued. Pulses/s Pulse (ms) Min Freq (Hz) Max Freq (Hz) Dom Freq (Hz) A 9.1��0.9 25��5 4.9��0.3 9.1��0.2 6.6��0.1 (8.5���10.7) (20���30) (4.5���5.2) (9.0���9.4) (6.5���6.7) B 7.6��1.4 20��0 5.1��0.4 8.2��0.0 6.4��0.2 (6.1���9.0) (20���20) (4.7���5.5) (8.2���8.2) (6.2���6.6) C 10.1��1.9 24��5 5.0��0.3 9.9��0.3 6.9��0.2 (7.7���12.3) (20���30) (4.5���5.4) (9.4���10.3) (6.7���7.2) Comparisons with other species. Brachycephalus sulfuratus sp. nov. shares the diminutive size and reduced number of functional toes with other species of Brachycephalus. The texture of the skin of the head and dorsum of B. sulfuratus sp. nov. is smooth, and without dermal ossification. This characteristic distinguishes the new species from B. ephippium, B. garbeanus, and B. margaritatus, all of which have the largest SVLs in the genus (pers. observation) and the most extreme condition of hyperossification, as revealed by the presence of a dorsal bony shield (Clemente-Carvalho et al. 2009). Brachycephalus sulfuratus sp. nov. also differs from B. alipioi, B. atelopoide, B. bufonoides, B. crispus, B. guarani, B. nodoterga, B. pitanga, B. vertebralis and B. toby, species with intermediate condition of hyperossification of the skull and skeleton (Clemente-Carvalho et al. 2009; Haddad et al. 2010; Pombal 2010; Clemente-Carvalho et al. 2012; Condez et al. 2014). The lack of hyperossification is shared among the new species and B. auroguttatus, B. boticario, B. brunneus, B. didactylus, B. ferruginus, B. fuscolineatus, B. hermogenesi, B. izecksohni, B. leopardus, B. mariaterezae, B. olivaceus, B. pernix, B. pombali, B. pulex, B. quiririensis, B. tridactylus, and B. verrucosus (Clemente-Carvalho et al. 2009; Napoli et al. 2011; Ribeiro et al. 2015; Pie & Ribeiro 2015). Except for B. didactylus, B. hermogenesi, B. pulex, and B. sulfuratus sp. nov., all of these species have a ���bufoniform��� body and in most of them the body background color is orange. The fleatoads, B. didactylus, B. hermogenesi, B. pulex, and B. sulfuratus sp. nov., have the smallest SVLs within the genus, a ���leptodactyliform��� body, and a brown general background color (pers. observation). The main morphological differences among these species are related to the loss of phalangeal elements and the reduced number of toes. Brachycephalus sulfuratus sp. nov. has the toe I externally absent and toes II, III, IV, and V distinct and functional. This condition is very distinct from B. pulex, which presents toes I, II, and V absent, and toes III and IV distinct and functional (Napoli et al. 2011). Brachycephalus didactylus is easily distinguished from B. sulfuratus sp. nov. by having toe I absent, toes II, III, and IV distinct and functional, and toe V vestigial (Izecksohn 1971). Toes of B. hermogenesi have the same configuration as B. sulfuratus sp. nov. (see Giaretta & Sawaya 1998). According to the original description of B. hermogenesi the tip of toe II is pointed, which could be considered distinct from the new species. However, after analyzing several individuals of both species we conclude that this character is variable (being rounded or slightly pointed) and not informative. The general color of Brachycephalus sulfuratus sp. nov. in life and preservative is very distinct from all currently known species of Brachycephalus, except for the flea-toads B. didactylus, B. hermogenesi, and B. pulex. These species exhibit a brown general body color with variable dorsal and ventral patterns of dark ornamentation (Izecksohn 1971; Giaretta & Sawaya 1998; Napoli et al. 2011). The m-shaped mark around the cloacal opening in dorsal view, the ventral inverted v-shaped mark in the chest, and variable patterns of stripes on the legs are shared among the four species. The new species differs from B. didactylus, B. hermogenesi, and B. pulex by having (in life) yellow blotches on the ventral surfaces of the throat, chest, arms, and forearms. Another small differences among the flea-toads relate to the x-shaped dorsal mark, which is diagnostic for B. pulex (Napoli et al. 2011) and can be slightly visible in B. didactylus (Izecksohn 1971) and some B. hermogenesi specimens (Giaretta & Sawaya 1998). In the specimens of B. sulfuratus sp. nov. the irregular mark on the middle of the dorsum and the two circular dark blotches on the dorsal view of pelvic girdle are coincident with the x-shaped dorsal mark, though in this species this mark is not clearly distinguished. Another remarkable feature of the new species is the singular inverted v-shaped mark around the cloacal region in ventral view (Figure 6A), which is not clearly distinguishable in B. pulex (Napoli et al. 2011) and generally rounded and not ornamented in B. didactylus (Izecksohn 1971) and B. hermogenesi (Giaretta & Sawaya 1998; Figure 6C). The ornamented marks around the cloacal region of B. sulfuratus sp. nov. are usually sharper when compared to B. hermogenesi (Figure 6). The m-shaped mark around the cloacal opening, which is dark and defined in B. sulfuratus sp. nov. (Figure 6B), is present but not clearly defined in B. hermogenesi (Figure 6D). The main structure of the advertisement call of B. sulfuratus sp. nov. is exceptional within Brachycephalus. It differs greatly from B. ephippium, B. pitanga, and B. crispus because it is composed of a set of 4���7 high-fr, Published as part of Condez, Thais Helena, Monteiro, Juliane Petry De Carli, Comitti, Estev��o Jasper, Garcia, Paulo Christiano De Anchietta, Amaral, Ivan Borel & Haddad, C��lio Fernando Baptista, 2016, A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil, pp. 40-56 in Zootaxa 4083 (1) on pages 42-52, DOI: 10.11646/zootaxa.4083.1.2, http://zenodo.org/record/1050887, {"references":["Clemente-Carvalho, R. B. G., Antoniazzi, M. M., Jared, C., Haddad, C. F. B, Alves, A. C. R., Rocha, H. S., Pereira, G. R., Oliveira, D. F., Lopes, R. T. & Reis, S. F. (2009) Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): microscopic structure and macroscopic patterns of variation. Journal of Morphology, 270 (11), 1285 - 1295. https: // dx. doi. org / 10.1002 / jmor. 10755","Haddad, C. F. B., Alves, A. C. R., Clemente-Carvalho, R. B. G. & Reis, S. F. (2010) A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae). Copeia, 2010, 410 - 420. https: // dx. doi. org / 10.1643 / CH- 09 - 102","Pombal, J. P. Jr. (2010) A posicao taxonomica das \" variedades \" de Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae). Boletim do Museu Nacional, Nova Serie, Zoologia, 526, 1 - 12.","Clemente-Carvalho, R. B. G., Giaretta, A. A., Condez, T. H., Haddad, C. F. B. & Reis, S. F. (2012) A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from the Atlantic Forest of southeastern Brazil. Herpetologica, 68, 365 - 374. http: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 11 - 00085.1","Condez, T. H., Clemente-Carvalho, R. B. G., Haddad, C. F. B. & Reis, S. F. (2014) A new species of Brachycephalus (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, southeastern Brazil. Herpetologica, 70 (1), 89 - 99. https: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00044","Napoli, M. F., Caramaschi, U., Cruz, C. A. G. & Dias, I. R. (2011) A new species of flea-toad, genus Brachycephalus Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil. Zootaxa, 2739, 33 - 40.","Izecksohn, E. (1971) Novo genero e nova especie de Brachycephalidae do estado do Rio de Janeiro, Brasil. Boletim do Museu Nacional, Rio de Janeiro, nova serie, Zoologia, 280, 1 - 12.","Giaretta, A. A. & Sawaya, R. J. (1998) Second species of Psyllophryne (Anura: Brachycephalidae). Copeia, 1998, 985 - 987. https: // dx. doi. org / 10.2307 / 1447345","Pombal, J. P. Jr., Sazima, I. & Haddad, C. F. B. (1994) Breeding behavior of the pumpkin toadlet, Brachycephalus ephippium (Brachycephalidae). Journal of Herpetology, 28, 516 - 519. https: // dx. doi. org / 10.2307 / 1564972","Araujo, C. B., Guerra, T. J., Amatuzzi, M. C. O. & Campos, L. A. (2012) Advertisement and territorial calls of Brachycephalus pitanga (Anura: Brachycephalidae). Zootaxa, 3302, 66 - 67.","Garey, M. V., Lima, A. M. X., Hartmann, M. T. & Haddad, C. F. B. (2012) A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from southern Brazil. Herpetologica, 68 (2), 266 - 271. https: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 11 - 00074.1","Verdade, V. K., Rodrigues, M. T., Cassimiro, J., Pavan, D., Liou, N. & Lange, M. (2008) Advertisement call, vocal activity, and geographic distribution of Brachycephalus hermogenesi (Giaretta & Sawaya, 1998) (Anura, Brachycephalidae). Journal of Herpetology, 42 (3), 542 - 549. http: // dx. doi. org / 10.1670 / 07 - 287.1","Clemente-Carvalho, R. G. B., Klaczo, J., Perez, I., Alves, A. C. R, Haddad, C. F. B. & Reis, S. F. (2011) Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 61, 79 - 89. https: // dx. doi. org / 10.1016 / j. ympev. 2011.05.017","Padial, J. M., Grant, T. & Frost, D. R. (2014) Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa, 3825 (1), 1 - 132. https: // dx. doi. org / 10.11646 / zootaxa. 3825.1.1","Pimenta, B. V. S, Bernils, R. S. & Pombal, J. P. Jr. (2007) Amphibia, Anura, Brachycephalidae, Brachycephalus hermogenesi: Filling gap and geographic distribution map. Notes on Geographic Distribution. Checklist, 3 (3), 277 - 279.","Oliveira, J. C. F., Coco, L., Pagotto, R. V., Pralon, E., Vrcibradic, D., Pombal, J. P. Jr. & Rocha, C. F. D. (2012) Amphibia, Anura, Brachycephalus didactylus (Izecksohn, 1971) and Zachaenus parvulus (Girard, 1853): Distribution extension. Checklist, 8 (2), 242 - 244.","Clemente-Carvalho, R. B. G., Monteiro, L. R., Bonato, V., Rocha, H. S., Pereira, G. R., Oliveira, D. F., Lopes, R. T., Haddad, C. F. B., Martins, E. G. & Reis, S. F. (2008) Geographic variation in cranial shape in the pumpkin toadlet (Brachycephalus ephippium): a geometric analysis. Journal of Herpetology, 42 (1), 176 - 185.","Clemente-Carvalho, R. B. G., Perez, S. I., Tonhati, C. H., Condez, T. H., Sawaya, R. J., Haddad, C. F. B. & Reis, S. F. (2015) Boundaries of morphological and molecular variation and the distribution range of a miniaturized froglet, Brachycephalus nodoterga (Anura: Brachycephalidae). Journal of Herpetology. [in press]"]}

Dendropsophus ozzyi, new species (Figs. 3���6) Holotype: MPEG 27811, an adult male, collected on vegetation of pond within the forest, at Comunidade Bragan��a, Rio Paraconi, Floresta Nacional de Pau-Rosa (FNPR), Mau��s, State of Amazonas, Brazil (03�� 56 ��� 50 ���S and 58 �� 26 ��� 36 ���W, 45 meters above sea level [m a.s.l.]) on 25 February 2009, by M.J. Sturaro and P.L.V. Peloso (Figs 3���4). Paratopotypes: MPEG 27809���27814, five adult males, collected on 20 February to 0 3 March 2009, other data same as holotype. Paratypes: INPA-H 32742���32748, seven adult males on 23 June 2010, and 21 March 2011; CFBH 35760 and 35761, two adult males, on 23 June 2010, all collected by M. Gordo; MPEG 22400���22404, four adult males and one adult female, on 0 7 March 2008, collected by S. Neckel-Oliveira and H. F. Silva-Filho; all from Igarap�� Prudente (02�� 27 ' 13.03 "S; 56 �� 10 ' 54.14 "W), Juruti, State of Par��, Brazil. CFBH 35762���35763, two adults males, on 19 march 2011; CFBH 35764, an adult male, on 7 May 2013; MZUSP 154084���154088, five adult males, on 19 March 2011; MNRJ 86921���86925, five adult males, on 0 7 May 2013, all collected by M. Gordo; MPEG 22352���22355, three adult males and one adult female, on 0 8 March 2008, collected by S. Neckel-Oliveira and H. F. Silva-Filho; all from Igarap�� Mutum (02�� 36 ' 46.09 "S; 56 �� 11 ' 38.53 "W), Juruti, State of Par��, Brazil. Generic placement: We assign the new species to the genus Dendropsophus on the basis of its overall morphological similarity with other species of the genus (Figs. 3���6). Overall morphology (particularly the combination of size, dorsum color, and flash colors) and advertisement calls of D. ozzyi are similar to those of species assigned to the D. microcephalus Group. Within this group, the new species is most similar to D. shiwiarum Ortega-Andrade and Ron 2013 because both species are unique in having pointed discs (Ortega- Andrade & Ron 2013). Dendropsophus shiwiarum is assigned to the D. microcephalus Group largely due to its resemblance to D. riveroi (Ortega-Andrade & Ron 2013); the later was, however, originally assigned to the D. minimus Group and only recently shown to be part of the D. microcephalus Group (Fouquet et al. 2011). For the above reasons, we tentatively assign D. ozzyi to the D. microcephalus Group. We could not evaluate the putative morphological synapomorphies of the genus suggested by Faivovich et al. (2005) and we do not know the number of chromosomes for the species described here; a diploid number of 2 N = 30 seems to be a synapomorphy for the genus (Suarez et al. 2013). We were also unable to check the proposed larval synapomorphies for the D. microcephalus Group (Duellman & Trueb, 1983; Faivovich et al. 2005), as no tadpoles have been collected. Thus, we remark that a formal test of the relationships of D. ozzyi is still lacking. Diagnosis. Dendropsophus ozzyi is diagnosable by the following combination of character states: (1) a medium sized member of the Dendropsophus microcephalus Group, (SVL 18.5���21.5 mm in adult males, N = 37); (2) head wider than body; (3) snout truncate in dorsal and lateral views; (4) nostrils protuberant; (5) large prominent eyes (EL/ HW 0.3���0.5, N = 37); (6) nictitating membrane bearing brownish pigmentation on its border; (7) small tympanum (TD / ED 0.3���0.5, N = 37); (8) axillary membrane present; (9) glandular nuptial pads on finger II; (10) hands webbing formula II 2 - ��� 2 - III 2 - ��� 2 - IV 2 + ��� 2 + V; (11) feet webbing formula I 2 ��� 2 + II 1 + ��� 3 - III 1 +��� 2 + IV 2 + ��� 1 + V; (12) fingers and toes bearing pointed discs; (13) no inner tarsal fold; (14) calcar tubercle absent, heel with small, inconspicuous tubercles; (15) a dorsal sheath covering 1 / 3 of the cloacal opening; (16) in life, dorsal surfaces plain light brown with randomly distributed dark smudges; palmar, plantar and ventral surfaces of the axillary membrane, thigh, and shank vivid orange, belly creamy white with clear white spots and a rectangular bright white area, gular region orangish cream with brown flecks, vocal sac transparent; (17) advertisement call composed of a single, not pulsed, tonal, and highly pitched (dominant frequency around 9 kHz) note. FIGURE 3. Dendropsophus ozzyi sp. nov. holotype (MPEG 27811). (A) Dorsal, and (B) lateral views of head; (C) palmar view of right hand, and (D) plantar view right foot. Scale bar = 5 mm. Comparison with other species. Dendropsophus ozzyi differs from all other species of Dendropsophus, except D. shiwiarum ���also in the D. microcephalus Group���by having pointed discs on fingers and toes. In the following paragraphs we first diagnose our new species from D. shiwiarum and then provided additional diagnostic characters that differentiate D. ozzyi from all other species of the genus arranged by species groups. Dendropsophus ozzyi differs from D. shiwiarum by (character states of the later in parenthesis; all retrieved from Ortega-Andrade and Ron 2013) by the absence of dark brown interorbital, canthal, and post-orbital stripes extending to mid-flank (present); absence of suborbital bars or spots (present); shanks not presenting bars (present); gular region (and not the vocal sac) orangish cream with brown flecks (bright yellow, unflecked); by the absence of a distinct supratympanic fold (present). These species also differ by their color in life, while D. ozzyi presents vivid orange flash colors, D. shiwiarum presents ���fleshy white��� (sic) flash colors. ���Fleshy��� can be interpreted as a variety of colors, including orange. From pictures in Ortega-Andrade & Ron (2013) we understand that the color in D. shiwiarum is better described as pinkish-white. Morphometrically, Dendropsophus shiwiarum and D. ozzyi are distinguishable by their eye-to-nostril distance (EN = 1.2���1.8 in D. shiwiarum and EN = 2.31���2.37 mm in D. ozzyi). Advertisement calls of the two species differ by the longer duration of D. ozzyi (248 ms vs. 52.83 �� 31.87 ms of D. shiwiarum [original description does not present range]) and especially by the higher dominant frequency of D. ozzyi (9130.1���10136.7 Hz vs. 3983.6���5254.1 Hz of D. shiwiarum). From other species of the Dendropsophus microcephalus Group, D. ozzyi is distinguished from species of the D. rubicundulus Clade sensu Faivovich et al. (2005) (traits of the later in parentheses) by the stouter body (body slender), light brown dorsum in live and preserved specimens (dorsum dark green in life and violet in preserved specimens) with unaligned spots (stripes or aligned spots). From species of the D. decipiens Clade sensu Faivovich et al. (2005) by the absence of dorsal patterns (all species framed, except D. berthalutzae that presents an ���>et al. 2000). The simple (monophasic) advertisement call also distinguishes D. ozzyi from most Dendropsophus species (see below). The high dominant frequency distinguishes D. ozzyi from all other congeners (Table 2), except from D. minusculus (9020���9360 Hz see Duellman & Pyles 1983) although call duration of D. ozzyi is much longer [165���386 ms vs. 10���46 ms (Duellman & Pyles 1983; Lescure & Marty 2000 respectively)]. Dendropsophus ozzyi is distinguished from species of the D. columbianus Group by the smaller SVL [combined SVL = 21.1 mm in D. columbianus, to 33.5 mm in D. bogerti (data of both species from Cochran & Goin 1970)], except that the maximum SVL of D. ozzyi is 0.5 mm longer than the minimum SVL of D. columbianus. Apart from that, members of the D. columbianus Group have flecked or marbled bellies, and dorsum with dark spots (Duellman 1989; Rivera-Correa & Guti��rrez-C��rdenas 2012). Additionally, advertisement calls of species of the D. columbianus Group are diphasic (see Duellman & Trueb 1983) while D. ozzyi has a simple call. Dendropsophus ozzyi is also smaller than species of the D. garagoensis Group [combined SVL ranges from 21.3 mm in D. virolinensis (see Kaplan & Ruiz-Carranza 1997) to 31.5 mm in D. praestans (see Duellman & Trueb 1983) specimens], except that the maximum SVL of D. ozzyi is 0.2 mm larger then the minimum SVL of D. virolinensis (21.3 mm). Species of the D. garagoensis Group also present different dorsal patterns, with paramedial white lines and dark blotches in the flanks (both traits absent in D. ozzyi) (Kaplan 1991; 1997; Kaplan & Ruiz- Carranza 1997). To date, advertisement calls of the D. garagoensis Group are unknown. From species of the D. labialis, D. marmoratus and D. parviceps Groups, the new species differs by the homogeneous light brown dorsum whereas (green in life fading to gray in preserved specimens in the D. labialis Group and lichenous, both in life and in preserved specimens, in the D. marmoratus and D. parviceps Groups). From the D. labialis and D. marmoratus Groups by the smaller SVL [combined SVL ranging from 27.5 mm in D. melanargyreus (VGDO unpub. data) to 61 mm in D. labialis, (from Guarnizo et al. 2012)]. Species of the D. marmoratus and D. parviceps Groups have black flash colors [species of the D. parviceps Group with or without bright yellow spots (see Duellman & Crump 1974)], and species of the D. labialis Group have blue colors (e.g., Guarnizo et al. 2012), while D. ozzyi has orange flash colors. Additionally, many species of the D. marmoratus and D. parviceps Groups have suborbital bars (e.g., Duellman & Crump 1974; Lutz & Bokermann 1963). These character states also distinguish D. ozzyi from D. yaracuyanus, a species without formal assignment to a species group (Mijares-Urrutia & Rivero 2000) but quite similar to species of the D. parviceps Group. Additionally, calls of species of the D. parviceps Group are complex (e.g., Amezquita & Hodl 2004; Duellman & Crump 1974; Marquez et al. 1993) while D. ozzyi has a simple call. Calls of the D. labialis (Guarnizo et al. 2012) and D. marmoratus Group (Orrico et al. 2009) are longer and have much lower dominant frequencies. Dendropsophus ozzyi is smaller than species of the D. leucophyllatus Group [combined SVL ranging from 23 mm in D. bifurcus (see Kaplan & Ruiz-Carranza 1997) to 40 mm in D. anceps specimens (see Duellman & Trueb 1983)]. Exceptions are D. rossalleni [19���22.3 mm (De la Riva & Duellman 1997)] and smaller individuals of D. elegans [20 mm (Lutz, 1973)] as D. ozzyi reaches a maximum SVL of 21.5 mm. Additionally, D. ozzyi does not have any interorbital bar or blotch of different color of the dorsum ground color, as observed in all species of the D. leucophyllatus Group. Furthermore, species of the D. leucophyllatus Group have pectoral glandular patches���a likely synapomorphy for the group (see comments in Rivera-Correa & Orrico 2013)���which are absent in D. ozzyi Additionally, species of the D. leucophyllatus Group present diphasic calls (e.g., Conte et al. 2010; Jungfer et al. 2010; Marquez et al. 1993) while D. ozzyi presents a monophasic call. Dendropsophus ozzyi is distinct from species of the D. minutus Group by possessing a small cloacal sheath partially covering the cloaca (long cloacal sheath covering entirely the cloaca in D. minutus Group) and by the absence of white supracloacal and tarsal lines (present in species of the D. minutus Group). These traits also distinguish D. ozzyi from D. stingi and D. amicorum that present such structure and are quite similar to species of the D. minutus Group although not assigned to any species group in the original description (see Kaplan 1994; Mijares-Urrutia 1998). These same traits also distinguish D. ozzyi from D. aperomeus, a species of the D. minimus Group (Duellman 1982). The Dendropsophus minimus Group has been consistently recovered as paraphyletic but to date it is still recognized because D. minimus has never been included in a phylogenetic analysis (Fouquet et al. 2011); therefore, the status of the group name remains unclear (Frost 2014). We group the comparisons of D. ozzyi with the remaining species still associated with this possibly paraphyletic arrangement for convenience only, however, not formally recognizing it. Dendropsophus minimus presents a concealed tympanum (clearly visible in D. ozzyi) a clear tarsal fold (absent in D. ozzyi), a rostral white line (absent in D. ozzyi) and a different dorsal pattern (Ahl 1933). Dendropsophus miyatai presents a concealed tympanum (visible in D. ozzyi), a bright red over bright yellow, dorsal pattern in life (light brown in D. ozzyi), and absence of fimbriae on hind limbs (present in D. ozzyi) (Vigle & Goberdhan-Vigle 1990). Additionally, calls of the D. minutus Group are complex, diphasic or even triphasic (e.g., Cardoso & Haddad 1984; Duellman 1978; Heyer et al. 1990; Kohler & Lotters 2001 b; Marquez et al. 1993; Morais et al. 2012) while D. ozzyi has a simple, monophasic, call. The following comparisons pertain to species of the genus not assigned to a group (Faivovich et al. 2005). Dendropsophus ozzyi differs from D. haraldschultzi by presenting a light brown smooth dorsum while D. haraldschultzi has a grayish cream dorsum with thin longitudinal stripes composed of small dots and with a series of glands, more densely distributed over the head (Bokermann 1962). From D. battersbyi, D. ozzyi can be distinguished mainly by the smaller SVL���males of D. ozzyi range between 19.45���21.53 mm vs. 33 mm of the male holotype of D. battersbyi, and by different overall coloration; while D. battersbyi is gray and minutely dotted with reddish brown on all the upper and lower surfaces except the forelimbs and belly, D. ozzyi is light brown. Dendropsophus battersbyi presents a dark canthal and supratympanic streak, loreal region whitish with dark dots, limbs crossbarred and a pair of callous, white spots on the buttocks���features absent in D. ozzyi (see Rivero 1961). Dendropsophus tintinnabulum presents a concealed tympanum (visible in D. ozzyi) and a white line dividing the dorsum from the flanks in preservative (VGDO observation on the syntypes). In life, D. tintinnabulum presents a grayish green dorsum and the venter is bluish green (Melin 1941); D. ozzyi has a light brown dorsum and the venter is orange and white. Description of holotype: Adult male, SVL 19.4 mm; head wider than long (HW /HL 1.18), widest below eyes; snout truncate in both dorsal and lateral views, eye-nostril distance longer than eye diameter (END / ED 1.17); canthus rostralis indistinct, straight; loreal region slightly concave; lips thin; internarial area slightly depressed; nostrils protuberant, directed dorsolaterally; interorbital area flat; IOD / ED 1.48, IOD / HW 0.49; eyes large and protuberant (ED /HL 0.40, ED / HW 0.33); ELW 1.38 mm. Pupil horizontally elliptical; nictitating membrane transparent, its border slightly pigmented in the same pattern of the eyelid. Supratympanic fold barely visible, semi-circular in outline, less visible on the posterior portion; tympanum small (TD /HL 0.17), distinct, nearly round, separated from eye by a distance of 1.3 tympanum diameter; choanae small, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores very small, the right bearing two small, unaligned teeth while the left bears three; tongue cordiform, notched behind, posterior 1 / 3 not adhered to floor of mouth; vocal slits long, extending from midlateral base of tongue to almost the angle of jaws; vocal sac developed, single, and subgular, extending over the forearm (Figs. 3���4). Arm slender, not hypertrophied; axillary membrane reaching half arm, delimited by the m. pectoralis portio axillaris (seen by transparency). Lateral border of arm with fimbria composed of small tubercles, not all clearly individualized. Fingers short, bearing discs; finger discs are slightly pointed, albeit disc of Finger II is slightly less, close to round; relative length of fingers II Coloration in preservative: All dorsal surfaces yellowish brown, except thighs, fingers II, III, IV and toes I, II, III, IV that are cream. Dorsal ground color is darker at the head, fading posteriorly. Dorsum bears 13 brown smudges of irregular shape. Canthus rostralis, eyelid and the anterior third of the flanks have brown and white markings. Iris black with m, Published as part of Orrico, Victor G. D., Peloso, Pedro L. V., Sturaro, Marcelo J., Da Silva-Filho, Heriberto F., Neckel-Oliveira, Selvino, Gordo, Marcelo, Faivovich, Juli��n & Haddad, C��lio F. B., 2014, A new " Bat-Voiced " species of Dendropsophus Fitzinger, 1843 (Anura, Hylidae) from the Amazon Basin, Brazil, pp. 341-361 in Zootaxa 3881 (4) on pages 343-354, DOI: 10.11646/zootaxa.3881.4.3, http://zenodo.org/record/230815, {"references":["Jungfer, K. - H., Reichle, S. & Piskurek, O. (2010) Description of a new cryptic southwestern Amazonian species of leaf-gluing treefrog, genus Dendropsophus (Amphibia: Anura: Hylidae). Salamandra, 46, 204 - 213.","Ortega-Andrade, H. M. & Ron, S. (2013) A new species of small tree frog, genus Dendropsophus (Anura: Hylidae) from the eastern Amazon lowlands of Ecuador. Zootaxa, 3652 (1), 163 - 178. http: // dx. doi. org / 10.11646 / zootaxa. 3652.1.6","Fouquet, A., Noonan, B. P., Blanc, M. & Orrico, V. G. D. (2011) Phylogenetic position of Dendropsophus gaucheri (Lescure and Marty 2000) highlights the need for an in - depth investigation of the phylogenetic relationships of Dendropsophus (Anura: Hylidae). Zootaxa, 3035, 59 - 67.","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, 1 - 240. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2005) 294 [0001: SROTFF] 2.0. CO; 2","Duellman, W. E. & Trueb, L. (1983) Frogs of the Hyla columbiana group: Taxonomy and phylogenetic relationships. In: Rhodin, A. G. J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary biology. Museum of Comparative Zoology, Harvard University, Cambridge, pp. 33 - 51.","Pugliese, A., Alves, A. C. R. & Carvalho-e-Silva, S. P. (2000) The tadpoles of Hyla oliveirai and Hyla decipiens with notes on the Hyla microcephala group (Anura, Hylidae). Alytes, 18, 73 - 80.","Duellman, W. E. & Pyles, R. A. (1983) Acoustic resource partitioning in anuran communities. Copeia, 1983, 639 - 649. http: // dx. doi. org / 10.2307 / 1444328","Lescure, J. & Marty, C. (2000) Atlas des Amphibiens de Guyane. Patrimoines Naturels, 45, 1 - 388.","Cochran, D. M. & Goin, C. J. (1970) Frogs of Colombia. Bulletin of the United States National Museum, 288, 1 - 655. http: // dx. doi. org / 10.5479 / si. 03629236.288.1","Duellman, W. E. (1989) New Species of Hylid Frogs from the Andes of Colombia and Venezuela. Occasional papers of the Museum of Natural History of the University of Kansas, 12 pp.","Rivera-Correa, M. & Gutierrez-Cardenas, P. D. A. (2012) A new highland species of treefrog of the Dendropsophus columbianus group (Anura: Hylidae) from the Andes of Colombia. Zootaxa, 3486, 50 - 62.","Kaplan, M. & Ruiz-Carranza, P. M. (1997) Two new species of Hyla from the Andes of Central Colombia and their relationships to other small Andean Hyla. Journal of Herpetology, 31, 230 - 244. http: // dx. doi. org / 10.2307 / 1565391","Kaplan, M. (1991) A new species of Hyla from the eastern slope of the cordillera oriental in northern Colombia. Journal of Herpetology, 25, 313 - 316. http: // dx. doi. org / 10.2307 / 1564589","Guarnizo, C. E., Escallon, C., Cannatella, D. & Amezquita, A. (2012) Congruence between acoustic traits and genealogical history reveals a new species of Dendropsophus (Anura: Hylidae) in the high Andes of Colombia. Herpetologica, 68, 523 - 540. http: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 10 - 00038","Duellman, W. E. & Crump, M. L. (1974) Speciation in frogs of the Hyla parviceps group in the upper amazon basin. Occasional papers of the Museum of Natural History of the University of Kansas, 23, 1 - 40.","Lutz, B. & Bokermann, W. C. A. (1963) A new tree Frog from Santa Catarina, Brazil. Copeia, 1963, 558 - 561. http: // dx. doi. org / 10.2307 / 1441477","Mijares-Urrutia, A. & Rivero, R. A. (2000) A new treefrog from the Sierra de Aroa, northern Venezuela. Journal of Herpetology, 34, 80 - 84. http: // dx. doi. org / 10.2307 / 1565242","Amezquita, A. & Hodl, W. (2004) How, when, and where to perform visual displays: the case of the Amazonian frog Hyla parviceps. Herpetologica, 60, 420 - 429. http: // dx. doi. org / 10.1655 / 02 - 51","Marquez, R., De la Riva, I. & Bosch, J. (1993) Advertisement calls of Bolivian species of Hyla (Amphibia, Anura, Hylidae). Biotropica, 25, 426 - 443. http: // dx. doi. org / 10.2307 / 2388866","Orrico, V. G. D., Lingnau, R. & Giasson, L. O. M. (2009) The advertisement call of Dendropsophus nahdereri (Anura, Hylidae, Dendropsophini). South American Journal of Herpetology, 4, 295 - 299. http: // dx. doi. org / 10.2994 / 057.004.0313","de la Riva, I. & Duellman, W. E. (1997) The identity and distribution of Hyla rossalleni Goin. Amphibia-Reptilia, 18, 433 - 436. http: // dx. doi. org / 10.1163 / 156853897 X 00486","Lutz, B. (1973) Brazilian species of Hyla. University of Texas Press, Austin, 265 pp.","Rivera-Correa, M. & Orrico, V. G. D. (2013) Description and phylogenetic relationships of a new species of treefrog of the Dendropsophus leucophyllatus group (Anura: Hylidae) from the Amazon basin of Colombia and with an exceptional color pattern. Zootaxa, 3686 (4), 447 - 460. http: // dx. doi. org / 10.11646 / zootaxa. 3686.4.3","Conte, C. E., Nomura, F., Machado, R. A., Kwet, A., Lingnau, R. & Rossa-Feres, D. d. C. (2010) Novos registros na distribuicao geografica de anuros na Floresta com Araucaria e consideracoes sobre suas vocalizacoes. Biota Neotropica, 10, 201 - 224. http: // dx. doi. org / 10.1590 / S 1676 - 06032010000200024","Kaplan, M. (1994) A new species of frog of the genus Hyla from the cordillera oriental in northern Colombia with comments on the taxonomy of Hyla minuta. Journal of Herpetology, 28, 79 - 87. http: // dx. doi. org / 10.2307 / 1564684","Mijares-Urrutia, A. (1998) Una nueva especie de rana arboricola (Amphibia: Hylidae) de un bosque nublado del oeste de Venezuela. Revista Brasileira de Biologia, 58, 659 - 663. http: // dx. doi. org / 10.1590 / S 0034 - 71081998000400013","Duellman, W. E. (1982) A new species of small yellow Hyla from Peru (Anura: Hylidae). Amphibia-Reptilia, 3, 153 - 160. http: // dx. doi. org / 10.1163 / 156853882 X 00383","Frost, D. (2014) Amphibian Species of the World: an Online Reference. Version 6.0 (9 January 2013). In: American Museum of Natural History, New York. Available from: http: // research. amnh. org / vz / herpetology / amphibia / (accessed February 26 2014)","Ahl, E. (1933) Uber einige neue Frosche aus Brasilien. Zoologischer Anzeiger Leipzig, 104, 25 - 30.","Vigle, G. O. & Goberdhan-Vigle, D. C. I. (1990) A new species of small colorful Hyla from the Lowland rainforest of Amazonian Ecuador. Herpetologica, 46, 467 - 473.","Cardoso, A. J. & Haddad, C. F. B. (1984) Variabilidade acustica em diferentes populacoes e interacoes agressivas de Hyla minuta (Amphibia, Anura). Ciencia e Cultura, 36, 1393 - 1399.","Duellman, W. E. (1978) The biology of an equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 65, 1 - 352.","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.","Kohler, J. & Lotters, S. (2001 b) Description of a small tree frog, genus Hyla (Anura: Hylidae), from humid Andean slopes of Bolivia. Salamandra, 37, 175 - 184.","Morais, A. R., Batista, V. G., Gambale, P. G., Signorelli, L. & Bastos, R. P. (2012) Acoustic communication in a Neotropical frog (Dendropsophus minutus): vocal repertoire, variability and individual discrimination. Herpetological Journal, 22, 249 - 257.","Bokermann, W. C. A. (1962) Cuatro nuevos hylidos del Brasil (Amphibia, Salientia, Hylidae). Neotropica, 8, 81 - 91.","Rivero, J. A. (1961) Salientia of venezuela. Bulletin of the Museum of Comparative Zoology, Harvard College, 126, 1 - 207.","Melin, D. (1941) Contributions to the knowledge of the amphibia of South America. Vol. 6. Goteborgs Vetensk Samh. Handl, 71 pp. [1 B]","Abrunhosa, P. A., Wogel, H. & Pombal, J. P. Jr. (2001) Vocalizacoes de quatro especies de anuros do estado do Rio de Janeiro (Amphibia, Hylidae, Leptodactylidae). Boletim do Museu Nacional - Nova Serie, Zoologia, 472, 1 - 12.","Tarano, Z. (2010) Advertisement Calls and Calling Habits of Frogs from a Flooded Savanna of Venezuela. South American Journal of Herpetology, 5, 221 - 240. http: // dx. doi. org / 10.2994 / 057.005.0308"]}

Elachistocleis cesarii (Miranda-Ribeiro, 1920) Engystoma ovale cesarii Miranda-Ribeiro, 1920 Engystoma ovale lineatum Miranda-Ribeiro, 1920 Lectotype. MZUSP 529, an adult male, collected at municipality of Piquete, state of S��o Paulo, southeastern Brazil (approximately 22 �� 36 ��� S; 47 �� 10 ��� W; 675 m above sea level), on November 1896 by Zech (Figure 1). Paralectotypes. ��� MZUSP 33 collected on January 1900 by Hempel, at Ipiranga (Ypiranga), municipality of S��o Paulo; MZUSP 36 (gravid female) collected on October 1896 by Bicego, at municipality of Perus; MZUSP 37 (gravid female) collected on 1897 by Bicego, at municipality of Cubat��o; MZUSP 38 collected on January 1898 by Bicego, at Bel��m, municipality of S��o Paulo; MZUSP 42 (gravid female) collected on October 1907 by Ruedervalt and Cheubel at Ipiranga (Ypiranga), municipality of S��o Paulo; MZUSP 264 (gravid female) collected together with the lectotype; MZUSP 715 collected on 1896 by Bicego, at Alto da Serra; MZUSP 2023 (gravid female) collected on 1897 by Bicego at municipality of Cubat��o; MZUSP 2024 - 27 (gravid females; except for 2025 which resembles an young male) collected on October 1896 by Bicego at municipality of Perus. All municipalities are in the state of S��o Paulo, southeastern Brazil. Diagnosis. Elachistocleis cesarii is recognized by the following combination of characters: 1) ventral coloration is white or yellow with gray marks and reticulations; 2) presence of postcommisural glands behind the corner of mouth; 3) tympanum concealed; 4) advertisement call with the mean dominant frequency of about 3.5 kHz. Comparison with other species. In life, Elachistocleis cesarii is readily distinguished from E. erythrogaster by the absence of the reddish coloration of the ventral portion of its body (see Figures 2 C, D and 3 C, D), its smallest body size, and the presence of a well defined femoral stripe (not well defined in E. erythrogaster). Besides this, the frequency of the advertisement call of E. cesarii is lower than that reported to E. erythrogaster (see Kwet & Di-Bernardo 1998; 1999). E. cesarii is distinguished from E. surinamensis by its smaller size (mean of 32.3 mm in males of E. surinamensis) and by differences in body coloration (see plate XIII in Kenny 1969, and Figures 3 C, D, and E). E. cesarii is distinguished from E. bicolor and E. ovalis by the absence of the immaculate, yellow, ventral coloration (spotted in E. cesarii) (see Lavilla et al. 2003 and Figures 2 A, B and 3 C, D). E. cesarii is distinguished from E. skotogaster by presenting postcommisural glands (absent in E. skotogaster; see Figure 2 E) and by its smaller size (minimum size of 27.5 in males of E. stokogaster). E. cesarii is distinguished from E. piauiensis by some advertisement call parameters: the call frequency is lower and call duration is shorter in E. cesarii than in E. piauiensis (see Table 2). Besides this, E. cesarii has a smaller postcommisural gland (more conspicuous in E. piauiensis) and the blotches of the inguinal region are less conspicuous in E. piauiensis than in E. cesarii. In figures 2 and 3 it is possible to observe dorsal and ventral color patterns of some species of Elachistocleis in life. Description of the lectotype. Body robust and ovoid. Texture of the dorsal skin and undersurface of the body smooth. Skin on ventrolateral surface of body and area around the cloacal region smooth. Cloacal flap absent; vent opening positioned posterodorsally. Head small and triangular, broader than long. Cranial crests absent. A conspicuous dorsolateral skin fold from the arms to the groin. Another skin fold crossing the chest between axillae. Eyes small, slightly prominent. Snout pointed in dorsal view and protruding in lateral view. Nostrils directed anterolaterally, slightly protuberant. Choanae relatively small and round. A well developed groove in the terminal part of the lower jaw. Internarial distance small. Canthus rostralis distinct, rounded. Loreal region slightly convex. Interorbital space slightly convex. Lips nearly horizontal when belly is touching the ground. Presence of postcommisural glands behind the corner of mouth. Tympanum concealed. Supratympanic fold absent. Upper jaw projects 1.59 mm beyond the lower jaw. Subgular vocal sac not expanded externally. Vocal slits present. Vomerine odontophores absent. Tongue large, oval. Arms and fingers robust and short. Legs extremely robust; almost equal lengths of thigh, tibia, and foot. Finger lengths IColor in life of individuals apart from the type series (Figure 3 C, D). ��� Dorsum and limbs are dark brownish gray with small white dots. The single, subgular vocal sac is darker than the venter. The chest is yellow with gray marks. Belly white or yellow, with gray marks and reticulations, which reach the flanks. Orange femoral stripe present on the posterior surface of the thigh. Orange inguinal spots present. There is a tiny white vertebral line from the snout to the vent. Color in preservative of individuals other than the type series (70 % ethanol). ��� Dorsum and limbs are dark grayish blue. Ventral region is gray with white spots. These spots reach up to the lateral portion of the animals. The throat is darker than chest and belly. The inguinal spots and femoral stripes on the posterior surface of the thigh are white. Vertebral line undistinguished. Variation. Females are larger than males (see Table 1). Females present variation in size, with the SVL varying from 19.4 up to 42.7 mm (Thom�� & Brasileiro, 2007). The vertebral stripe may be absent. The ventral spots may not reach the flanks and the groin region of some individuals. The throat of males is darker than that of females. Call descriptions. Advertisement calls of E. cesarii were recorded in Itap��, municipality of Rio Claro, S��o Paulo state, Brazil, on 29 January 2002, at air temperature of 23.1 ��C (Figure 4). The call consists of a loud, high pitched, sustained whistle, ranging from 1.49 to 2.66 s of duration, with a mean dominant frequency of 3,515 Hz, with a mean pulse rate of 238.2 per second, and 460.2 pulses per note in average. The frequency ranged from 3.01 to 4.17 kHz. The duration of the pulses did not vary much (mean of 4.23 �� 0.16 ms; n = 50). Further spectral characteristics are presented in Table 2 (see also Figure 4). Advertisement calls of E. piauiensis were recorded in the municipalities of Pacatuba on 13 March 2007 (air temperature 22 ��C), Caucaia on 15 May 2005 (air temperature 27 ��C), and Vi��osa do Cear�� on 24 April 2007 (air temperature 25 ��C), all of them in the state of Cear��, Northeastern Brazil. The call structure of E. piauiensis is very similar to that of E. cesarii and consist of a loud, high pitched, sustained whistle (ranging from 2.45 to 2.51 s of duration). The duration of the pulses did not vary much (3.00 �� 0.23 ms; n = 50). However, the mean dominant frequency is higher in E. piauiensis (4.53 �� 0.06 Hz) compared to the new species and the frequency ranged from 3.18 to 5.10 kHz (Table 2; Figure 4). Tadpoles. Tadpoles are unknown; however, 10 tadpoles collected in the municipality of Vit��ria Brasil, state of S��o Paulo, Brazil, and briefly described under the name Elachistocleis sp. by Rossa-Feres and Nomura (2006), could be E. cesarii. Natural history notes. Elachistocleis cesarii calls during the wet and hot season of the year (summer) as a prolonged breeder; however, after heavy rains during the hot nights it may behave like an explosive breeder (sensu Wells 1977). Males call with the legs in the water and the body out of water, holding the emergent vegetation with the forelimbs. The amplexus is axillar. We collected two amplected pairs that laid eggs in plastic bags. One female deposited 736 eggs and the second deposited 1126 eggs. In nature, the eggs are laid in the water surface as a gelatinous mass attached to floating or emergent vegetation (Figure 3 F). Newly metamorphosed specimens have a mean SVL of 11.83 mm �� 0.38 (11.4 ��� 12.3; N = 6). The ecology of this species has been studied by Thom�� and Brasileiro (2007) and a short communication about the defensive posture was published by Toledo (2004), in both articles under the name Elachistocleis cf. ovalis. Geographical distribution. Besides the localities of the type series, the species is known in the municipalities of Rio Claro, Campinas, Itirapina, S��o Paulo, Angatuba, and Ubatuba, all in the state of S��o Paulo. In the state of Minas Gerais it can be found in the municipalities of Jaboticatubas, Santana do Riacho, Munhoz, and Vi��osa. In the state of Goi��s it can be found in Pontalina (Figure 5). Etymology. The meaning of the specific name of E. cesarii was not stated in the original description. Discussion. The taxonomic status of Elachistocleis ovalis remains unclear. This fact limits the recognition of several related still undescribed species in South America. These species are incorrectly being referred to as E. ovalis and even E. bicolor in the past and current literature, as well as in museum collections. The lack of correct identification of all those taxa has direct implication in conservation, as some of them may be endangered or may be endemic to certain localities, countries or biomes. Therefore, a taxonomic review of the genus urges attention and may reveal a great hidden biodiversity. Additional specimens examined. Elachistocleis bicolor: CFBH 3841; 3859; 4010 S��o Domingos, state of Santa Catarina, Brazil; CFBH 4225 Tabapu��, state of S��o Paulo, Brazil; CFBH 13608 - 10 Tr��s Lagoas, state of Mato Grosso do Sul, Brazil; CFBH 18199 Bom Jesus, state of Rio Grande do Sul, Brazil. E. cesarii: CFHB 4132 ��� 37; 4147 ��� 48; 4209; 4230 ��� 31; 4233 ��� 37; 4256 ��� 7 Rio Claro, state of S��o Paulo, Brazil; ZUEC 8054 Campinas, state of S��o Paulo, Brazil; ZUEC 12031 Itirapina, state of S��o Paulo, Brazil, CFBH 4997 Itirapina, state of S��o Paulo, Brazil; CFBH 3768 Pontalina, state of Goi��s, Brazil; CFBH 6575-6578 Rio Claro, state of S��o Paulo, Brazil; CFBH 10907 Ubatuba, state of S��o Paulo, Brazil; CFBH 23136 Angatuba, state of S��o Paulo, Brazil; ZUEC 599-600 S��o Paulo, state of S��o Paulo, Brazil; ZUEC 4246 -48, 4250-51, 4263 Munhoz, state of Minas Gerais, Brazil; ZUEC 1417 -18, 2848, 2862 -63, 3031-32, 3069, 3072 - 76 Jaboticatubas, state of Minas Gerais, Brazil; ZUEC 6157 (adult male recorded) Vi��osa, state of Minas Gerais, Brazil; ZUEC 4995 - 97 Santana do Riacho, state of Minas Gerais, Brazil. E. erythrogaster: MNRJ 39098 adult male, Paratype. E. piauiensis: CFBH: 15880; 15902 -03; 23465 - 71 Vi��osa do Cear��, state of Cear��, Brazil. E. surinamensis: ZUEC 9422 ��� 23 Rapidos de Komoiran, Estado Bolivar, Venezuela., Published as part of Toledo, Lu��s Felipe, Loebmann, Daniel & Haddad, C��lio F. B., 2010, Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae), pp. 50-60 in Zootaxa 2418 on pages 52-59, DOI: 10.5281/zenodo.194434, {"references":["Miranda-Ribeiro, A. (1920) Os Engystomatideos do Museu Paulista (com um genero e tres especies novos). Revista do Museu Paulista, 12, 280 - 288, lams. 1 - 2.","Kwet, A. & Di - Bernardo, M. (1998) Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and Environment, 33, 7 - 18.","Kenny, J. S. (1969) The Amphibia of Trinidad. In: Hummelinck, P. W. (Ed.). Studies on the fauna of Curacao and other Caribbean islands, 108, pp. 1 - 78.","Lavilla, E. O., Vaira, M. & Ferrari, L. (2003) A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis ovalis - E. bicolor controversy. Amphibia-Reptilia, 24, 269 - 284.","Thome, T. C. & Brasileiro, C. A. (2007) Dimorfismo sexual, uso do ambiente e abundancia sazonal de Elachistocleis cf. ovalis (Anura: Microhylidae) em um remanescente de Cerrado do estado de Sao Paulo, sudeste do Brasil. Biota Neotropica, 7 (1), bn 00307012007.","Rossa-Feres, D. C. & Nomura, F. (2006) Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of Sao Paulo state, Brazil. Biota Neotropica, 6 (1), bn 00706012006.","Wells, K. D. (1977) The social behaviour of anuran amphibians. Animal Behaviour, 25, 666 - 693.","Toledo, L. F. (2004) Elachistocleis cf. ovalis (Common Oval Frog). Death feigning. Herpetological Review, 35 (4), 371 - 372."]}

Ischnocnema vizottoi sp. nov. (Figures 1, 2, 3, and 4) Holotype. CFBH 8222, adult male, collected at Parque Estadual de Campos do Jord��o (22 �� 39`50 ``S, 45 �� 27`03`` W, 1540 m elevation), Municipality of Campos do Jord��o, S��o Paulo, Brazil, on 17���21 December 2004 by I. A. Martins, A. F. L. Leite, and F. B. R. Gomes. Paratopotypes. CFBH 8205 ���08, 8210 ���21, 16 adult males, and CFBH 8209, adult female, collected with the holotype. CCLZU / IAM 2151, adult male collected by I. A. Martins, F. B. R. Gomes, and A. F. B. Junqueira, on 3 March 2005. CCLZU / IAM 2161, adult male collected by I. A. Martins, F. B. R. Gomes, and T. L. P. C. Perroni, on 8 April 2005. CCLZU / IAM 2149, 2159, two adult females, and CCLZU / IAM 2158, 2160, two adult males, collected by C. F. B. Haddad, I. A. Martins, and F. B. R. Gomes, on 11���14 November 2005. CCLZU / IAM 2148, 2150, two adult females, and CCLZU / IAM 2152 ��� 57, six adult males collected on 21���23 December 2005 by I. A. Martins, F. B. R. Gomes, and A. F. B. Junqueira. Diagnosis. A member of the Ischnocnema lactea Species Series. It differs from other members of the Species Series by the following combination of traits: (1) diminutive size (males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL) (Table 1); (2) iris yellow in live specimens; (3) snout sub-elliptical in dorsal view and acuminate-rounded in lateral view; (4) vocal sac expanded externally; (5) tympanic membrane differentiated; (6) skin texture granular on the flanks and venter; (7) adhesive disks rounded; (8) calcar appendage absent or small; (9) external face of the thigh with a light line in the central area or irregular light blotches; (10) notes of the advertisement call with two harmonics; (11) note duration 38���72 ms; (12) mean dominant frequency of the advertisement call 3611 �� 103 Hz (3417���3763 Hz). TABLE 1. Measurements of Ischnocnema vizottoi sp. nov. (holotype and paratopotypes). Mean (x), standard deviation (SD), and range (in millimeters) for males and females. Comparison with other species. The new species differs from Ischnocnema sambaqui by its smaller size (I. vizottoi sp. nov. males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL; I. sambaqui males 32.3 ���40.0 mm SVL), finger adhesive disks rounded (T-shape in I. sambaqui), narrower head, and higher dominant frequency of advertisement call (I. vizottoi sp. nov. 3417���3763 Hz, I. sambaqui 1800���2050 Hz; Castanho & Haddad, 2000). The new species differs from Ischnocnema manezinho by its smaller size (I. vizottoi sp. nov. males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL; I. manezinho males 22.7���28.1 mm SVL and females 29.8���34.8 mm SVL; Garcia, 1996), and finger adhesive disks rounded (T-shaped in I. manezinho). The new species differs from Ischnocnema bolbodactyla by its smoother dorsal skin, shorter snout, adhesive disks less developed, absence of bright coloration on the inguinal region (inguinal region bright orange in I. bolbodactyla), and advertisement call formed by a single note (formed by series of 3���4 notes in I. bolbodactyla). The new species differs from Ischnocnema lactea by its smaller size (I. vizottoi sp. nov. males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL; I. lactea female lectotype 32mm SVL) and finger adhesive disks rounded (T-shaped in I. lactea). The new species differs from Ischnocnema holti by its smaller size (I. vizottoi sp. nov. males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL; I. holti 19 mm SVL; Cochran, 1955; male mean SVL 21.2 mm and female SVL 25.6mm; Targino & Carvalho-e-Silva, 2008) and snout outline sub-elliptical from above and acuminate-rounded in profile (snout in dorsal and lateral views rounded in I. holti). The new species differs from Ischnocnema concolor by its smaller size (I. vizottoi sp. nov. males 13.3���16.6 mm SVL and females 18.4���22.1 mm SVL; I. concolor males 15.0��� 18.4mm SVL; Targino et al., 2009), by a light line or irregular light blotches on the external face of the central area of the thigh (I. concolor lacks a light line or irregular light blotches in the central area of the thigh; Targino et al., 2009), and advertisement call traits [the notes in the advertisement call of I. vizottoi have two harmonics; minimum and maximum fundamental frequency (H 1) of 2780���3555 Hz, 3634���4097 Hz, respectively; the notes in the advertisement call of I. concolor have three harmonics; minimum and maximum fundamental frequency (H 1) of 2616���2989 Hz, 3150���3470 Hz, respectively; Martins, I.A. unpublished data]. The new species differs from Ischnocnema melanopygia by the smaller or absent calcar appendage (calcar appendage well developed in I. melanopygia; Targino et al., 2009) and by the absence of a black stripe on the perianal region, posterior area of tarsus, and feet (black stripe present in I. melanopygia; Targino et al., 2009). Description of the holotype. Body robust (Figure 1); snout sub-elliptical in dorsal view (Figure 2 A) and acuminate-rounded in lateral view (Figure 2 B); nostrils protuberant, directed laterally; canthus rostralis distinct; loreal region concave; eye large, protruding; tympanum distinct, large, diameter about 1 / 2 of eye diameter; small supratympanic fold extending from the back of the eye to near the arm insertion; vocal sac single, externally expanded; vocal slits present; tongue large, shallowly notched posteriorly; vomerine teeth in two small, oblique, and narrowly separated series of approximately 10 teeth each, between and behind choanae; choanae small, slightly rounded. Forearm moderately robust, arm slender; fingers long; thumb without nuptial pad; subarticular tubercles single, round; outer metacarpal tubercle cordiform, inner metacarpal tubercle elliptical; supernumerary tubercles on the palmar region (Figure 2 C); finger lengths IColor of the holotype in preservative. Dorsal surfaces brown. Vocal sac white with black punctuations. Inter-orbital mark dark brown. Elbow dark brown; dark brown marks on the forearms. Irregular light blotches on the external surface of the thigh. Two dark brown blotches on the sacrum region (Figure 1). Measurements of the holotype. SVL 14.50; HL 5.23; HW 5.00; ED 1.71; IOD 1.9; END 1.23; NSD 0.95; IND 1.33; TD 0.85; FL 6.75; TL 6.93; FOL 6.74; 3 FD 0.55; 4 TD 0.55. Color in life. The specimens of Ischnocnema vizottoi sp. nov. present wide variation of dorsal coloration (Figure 3). Dorsal coloration of different specimens can be green, red, black, brown, and light gray (Figure 3). A longitudinal light line extending from the snout to the vent is present in several specimens. A white or beige inter-orbital stripe is almost always visible, with a dark stripe behind the light inter-orbital stripe. Two dark blotches can be present in the middle of dorsum in some individuals. The external face of the thigh presents a light line in the central area or irregular light blotches. The subgular region may be lightly to very dark pigmented. In several specimens it is possible to see a light ventral line, from the anterior border of the chin to the chest. The ventral regions of the thigh and tibia present light spots with dark borders. Variation. Measurements of 27 males and five females are given in Table 1. The head is generally wider than long, but in some specimens it is longer than wide. The tympanum is distinct in the majority of the specimens. The vocal sac may be well expanded externally or indistinct. The calcar appendage may be smaller or absent. A light to dark stripe is generally present in the inter-orbital region. A light vertebral line is present in the majority of the specimens (Figures 3 and 4). The external face of the thigh may show a light line in the central area or irregular light blotches. Vocalization. The advertisement call of Ischnocnema vizottoi sp. nov. is composed of simple notes, with a fairly irregular rhythm and varied intervals (Figure 5). The average duration of the note is 52.7 �� 10.2 ms (38���72 ms), with a mean repetition rate of 9.5 �� 2.2 notes per minute. The advertisement call of Ischnocnema vizottoi sp. nov. has notes with two harmonics (Figure 5 B, D). The advertisement call has sharply rising frequencies at the beginning and sharply falling frequencies at the end (Figure 5). Most of the energy of these notes is concentrated in the first harmonic (fundamental frequency = H 1; Figure 5 E). The mean frequencies of the first harmonic (H 1) minimum frequencies is between 3204 �� 158 (amplitude 2780���3555 Hz) and maximum frequencies 3837 �� 95 Hz (amplitude 3634���4097 Hz), with most energy concentrated around 3611 �� 103 Hz (amplitude 3417���3763 Hz). The second harmonic (H 2) has minimum and maximum mean frequencies between 6967 �� 207 (amplitude 6585���7317 HZ), 7379 �� 119 Hz (amplitude 7146���7618 Hz) respectively, and most energy of the second harmonic is concentrated around 6996 �� 212 Hz (Figure 5). Distribution. The new species has been found in Serra da Mantiqueira at the localities of Parque Estadual de Campos do Jord��o, Municipality of Campos do Jord��o, S��o Paulo State, (type locality); Lavrinhas farm, Municipality of Pindamonhangaba, S��o Paulo State; Municipality of S��o Bento do Sapuca��, S��o Paulo State; Monte Verde, Municipality of Camanducaia, Minas Gerais State; and Serra do Mar, at Parque Nacional da Serra da Bocaina, Municipality of S��o Jos�� do Barreiro, S��o Paulo State. All the localities are in the Atlantic forest, above 1300 elevation (Figure 6). Etymology. The specific name honors Dr. Luiz Dino Vizotto, for his pioneer studies of Brazilian anurans and for his contribution to the knowledge of the Brazilian vertebrate fauna. Natural history. Ischnocnema vizottoi sp. nov. was observed in places of high altitude, from 1300 to 2100 m above sea level, inside forests, forest edges, and open fields. Males vocalize on the forest floor or perched 10 to 80 cm from the ground. The distance among neighboring males varied from 0.5 to 8 m. The calling season is from September to April; however, the vocalization was most intense from October to January, when gravid females were observed. Three analyzed females had from 10 to 18 oocytes with mean diameter of 2.45 �� 0.15 mm (N = 27). Oocytes in different developmental stages suggest that females may lay more than one clutch per season. Remarks. The Atlantic forest is the biome with the greatest endemism in species of anurans in the world, at the same time having high diversity of species and of phylogenetic groups (Duellman, 1999, Frost et al., 2006). This high diversity and endemism can be explained by the physical environment, a reflex of the complex geomorphologic, climatic, and phytogeographic characteristics of this formation. However, the species diversity of anurans in the Atlantic Forest of Brazil is still poorly known, particularly in places with more than 1000 m of altitude (Rossa-Feres et al., 2008). The number of species in the genus Ischnocnema will probably increase in the next years, considering the recent descriptions of new species (Giaretta et al., 2007; Targino et al., 2009) and mainly the large number of unnamed species that can be found in the shelves of Brazilian collections (personal observations). The use of molecular information and vocalizations, allied to morphological studies, will increase our comprehension on the evolution of this genus and certainly uncover a great diversity of cryptic species., Published as part of Martins, Itamar A. & Haddad, C��lio F. B., 2010, A new species of Ischnocnema from highlands of the Atlantic Forest, Southeastern Brazil (Terrarana, Brachycephalidae), pp. 55-65 in Zootaxa 2617 on pages 56-64, DOI: 10.5281/zenodo.197955, {"references":["Castanho, L. M. & Haddad, C. F. B. (2000) New species of Eleutherodactylus (Amphibia: Leptodactylidae) from Guaraquecaba, Atlantic Forest of Brazil. Copeia, 2000, 777 - 781.","Garcia, P. C. A. (1996) Nova especie de Eleutherodactylus Dumeril & Bribon, 1891 do Estado de Santa Catarina, Brasil (Amphibia; Anura; Leptodactylidae). Biociencias, Porto Alegre, 4, 57 - 68.","Cochran, D. M. (1955) Frogs of southeastern Brazil. United States National Museum Bulletin, 206, 1 - 423.","Targino, M. & Carvalho-e-Silva, S. P. (2008) Redescricao de Ischnocnema holti (Cochran, 1948) (Amphibia, Anura, Brachycephalidae). Revista Brasileira de Zoologia, 25, 716 - 723.","Targino, M., Costa, P. N. & Carvalho-e-Silva, S. P. (2009) Two new species of the Ischnocnema lactea species series from Itatiaia highlands, southeastern Brazil (Amphibia, Anura, Brachycephalidae). South American Journal of Herpetology, 4, 139 - 150.","Duellman, W. E. (1999) Global distribution of amphibians: patterns, conservation, and future challenges. In: Duellman, W. E. (ed.). Patterns of Distribution of Amphibians: A Global Perspective. Johns Hopkin Univ. Press, Baltimore, Maryland, pp. 1 - 30.","Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., de Sa´, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 370.","Rossa-Feres, D. C., Martins, M., Marques, O. A. V., Martins, I. A., Sawaya, R. J. & Haddad, C. F. B. (2008) Herpetofauna In: Rodrigues, R. R. & Bononi, V. L. R. (Org.), Diretrizes para conservacao e restauracao da biodiversidade no Estado de Sao Paulo. Sao Paulo. Imprensa Oficial do Estado de Sao Paulo, pp. 82 - 94.","Giaretta, A. A., Ribeiro, D. T. & Oliveira, L. E. (2007) A new species of Ischnocnema (Anura: Eleutherodactylinae) from open areas of the Cerrado Biome in southeastern Brazil. Zootaxa, 1666, 43 - 51."]}

Toledo, L. F., Haddad, C. F. B. (2005): Reproductive biology of Scinax fuscomarginatus (Anura, Hylidae) in south-eastern Brazil. Journal of Natural History 39 (32): 3029-3037, DOI: 10.1080/00222930500221403, URL: http://dx.doi.org/10.1080/00222930500221403